35
MARK F. O'BRIEN Vol. 16, No. 2 Summer 1983 THE GREAT LAKES ENTOMOLOGIST PUBLISHED BY THE MICHIGAN EN1"OMOLOGICAL SOC I Erry

 · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

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Page 1:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

MARK F OBRIEN

Vol 16 No 2 Summer 1983

THE

GREAT LAKES ENTOMOLOGIST

PUBLISHED BY

THE MICHIGAN EN1OMOLOGICAL

SOCIErry

THE GREAT LAKES ENTOMOLOGIST

Published by the Michigan Entomological Society

Volume 16 No2

ISSN 0090-0222

TABLE OF CONTENTS

Seasonal Flight Patterns of Hemiptera in a North Carolina Black Walnut Plantation 7 Miridae

J E McPherson B C Weber and T J Henry 35

Effects of Various Split Developmental Photophases and Constant Light During Each 24 Hour Period on Adult Morphology in Thyanta calceata (Hemiptera Pentatomidae)

J E McPherson T E Vogt and S M Paskewitz 43

Buprestidae Cerambycidae and Scolytidae Associated with Successive Stages of Agrilus bilineatus (Coleoptera Buprestidae) Infestation of Oaks in Wisconsin

R A Haack D M Benjamin and K D Haack 47

A Pyralid Moth (Lepidoptera) as Pollinator of Blunt-leaf Orchid Edward G Voss and Richard E Riefner Jr 57

Checklist of American Uloboridae (Arachnida Araneae) Brent D Ope II 61

COVER ILLUSTRATION

Blister beetles (Meloidae) feeding on Siberian pea-tree (Caragana arborescens) Photoshygraph by Louis F Wilson North Central Forest Experiment Station USDA Forest Serice East Lansing Michigan

THE MICHIGAN ENTOMOLOGICAL SOCIETY

1982-83 OFFICERS

President Ronald J Priest President-Elect Gary A Dunn Executive Secretary M C Nielsen Journal Editor D C L Gosling Newsletter Editor Louis F Wilson

The Michigan Entomological Society traces its origins to the old Detroit Entomological Society and was organized on 4 November 1954 to promote the science ofentomology in all its branches and by all feasible means and to advance cooperation and good fellowship among persons interested in entomology The Society attempts to facilitate the exchange of ideas and information in both amateur and professional circles and encourages the study of insects by youth Membership in the Society which serves the North Central States and adjacent Canada is open to all persons interested in entomology There are four paying classes of membership

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Inquiries about back numbers subscriptions and Society business should be directed to the Executive Secretary Michigan Entomological Society Department of Entomology Michigan State University East Lansing Michigan 48824 USA Manuscripts and related correspondence should be directed to the Editor (see inside back cover)

Copyright copy 1983 The Michigan Entomological Society

1983 THE GREAT LAKES ENTOMOLOGIST 35

SEASONAL FLIGHT PATTERNS OF HEMIPTERA IN A NORTH CAROLINA BLACK WALNUT PLANTATION 7 MIRIDAE

J E McPherson I B C Weber2 and T 1 Henry3

ABSTRACT

The seasonal flight patterns of 79 species of Miridae collected in window traps in a North Carolina black walnut plantation are described Flying height distributions and seasonal flight activities of Deraeocoris nebulosus (Uhler) Keltonia sulphurea (Reuter) Lygus lineoaris (Palisot de Beauvois) Plagiognathus poUtus Uhler and Reuteroscopus ornatus (Reuter) are considered in detail

This is the last in a series of papers on seasonal flight patterns of Hemiptera in a black walnut (Jugans nigra L) plantation near Asheville North Carolina and deals with the family Miridae earlier papers dealt with the Pentatomoidea (1980) Coreoidea (l98Ia) Reduvioidea (l981b) Cimicoidea (l98Ic) Lygaeoidea (l98Id) and Tingidae and Aradidae (198Ie) The study was conducted from 24 March to 14 October in 1977 and from 24 March to 13 October in 1978 Specimens were collected weekly by window trapping traps were suspended at 1 2 3 4 5 6 and 7 m The study site and trap construction are discussed in detail by McPherson and Weber (1980) All hemipteran specimens collected during this study are deposited in the Entomology Collection Zoology Research Museum Southern Illinois University Carbondale

RESULTS AND DISCUSSION

Seventy nine mirid species were collected during the two years of this study with the subfamily Mirinae being best represented numbers of specimens eollected for all taxa ranged from I to 2673 (Table I)

Most of the species were collected in numbers too low to permit conclusions about seasonal flight patterns However Deraeocoris nebulosus (Uhler) Keltollia suphurea (Reuter) Lygus lineolaris (Palisot de Beauvois) Pagiogllathus POlilUS Uhler and Reliteroshyscopus ornatus (Reuter) were collected in sufficient numbers (Table 1) to allow a more detailed discussion of flying height distributions and seasonal flight activities

D Ilebulosus a predaceous species is generally associated with trees and shrubs (Wheeler et al 1975) It has been reported to attack various species of mites and insects including among others European red mite Panollychus ulmi (Koch) woolly apple aphid Eriosoma lanigerum (Hausmann) clover aphid Nearctaphis bakeri (Cowen) hop aphid Phorodoll humuli (Schrank) cotton aphid Aphis gossypii Glover terrapin scale Lecanium nigrofasciatum Pergande eyespotted bud moth Spiionota ocellana (Denis and Schiffershymuller) and possibly codling moth Laspeyresia pomollella (L) (see literature survey of Wheeler et al 1975) Wheeler et al (1975) in their study of this insect also reported its feeding on the mite Oliogonychus bicoior (Banks) the oak lace bug Corythucha arcuata

I Department of Zoology Southern Illinois University Carbondale IL 6290 L 2USDA Forest Service North Central Forest EXperiment Station Forestry Sciences Laboratory

Carbondale IL 62901 3Systematic Entomology Laboratory lIBllI Agricultuml Research Service USDA co National Mushy

seum of Natural History Washington DC 20560

wTable I Seasonal flight activity of Miridae during 1977-78 in a North Carolina black walnut plantation 0

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

MIRINAE Adelphocoris lineolatus (Goeze) 2 200 27 May Adelphocoris rapidus (Say) 11 182 plusmn 033 1-4 26 May-6 Oct Agnocoris pulverulentus (Uhler) 2 450 plusmn 250 2-7 26 May-30 June Capsus ater (L) Dichrooscytus elegans Heidemann Dichrooscytus suspectus Reuter Garganus Jusiformis (Say)

4 1 1 3

425 plusmn 095 700 400 167 067

3-7

1-3

20 May-9 June 23 June 2 June

24 June

r tTl Clg

Leptopterna dolabrata (L) 5 320 plusmn 102 1-6 13 May-26 May ~ Lygocoris caryae (Knight) Lygocoris geneseensis (Knight) Lygocoris sp Lygus lineolaris (Palisot de Beauvois)

11 3 1

929

382 052 600 plusmn 100 400 260 plusmn 0002

2-7 4-7

1-7

13 May-7 July 13 May-17 June 27 May 31 March-13 Oct

r Jgt iA tTl en

Megaloceroea recticornis (Geoffroy) Phytocoris prob americanus Carvalho

1 2

100 350 150 2-5

16 June 7 July

tTl Z

Phytocoris confluens Reuter Phytocoris conspurcatus Knight Phytocoris ereclUs Van Duzee Phytocoris Jenestratus Reuter Phytocoris intermedius Henry Phytocoris minutulus Knight

I I 2 I 3 7

500 300 450 plusmn 050 700 533 plusmn 067 200 plusmn 053

4-5

4-6 1-5

30 June 7 Oct

23 Jooe-5 Aug 20 May 20 May-16 June 30 J une-6 Oct

as a r a Cl-en

Phytocoris penipectus Knight 1 600 13 May Phytocoris puella Reuter 2 550 050 5-6 23 June-30 June Phytocoris salicis Knight I 500 7 July Phytocoris tibialis Reuter Polymerus basalis (Reuter)

8 21

250 190

073 033

1-7 1-7

30 June-IS Sept 1 April-13 Oct

lt P

Prepops Jraterculus (Knight) 1 700 2 Sept ~ Stenodema trispinosa Reuter Stenotus binotatus (Fabricius)

4 6

275 plusmn 144 400 plusmn 093

1-7 1-7

1 April-6 May 2 June-30 June

Z

Taylorilygus pallidulus (Blanchard) 36 211 plusmn 021 1-6 30 June-7 Oct N

Trigonotylus coelestiaUum (Kirkaldy) Trigonotylus doddi (Distant) Tropidosteptes rufusculus (Knight)

ORTHOTYLINAE Ceratocapsus setosus Reuter Ceratocapsus sp Diaphnocoris provancheri (Burque) Halticus bractatus (Say) Halticus intermedius Uhler Heterocordylas maUnus Reuter [nacora stalii Reuter Lopidea heidemanni Knight Lopidea media (Say) Lopidea robiniae (Uhler) Orthotylus modestus Van IAlZee Orthotylus ornatas Van Duzee Orthotylus ramus Knight Pilophorus crassipes Heidemann Pseudoxenetus scutellatus (Uhler) Sericophanes heidemanni Poppiu8 Slaterocoris atritibialis (Knight) Slaterocoris stygicus (Say)

PHYLINAE Chlamydatus suavis (Reuter) Criocoris sa liens (Reuter) Icodema nigrolineatum (Knight) Keltonia sulphurea (Reuter) Lepidopsalus c1aricomis Knight Lepidopsallus miniatus Knight Lepidopsalus rostratus Knight Microphylelus modestus Reuter Plagiognathus albatus (Van Duzee) Plagiognathus cameolus Knight Plagiognathus dispar Knight

37 4 2

2 1 6

24 3 1

82 33 3 1 7 3 5 4 2

12 6 1

5 2 2

194 14 6

13 2 1 4 1

459 plusmn 033 425 plusmn 103 600 plusmn 100

250 050 600 283 060 275 plusmn 034 333 t 120 700 209 t 019 206 033 267 plusmn 088 700 514 059 500 plusmn 058 480 plusmn 080 600 plusmn 058 450 plusmn 250 300 plusmn 051 217 075 100

360 plusmn 125 200 350 plusmn 150 132 plusmn 0Q7 336 027 367 088 377 046 500 100 100 250 plusmn 096 500

1-7 2-7 5-7

2-3

1-5 1-6 1-5

1-7 1-7 1-4

2-7 4-6 3-7 5-7 2-7 1-6 1-5

1-7

2-5 1-7 2-5 1-7 2-7 4-6

1-5

29 April-13 Oct 0 0030 June-6 Oct vgt

22 April-28 April

29 Ju1y-5 Aug 15 July 17 June-8 Sept 5 May-29 Sept

24 June-15 July gt-l9 June r

27 May-7 Oct ttI Q19 May-18 Aug ~

23 June-24 June ttI 7 July gtshy

gt-l 20 May-9 June t 6 May-26 May gtshy

i27 May-7 July ttI (Il10 lune-6 Oct

13 May-27 May ttI Z5 May-13 Oct gt-l

29 April-16 June 0 a26 May 0 t 0 Q

12 May-6 Oct til 19 May-20 Mv gt-l

24 June-14 27 May-7 Oct 6 May-3 June

22 April-16 June 13 May-l July 27 May 15 July 6 May-12 May

vgt10 June -j

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

~ ~ 5 0

M A M J J A S 0

25

iii 20 gt

5 15 E

0 10c

(J 5 0

M

sect 20 15 5

~ 10 c (J 5

0

M

8

r~ 1 A

)1J A M J J A S 0

10 ~ Ii1

I(

I I I

A M J J A SO

25----------------- 7sect 20 0 15 E

0 ~ 10 c IIgt

l(J

5

M A M J J A S 0

25-----------------

9

M A M J J A S 0

42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 2:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

THE GREAT LAKES ENTOMOLOGIST

Published by the Michigan Entomological Society

Volume 16 No2

ISSN 0090-0222

TABLE OF CONTENTS

Seasonal Flight Patterns of Hemiptera in a North Carolina Black Walnut Plantation 7 Miridae

J E McPherson B C Weber and T J Henry 35

Effects of Various Split Developmental Photophases and Constant Light During Each 24 Hour Period on Adult Morphology in Thyanta calceata (Hemiptera Pentatomidae)

J E McPherson T E Vogt and S M Paskewitz 43

Buprestidae Cerambycidae and Scolytidae Associated with Successive Stages of Agrilus bilineatus (Coleoptera Buprestidae) Infestation of Oaks in Wisconsin

R A Haack D M Benjamin and K D Haack 47

A Pyralid Moth (Lepidoptera) as Pollinator of Blunt-leaf Orchid Edward G Voss and Richard E Riefner Jr 57

Checklist of American Uloboridae (Arachnida Araneae) Brent D Ope II 61

COVER ILLUSTRATION

Blister beetles (Meloidae) feeding on Siberian pea-tree (Caragana arborescens) Photoshygraph by Louis F Wilson North Central Forest Experiment Station USDA Forest Serice East Lansing Michigan

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Copyright copy 1983 The Michigan Entomological Society

1983 THE GREAT LAKES ENTOMOLOGIST 35

SEASONAL FLIGHT PATTERNS OF HEMIPTERA IN A NORTH CAROLINA BLACK WALNUT PLANTATION 7 MIRIDAE

J E McPherson I B C Weber2 and T 1 Henry3

ABSTRACT

The seasonal flight patterns of 79 species of Miridae collected in window traps in a North Carolina black walnut plantation are described Flying height distributions and seasonal flight activities of Deraeocoris nebulosus (Uhler) Keltonia sulphurea (Reuter) Lygus lineoaris (Palisot de Beauvois) Plagiognathus poUtus Uhler and Reuteroscopus ornatus (Reuter) are considered in detail

This is the last in a series of papers on seasonal flight patterns of Hemiptera in a black walnut (Jugans nigra L) plantation near Asheville North Carolina and deals with the family Miridae earlier papers dealt with the Pentatomoidea (1980) Coreoidea (l98Ia) Reduvioidea (l981b) Cimicoidea (l98Ic) Lygaeoidea (l98Id) and Tingidae and Aradidae (198Ie) The study was conducted from 24 March to 14 October in 1977 and from 24 March to 13 October in 1978 Specimens were collected weekly by window trapping traps were suspended at 1 2 3 4 5 6 and 7 m The study site and trap construction are discussed in detail by McPherson and Weber (1980) All hemipteran specimens collected during this study are deposited in the Entomology Collection Zoology Research Museum Southern Illinois University Carbondale

RESULTS AND DISCUSSION

Seventy nine mirid species were collected during the two years of this study with the subfamily Mirinae being best represented numbers of specimens eollected for all taxa ranged from I to 2673 (Table I)

Most of the species were collected in numbers too low to permit conclusions about seasonal flight patterns However Deraeocoris nebulosus (Uhler) Keltollia suphurea (Reuter) Lygus lineolaris (Palisot de Beauvois) Pagiogllathus POlilUS Uhler and Reliteroshyscopus ornatus (Reuter) were collected in sufficient numbers (Table 1) to allow a more detailed discussion of flying height distributions and seasonal flight activities

D Ilebulosus a predaceous species is generally associated with trees and shrubs (Wheeler et al 1975) It has been reported to attack various species of mites and insects including among others European red mite Panollychus ulmi (Koch) woolly apple aphid Eriosoma lanigerum (Hausmann) clover aphid Nearctaphis bakeri (Cowen) hop aphid Phorodoll humuli (Schrank) cotton aphid Aphis gossypii Glover terrapin scale Lecanium nigrofasciatum Pergande eyespotted bud moth Spiionota ocellana (Denis and Schiffershymuller) and possibly codling moth Laspeyresia pomollella (L) (see literature survey of Wheeler et al 1975) Wheeler et al (1975) in their study of this insect also reported its feeding on the mite Oliogonychus bicoior (Banks) the oak lace bug Corythucha arcuata

I Department of Zoology Southern Illinois University Carbondale IL 6290 L 2USDA Forest Service North Central Forest EXperiment Station Forestry Sciences Laboratory

Carbondale IL 62901 3Systematic Entomology Laboratory lIBllI Agricultuml Research Service USDA co National Mushy

seum of Natural History Washington DC 20560

wTable I Seasonal flight activity of Miridae during 1977-78 in a North Carolina black walnut plantation 0

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

MIRINAE Adelphocoris lineolatus (Goeze) 2 200 27 May Adelphocoris rapidus (Say) 11 182 plusmn 033 1-4 26 May-6 Oct Agnocoris pulverulentus (Uhler) 2 450 plusmn 250 2-7 26 May-30 June Capsus ater (L) Dichrooscytus elegans Heidemann Dichrooscytus suspectus Reuter Garganus Jusiformis (Say)

4 1 1 3

425 plusmn 095 700 400 167 067

3-7

1-3

20 May-9 June 23 June 2 June

24 June

r tTl Clg

Leptopterna dolabrata (L) 5 320 plusmn 102 1-6 13 May-26 May ~ Lygocoris caryae (Knight) Lygocoris geneseensis (Knight) Lygocoris sp Lygus lineolaris (Palisot de Beauvois)

11 3 1

929

382 052 600 plusmn 100 400 260 plusmn 0002

2-7 4-7

1-7

13 May-7 July 13 May-17 June 27 May 31 March-13 Oct

r Jgt iA tTl en

Megaloceroea recticornis (Geoffroy) Phytocoris prob americanus Carvalho

1 2

100 350 150 2-5

16 June 7 July

tTl Z

Phytocoris confluens Reuter Phytocoris conspurcatus Knight Phytocoris ereclUs Van Duzee Phytocoris Jenestratus Reuter Phytocoris intermedius Henry Phytocoris minutulus Knight

I I 2 I 3 7

500 300 450 plusmn 050 700 533 plusmn 067 200 plusmn 053

4-5

4-6 1-5

30 June 7 Oct

23 Jooe-5 Aug 20 May 20 May-16 June 30 J une-6 Oct

as a r a Cl-en

Phytocoris penipectus Knight 1 600 13 May Phytocoris puella Reuter 2 550 050 5-6 23 June-30 June Phytocoris salicis Knight I 500 7 July Phytocoris tibialis Reuter Polymerus basalis (Reuter)

8 21

250 190

073 033

1-7 1-7

30 June-IS Sept 1 April-13 Oct

lt P

Prepops Jraterculus (Knight) 1 700 2 Sept ~ Stenodema trispinosa Reuter Stenotus binotatus (Fabricius)

4 6

275 plusmn 144 400 plusmn 093

1-7 1-7

1 April-6 May 2 June-30 June

Z

Taylorilygus pallidulus (Blanchard) 36 211 plusmn 021 1-6 30 June-7 Oct N

Trigonotylus coelestiaUum (Kirkaldy) Trigonotylus doddi (Distant) Tropidosteptes rufusculus (Knight)

ORTHOTYLINAE Ceratocapsus setosus Reuter Ceratocapsus sp Diaphnocoris provancheri (Burque) Halticus bractatus (Say) Halticus intermedius Uhler Heterocordylas maUnus Reuter [nacora stalii Reuter Lopidea heidemanni Knight Lopidea media (Say) Lopidea robiniae (Uhler) Orthotylus modestus Van IAlZee Orthotylus ornatas Van Duzee Orthotylus ramus Knight Pilophorus crassipes Heidemann Pseudoxenetus scutellatus (Uhler) Sericophanes heidemanni Poppiu8 Slaterocoris atritibialis (Knight) Slaterocoris stygicus (Say)

PHYLINAE Chlamydatus suavis (Reuter) Criocoris sa liens (Reuter) Icodema nigrolineatum (Knight) Keltonia sulphurea (Reuter) Lepidopsalus c1aricomis Knight Lepidopsallus miniatus Knight Lepidopsalus rostratus Knight Microphylelus modestus Reuter Plagiognathus albatus (Van Duzee) Plagiognathus cameolus Knight Plagiognathus dispar Knight

37 4 2

2 1 6

24 3 1

82 33 3 1 7 3 5 4 2

12 6 1

5 2 2

194 14 6

13 2 1 4 1

459 plusmn 033 425 plusmn 103 600 plusmn 100

250 050 600 283 060 275 plusmn 034 333 t 120 700 209 t 019 206 033 267 plusmn 088 700 514 059 500 plusmn 058 480 plusmn 080 600 plusmn 058 450 plusmn 250 300 plusmn 051 217 075 100

360 plusmn 125 200 350 plusmn 150 132 plusmn 0Q7 336 027 367 088 377 046 500 100 100 250 plusmn 096 500

1-7 2-7 5-7

2-3

1-5 1-6 1-5

1-7 1-7 1-4

2-7 4-6 3-7 5-7 2-7 1-6 1-5

1-7

2-5 1-7 2-5 1-7 2-7 4-6

1-5

29 April-13 Oct 0 0030 June-6 Oct vgt

22 April-28 April

29 Ju1y-5 Aug 15 July 17 June-8 Sept 5 May-29 Sept

24 June-15 July gt-l9 June r

27 May-7 Oct ttI Q19 May-18 Aug ~

23 June-24 June ttI 7 July gtshy

gt-l 20 May-9 June t 6 May-26 May gtshy

i27 May-7 July ttI (Il10 lune-6 Oct

13 May-27 May ttI Z5 May-13 Oct gt-l

29 April-16 June 0 a26 May 0 t 0 Q

12 May-6 Oct til 19 May-20 Mv gt-l

24 June-14 27 May-7 Oct 6 May-3 June

22 April-16 June 13 May-l July 27 May 15 July 6 May-12 May

vgt10 June -j

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

~ ~ 5 0

M A M J J A S 0

25

iii 20 gt

5 15 E

0 10c

(J 5 0

M

sect 20 15 5

~ 10 c (J 5

0

M

8

r~ 1 A

)1J A M J J A S 0

10 ~ Ii1

I(

I I I

A M J J A SO

25----------------- 7sect 20 0 15 E

0 ~ 10 c IIgt

l(J

5

M A M J J A S 0

25-----------------

9

M A M J J A S 0

42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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1983 THE GREAT LAKES ENTOMOLOGIST 35

SEASONAL FLIGHT PATTERNS OF HEMIPTERA IN A NORTH CAROLINA BLACK WALNUT PLANTATION 7 MIRIDAE

J E McPherson I B C Weber2 and T 1 Henry3

ABSTRACT

The seasonal flight patterns of 79 species of Miridae collected in window traps in a North Carolina black walnut plantation are described Flying height distributions and seasonal flight activities of Deraeocoris nebulosus (Uhler) Keltonia sulphurea (Reuter) Lygus lineoaris (Palisot de Beauvois) Plagiognathus poUtus Uhler and Reuteroscopus ornatus (Reuter) are considered in detail

This is the last in a series of papers on seasonal flight patterns of Hemiptera in a black walnut (Jugans nigra L) plantation near Asheville North Carolina and deals with the family Miridae earlier papers dealt with the Pentatomoidea (1980) Coreoidea (l98Ia) Reduvioidea (l981b) Cimicoidea (l98Ic) Lygaeoidea (l98Id) and Tingidae and Aradidae (198Ie) The study was conducted from 24 March to 14 October in 1977 and from 24 March to 13 October in 1978 Specimens were collected weekly by window trapping traps were suspended at 1 2 3 4 5 6 and 7 m The study site and trap construction are discussed in detail by McPherson and Weber (1980) All hemipteran specimens collected during this study are deposited in the Entomology Collection Zoology Research Museum Southern Illinois University Carbondale

RESULTS AND DISCUSSION

Seventy nine mirid species were collected during the two years of this study with the subfamily Mirinae being best represented numbers of specimens eollected for all taxa ranged from I to 2673 (Table I)

Most of the species were collected in numbers too low to permit conclusions about seasonal flight patterns However Deraeocoris nebulosus (Uhler) Keltollia suphurea (Reuter) Lygus lineolaris (Palisot de Beauvois) Pagiogllathus POlilUS Uhler and Reliteroshyscopus ornatus (Reuter) were collected in sufficient numbers (Table 1) to allow a more detailed discussion of flying height distributions and seasonal flight activities

D Ilebulosus a predaceous species is generally associated with trees and shrubs (Wheeler et al 1975) It has been reported to attack various species of mites and insects including among others European red mite Panollychus ulmi (Koch) woolly apple aphid Eriosoma lanigerum (Hausmann) clover aphid Nearctaphis bakeri (Cowen) hop aphid Phorodoll humuli (Schrank) cotton aphid Aphis gossypii Glover terrapin scale Lecanium nigrofasciatum Pergande eyespotted bud moth Spiionota ocellana (Denis and Schiffershymuller) and possibly codling moth Laspeyresia pomollella (L) (see literature survey of Wheeler et al 1975) Wheeler et al (1975) in their study of this insect also reported its feeding on the mite Oliogonychus bicoior (Banks) the oak lace bug Corythucha arcuata

I Department of Zoology Southern Illinois University Carbondale IL 6290 L 2USDA Forest Service North Central Forest EXperiment Station Forestry Sciences Laboratory

Carbondale IL 62901 3Systematic Entomology Laboratory lIBllI Agricultuml Research Service USDA co National Mushy

seum of Natural History Washington DC 20560

wTable I Seasonal flight activity of Miridae during 1977-78 in a North Carolina black walnut plantation 0

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

MIRINAE Adelphocoris lineolatus (Goeze) 2 200 27 May Adelphocoris rapidus (Say) 11 182 plusmn 033 1-4 26 May-6 Oct Agnocoris pulverulentus (Uhler) 2 450 plusmn 250 2-7 26 May-30 June Capsus ater (L) Dichrooscytus elegans Heidemann Dichrooscytus suspectus Reuter Garganus Jusiformis (Say)

4 1 1 3

425 plusmn 095 700 400 167 067

3-7

1-3

20 May-9 June 23 June 2 June

24 June

r tTl Clg

Leptopterna dolabrata (L) 5 320 plusmn 102 1-6 13 May-26 May ~ Lygocoris caryae (Knight) Lygocoris geneseensis (Knight) Lygocoris sp Lygus lineolaris (Palisot de Beauvois)

11 3 1

929

382 052 600 plusmn 100 400 260 plusmn 0002

2-7 4-7

1-7

13 May-7 July 13 May-17 June 27 May 31 March-13 Oct

r Jgt iA tTl en

Megaloceroea recticornis (Geoffroy) Phytocoris prob americanus Carvalho

1 2

100 350 150 2-5

16 June 7 July

tTl Z

Phytocoris confluens Reuter Phytocoris conspurcatus Knight Phytocoris ereclUs Van Duzee Phytocoris Jenestratus Reuter Phytocoris intermedius Henry Phytocoris minutulus Knight

I I 2 I 3 7

500 300 450 plusmn 050 700 533 plusmn 067 200 plusmn 053

4-5

4-6 1-5

30 June 7 Oct

23 Jooe-5 Aug 20 May 20 May-16 June 30 J une-6 Oct

as a r a Cl-en

Phytocoris penipectus Knight 1 600 13 May Phytocoris puella Reuter 2 550 050 5-6 23 June-30 June Phytocoris salicis Knight I 500 7 July Phytocoris tibialis Reuter Polymerus basalis (Reuter)

8 21

250 190

073 033

1-7 1-7

30 June-IS Sept 1 April-13 Oct

lt P

Prepops Jraterculus (Knight) 1 700 2 Sept ~ Stenodema trispinosa Reuter Stenotus binotatus (Fabricius)

4 6

275 plusmn 144 400 plusmn 093

1-7 1-7

1 April-6 May 2 June-30 June

Z

Taylorilygus pallidulus (Blanchard) 36 211 plusmn 021 1-6 30 June-7 Oct N

Trigonotylus coelestiaUum (Kirkaldy) Trigonotylus doddi (Distant) Tropidosteptes rufusculus (Knight)

ORTHOTYLINAE Ceratocapsus setosus Reuter Ceratocapsus sp Diaphnocoris provancheri (Burque) Halticus bractatus (Say) Halticus intermedius Uhler Heterocordylas maUnus Reuter [nacora stalii Reuter Lopidea heidemanni Knight Lopidea media (Say) Lopidea robiniae (Uhler) Orthotylus modestus Van IAlZee Orthotylus ornatas Van Duzee Orthotylus ramus Knight Pilophorus crassipes Heidemann Pseudoxenetus scutellatus (Uhler) Sericophanes heidemanni Poppiu8 Slaterocoris atritibialis (Knight) Slaterocoris stygicus (Say)

PHYLINAE Chlamydatus suavis (Reuter) Criocoris sa liens (Reuter) Icodema nigrolineatum (Knight) Keltonia sulphurea (Reuter) Lepidopsalus c1aricomis Knight Lepidopsallus miniatus Knight Lepidopsalus rostratus Knight Microphylelus modestus Reuter Plagiognathus albatus (Van Duzee) Plagiognathus cameolus Knight Plagiognathus dispar Knight

37 4 2

2 1 6

24 3 1

82 33 3 1 7 3 5 4 2

12 6 1

5 2 2

194 14 6

13 2 1 4 1

459 plusmn 033 425 plusmn 103 600 plusmn 100

250 050 600 283 060 275 plusmn 034 333 t 120 700 209 t 019 206 033 267 plusmn 088 700 514 059 500 plusmn 058 480 plusmn 080 600 plusmn 058 450 plusmn 250 300 plusmn 051 217 075 100

360 plusmn 125 200 350 plusmn 150 132 plusmn 0Q7 336 027 367 088 377 046 500 100 100 250 plusmn 096 500

1-7 2-7 5-7

2-3

1-5 1-6 1-5

1-7 1-7 1-4

2-7 4-6 3-7 5-7 2-7 1-6 1-5

1-7

2-5 1-7 2-5 1-7 2-7 4-6

1-5

29 April-13 Oct 0 0030 June-6 Oct vgt

22 April-28 April

29 Ju1y-5 Aug 15 July 17 June-8 Sept 5 May-29 Sept

24 June-15 July gt-l9 June r

27 May-7 Oct ttI Q19 May-18 Aug ~

23 June-24 June ttI 7 July gtshy

gt-l 20 May-9 June t 6 May-26 May gtshy

i27 May-7 July ttI (Il10 lune-6 Oct

13 May-27 May ttI Z5 May-13 Oct gt-l

29 April-16 June 0 a26 May 0 t 0 Q

12 May-6 Oct til 19 May-20 Mv gt-l

24 June-14 27 May-7 Oct 6 May-3 June

22 April-16 June 13 May-l July 27 May 15 July 6 May-12 May

vgt10 June -j

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

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M A M J J A S 0

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0 10c

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~ 10 c (J 5

0

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8

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10 ~ Ii1

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A M J J A SO

25----------------- 7sect 20 0 15 E

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M A M J J A S 0

25-----------------

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42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

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47 I 8113 b

i

gt-lr til Cl l til

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v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 4:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 35

SEASONAL FLIGHT PATTERNS OF HEMIPTERA IN A NORTH CAROLINA BLACK WALNUT PLANTATION 7 MIRIDAE

J E McPherson I B C Weber2 and T 1 Henry3

ABSTRACT

The seasonal flight patterns of 79 species of Miridae collected in window traps in a North Carolina black walnut plantation are described Flying height distributions and seasonal flight activities of Deraeocoris nebulosus (Uhler) Keltonia sulphurea (Reuter) Lygus lineoaris (Palisot de Beauvois) Plagiognathus poUtus Uhler and Reuteroscopus ornatus (Reuter) are considered in detail

This is the last in a series of papers on seasonal flight patterns of Hemiptera in a black walnut (Jugans nigra L) plantation near Asheville North Carolina and deals with the family Miridae earlier papers dealt with the Pentatomoidea (1980) Coreoidea (l98Ia) Reduvioidea (l981b) Cimicoidea (l98Ic) Lygaeoidea (l98Id) and Tingidae and Aradidae (198Ie) The study was conducted from 24 March to 14 October in 1977 and from 24 March to 13 October in 1978 Specimens were collected weekly by window trapping traps were suspended at 1 2 3 4 5 6 and 7 m The study site and trap construction are discussed in detail by McPherson and Weber (1980) All hemipteran specimens collected during this study are deposited in the Entomology Collection Zoology Research Museum Southern Illinois University Carbondale

RESULTS AND DISCUSSION

Seventy nine mirid species were collected during the two years of this study with the subfamily Mirinae being best represented numbers of specimens eollected for all taxa ranged from I to 2673 (Table I)

Most of the species were collected in numbers too low to permit conclusions about seasonal flight patterns However Deraeocoris nebulosus (Uhler) Keltollia suphurea (Reuter) Lygus lineolaris (Palisot de Beauvois) Pagiogllathus POlilUS Uhler and Reliteroshyscopus ornatus (Reuter) were collected in sufficient numbers (Table 1) to allow a more detailed discussion of flying height distributions and seasonal flight activities

D Ilebulosus a predaceous species is generally associated with trees and shrubs (Wheeler et al 1975) It has been reported to attack various species of mites and insects including among others European red mite Panollychus ulmi (Koch) woolly apple aphid Eriosoma lanigerum (Hausmann) clover aphid Nearctaphis bakeri (Cowen) hop aphid Phorodoll humuli (Schrank) cotton aphid Aphis gossypii Glover terrapin scale Lecanium nigrofasciatum Pergande eyespotted bud moth Spiionota ocellana (Denis and Schiffershymuller) and possibly codling moth Laspeyresia pomollella (L) (see literature survey of Wheeler et al 1975) Wheeler et al (1975) in their study of this insect also reported its feeding on the mite Oliogonychus bicoior (Banks) the oak lace bug Corythucha arcuata

I Department of Zoology Southern Illinois University Carbondale IL 6290 L 2USDA Forest Service North Central Forest EXperiment Station Forestry Sciences Laboratory

Carbondale IL 62901 3Systematic Entomology Laboratory lIBllI Agricultuml Research Service USDA co National Mushy

seum of Natural History Washington DC 20560

wTable I Seasonal flight activity of Miridae during 1977-78 in a North Carolina black walnut plantation 0

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

MIRINAE Adelphocoris lineolatus (Goeze) 2 200 27 May Adelphocoris rapidus (Say) 11 182 plusmn 033 1-4 26 May-6 Oct Agnocoris pulverulentus (Uhler) 2 450 plusmn 250 2-7 26 May-30 June Capsus ater (L) Dichrooscytus elegans Heidemann Dichrooscytus suspectus Reuter Garganus Jusiformis (Say)

4 1 1 3

425 plusmn 095 700 400 167 067

3-7

1-3

20 May-9 June 23 June 2 June

24 June

r tTl Clg

Leptopterna dolabrata (L) 5 320 plusmn 102 1-6 13 May-26 May ~ Lygocoris caryae (Knight) Lygocoris geneseensis (Knight) Lygocoris sp Lygus lineolaris (Palisot de Beauvois)

11 3 1

929

382 052 600 plusmn 100 400 260 plusmn 0002

2-7 4-7

1-7

13 May-7 July 13 May-17 June 27 May 31 March-13 Oct

r Jgt iA tTl en

Megaloceroea recticornis (Geoffroy) Phytocoris prob americanus Carvalho

1 2

100 350 150 2-5

16 June 7 July

tTl Z

Phytocoris confluens Reuter Phytocoris conspurcatus Knight Phytocoris ereclUs Van Duzee Phytocoris Jenestratus Reuter Phytocoris intermedius Henry Phytocoris minutulus Knight

I I 2 I 3 7

500 300 450 plusmn 050 700 533 plusmn 067 200 plusmn 053

4-5

4-6 1-5

30 June 7 Oct

23 Jooe-5 Aug 20 May 20 May-16 June 30 J une-6 Oct

as a r a Cl-en

Phytocoris penipectus Knight 1 600 13 May Phytocoris puella Reuter 2 550 050 5-6 23 June-30 June Phytocoris salicis Knight I 500 7 July Phytocoris tibialis Reuter Polymerus basalis (Reuter)

8 21

250 190

073 033

1-7 1-7

30 June-IS Sept 1 April-13 Oct

lt P

Prepops Jraterculus (Knight) 1 700 2 Sept ~ Stenodema trispinosa Reuter Stenotus binotatus (Fabricius)

4 6

275 plusmn 144 400 plusmn 093

1-7 1-7

1 April-6 May 2 June-30 June

Z

Taylorilygus pallidulus (Blanchard) 36 211 plusmn 021 1-6 30 June-7 Oct N

Trigonotylus coelestiaUum (Kirkaldy) Trigonotylus doddi (Distant) Tropidosteptes rufusculus (Knight)

ORTHOTYLINAE Ceratocapsus setosus Reuter Ceratocapsus sp Diaphnocoris provancheri (Burque) Halticus bractatus (Say) Halticus intermedius Uhler Heterocordylas maUnus Reuter [nacora stalii Reuter Lopidea heidemanni Knight Lopidea media (Say) Lopidea robiniae (Uhler) Orthotylus modestus Van IAlZee Orthotylus ornatas Van Duzee Orthotylus ramus Knight Pilophorus crassipes Heidemann Pseudoxenetus scutellatus (Uhler) Sericophanes heidemanni Poppiu8 Slaterocoris atritibialis (Knight) Slaterocoris stygicus (Say)

PHYLINAE Chlamydatus suavis (Reuter) Criocoris sa liens (Reuter) Icodema nigrolineatum (Knight) Keltonia sulphurea (Reuter) Lepidopsalus c1aricomis Knight Lepidopsallus miniatus Knight Lepidopsalus rostratus Knight Microphylelus modestus Reuter Plagiognathus albatus (Van Duzee) Plagiognathus cameolus Knight Plagiognathus dispar Knight

37 4 2

2 1 6

24 3 1

82 33 3 1 7 3 5 4 2

12 6 1

5 2 2

194 14 6

13 2 1 4 1

459 plusmn 033 425 plusmn 103 600 plusmn 100

250 050 600 283 060 275 plusmn 034 333 t 120 700 209 t 019 206 033 267 plusmn 088 700 514 059 500 plusmn 058 480 plusmn 080 600 plusmn 058 450 plusmn 250 300 plusmn 051 217 075 100

360 plusmn 125 200 350 plusmn 150 132 plusmn 0Q7 336 027 367 088 377 046 500 100 100 250 plusmn 096 500

1-7 2-7 5-7

2-3

1-5 1-6 1-5

1-7 1-7 1-4

2-7 4-6 3-7 5-7 2-7 1-6 1-5

1-7

2-5 1-7 2-5 1-7 2-7 4-6

1-5

29 April-13 Oct 0 0030 June-6 Oct vgt

22 April-28 April

29 Ju1y-5 Aug 15 July 17 June-8 Sept 5 May-29 Sept

24 June-15 July gt-l9 June r

27 May-7 Oct ttI Q19 May-18 Aug ~

23 June-24 June ttI 7 July gtshy

gt-l 20 May-9 June t 6 May-26 May gtshy

i27 May-7 July ttI (Il10 lune-6 Oct

13 May-27 May ttI Z5 May-13 Oct gt-l

29 April-16 June 0 a26 May 0 t 0 Q

12 May-6 Oct til 19 May-20 Mv gt-l

24 June-14 27 May-7 Oct 6 May-3 June

22 April-16 June 13 May-l July 27 May 15 July 6 May-12 May

vgt10 June -j

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

~ ~ 5 0

M A M J J A S 0

25

iii 20 gt

5 15 E

0 10c

(J 5 0

M

sect 20 15 5

~ 10 c (J 5

0

M

8

r~ 1 A

)1J A M J J A S 0

10 ~ Ii1

I(

I I I

A M J J A SO

25----------------- 7sect 20 0 15 E

0 ~ 10 c IIgt

l(J

5

M A M J J A S 0

25-----------------

9

M A M J J A S 0

42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 5:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

wTable I Seasonal flight activity of Miridae during 1977-78 in a North Carolina black walnut plantation 0

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

MIRINAE Adelphocoris lineolatus (Goeze) 2 200 27 May Adelphocoris rapidus (Say) 11 182 plusmn 033 1-4 26 May-6 Oct Agnocoris pulverulentus (Uhler) 2 450 plusmn 250 2-7 26 May-30 June Capsus ater (L) Dichrooscytus elegans Heidemann Dichrooscytus suspectus Reuter Garganus Jusiformis (Say)

4 1 1 3

425 plusmn 095 700 400 167 067

3-7

1-3

20 May-9 June 23 June 2 June

24 June

r tTl Clg

Leptopterna dolabrata (L) 5 320 plusmn 102 1-6 13 May-26 May ~ Lygocoris caryae (Knight) Lygocoris geneseensis (Knight) Lygocoris sp Lygus lineolaris (Palisot de Beauvois)

11 3 1

929

382 052 600 plusmn 100 400 260 plusmn 0002

2-7 4-7

1-7

13 May-7 July 13 May-17 June 27 May 31 March-13 Oct

r Jgt iA tTl en

Megaloceroea recticornis (Geoffroy) Phytocoris prob americanus Carvalho

1 2

100 350 150 2-5

16 June 7 July

tTl Z

Phytocoris confluens Reuter Phytocoris conspurcatus Knight Phytocoris ereclUs Van Duzee Phytocoris Jenestratus Reuter Phytocoris intermedius Henry Phytocoris minutulus Knight

I I 2 I 3 7

500 300 450 plusmn 050 700 533 plusmn 067 200 plusmn 053

4-5

4-6 1-5

30 June 7 Oct

23 Jooe-5 Aug 20 May 20 May-16 June 30 J une-6 Oct

as a r a Cl-en

Phytocoris penipectus Knight 1 600 13 May Phytocoris puella Reuter 2 550 050 5-6 23 June-30 June Phytocoris salicis Knight I 500 7 July Phytocoris tibialis Reuter Polymerus basalis (Reuter)

8 21

250 190

073 033

1-7 1-7

30 June-IS Sept 1 April-13 Oct

lt P

Prepops Jraterculus (Knight) 1 700 2 Sept ~ Stenodema trispinosa Reuter Stenotus binotatus (Fabricius)

4 6

275 plusmn 144 400 plusmn 093

1-7 1-7

1 April-6 May 2 June-30 June

Z

Taylorilygus pallidulus (Blanchard) 36 211 plusmn 021 1-6 30 June-7 Oct N

Trigonotylus coelestiaUum (Kirkaldy) Trigonotylus doddi (Distant) Tropidosteptes rufusculus (Knight)

ORTHOTYLINAE Ceratocapsus setosus Reuter Ceratocapsus sp Diaphnocoris provancheri (Burque) Halticus bractatus (Say) Halticus intermedius Uhler Heterocordylas maUnus Reuter [nacora stalii Reuter Lopidea heidemanni Knight Lopidea media (Say) Lopidea robiniae (Uhler) Orthotylus modestus Van IAlZee Orthotylus ornatas Van Duzee Orthotylus ramus Knight Pilophorus crassipes Heidemann Pseudoxenetus scutellatus (Uhler) Sericophanes heidemanni Poppiu8 Slaterocoris atritibialis (Knight) Slaterocoris stygicus (Say)

PHYLINAE Chlamydatus suavis (Reuter) Criocoris sa liens (Reuter) Icodema nigrolineatum (Knight) Keltonia sulphurea (Reuter) Lepidopsalus c1aricomis Knight Lepidopsallus miniatus Knight Lepidopsalus rostratus Knight Microphylelus modestus Reuter Plagiognathus albatus (Van Duzee) Plagiognathus cameolus Knight Plagiognathus dispar Knight

37 4 2

2 1 6

24 3 1

82 33 3 1 7 3 5 4 2

12 6 1

5 2 2

194 14 6

13 2 1 4 1

459 plusmn 033 425 plusmn 103 600 plusmn 100

250 050 600 283 060 275 plusmn 034 333 t 120 700 209 t 019 206 033 267 plusmn 088 700 514 059 500 plusmn 058 480 plusmn 080 600 plusmn 058 450 plusmn 250 300 plusmn 051 217 075 100

360 plusmn 125 200 350 plusmn 150 132 plusmn 0Q7 336 027 367 088 377 046 500 100 100 250 plusmn 096 500

1-7 2-7 5-7

2-3

1-5 1-6 1-5

1-7 1-7 1-4

2-7 4-6 3-7 5-7 2-7 1-6 1-5

1-7

2-5 1-7 2-5 1-7 2-7 4-6

1-5

29 April-13 Oct 0 0030 June-6 Oct vgt

22 April-28 April

29 Ju1y-5 Aug 15 July 17 June-8 Sept 5 May-29 Sept

24 June-15 July gt-l9 June r

27 May-7 Oct ttI Q19 May-18 Aug ~

23 June-24 June ttI 7 July gtshy

gt-l 20 May-9 June t 6 May-26 May gtshy

i27 May-7 July ttI (Il10 lune-6 Oct

13 May-27 May ttI Z5 May-13 Oct gt-l

29 April-16 June 0 a26 May 0 t 0 Q

12 May-6 Oct til 19 May-20 Mv gt-l

24 June-14 27 May-7 Oct 6 May-3 June

22 April-16 June 13 May-l July 27 May 15 July 6 May-12 May

vgt10 June -j

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

~ ~ 5 0

M A M J J A S 0

25

iii 20 gt

5 15 E

0 10c

(J 5 0

M

sect 20 15 5

~ 10 c (J 5

0

M

8

r~ 1 A

)1J A M J J A S 0

10 ~ Ii1

I(

I I I

A M J J A SO

25----------------- 7sect 20 0 15 E

0 ~ 10 c IIgt

l(J

5

M A M J J A S 0

25-----------------

9

M A M J J A S 0

42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 6:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

Trigonotylus coelestiaUum (Kirkaldy) Trigonotylus doddi (Distant) Tropidosteptes rufusculus (Knight)

ORTHOTYLINAE Ceratocapsus setosus Reuter Ceratocapsus sp Diaphnocoris provancheri (Burque) Halticus bractatus (Say) Halticus intermedius Uhler Heterocordylas maUnus Reuter [nacora stalii Reuter Lopidea heidemanni Knight Lopidea media (Say) Lopidea robiniae (Uhler) Orthotylus modestus Van IAlZee Orthotylus ornatas Van Duzee Orthotylus ramus Knight Pilophorus crassipes Heidemann Pseudoxenetus scutellatus (Uhler) Sericophanes heidemanni Poppiu8 Slaterocoris atritibialis (Knight) Slaterocoris stygicus (Say)

PHYLINAE Chlamydatus suavis (Reuter) Criocoris sa liens (Reuter) Icodema nigrolineatum (Knight) Keltonia sulphurea (Reuter) Lepidopsalus c1aricomis Knight Lepidopsallus miniatus Knight Lepidopsalus rostratus Knight Microphylelus modestus Reuter Plagiognathus albatus (Van Duzee) Plagiognathus cameolus Knight Plagiognathus dispar Knight

37 4 2

2 1 6

24 3 1

82 33 3 1 7 3 5 4 2

12 6 1

5 2 2

194 14 6

13 2 1 4 1

459 plusmn 033 425 plusmn 103 600 plusmn 100

250 050 600 283 060 275 plusmn 034 333 t 120 700 209 t 019 206 033 267 plusmn 088 700 514 059 500 plusmn 058 480 plusmn 080 600 plusmn 058 450 plusmn 250 300 plusmn 051 217 075 100

360 plusmn 125 200 350 plusmn 150 132 plusmn 0Q7 336 027 367 088 377 046 500 100 100 250 plusmn 096 500

1-7 2-7 5-7

2-3

1-5 1-6 1-5

1-7 1-7 1-4

2-7 4-6 3-7 5-7 2-7 1-6 1-5

1-7

2-5 1-7 2-5 1-7 2-7 4-6

1-5

29 April-13 Oct 0 0030 June-6 Oct vgt

22 April-28 April

29 Ju1y-5 Aug 15 July 17 June-8 Sept 5 May-29 Sept

24 June-15 July gt-l9 June r

27 May-7 Oct ttI Q19 May-18 Aug ~

23 June-24 June ttI 7 July gtshy

gt-l 20 May-9 June t 6 May-26 May gtshy

i27 May-7 July ttI (Il10 lune-6 Oct

13 May-27 May ttI Z5 May-13 Oct gt-l

29 April-16 June 0 a26 May 0 t 0 Q

12 May-6 Oct til 19 May-20 Mv gt-l

24 June-14 27 May-7 Oct 6 May-3 June

22 April-16 June 13 May-l July 27 May 15 July 6 May-12 May

vgt10 June -j

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

~ ~ 5 0

M A M J J A S 0

25

iii 20 gt

5 15 E

0 10c

(J 5 0

M

sect 20 15 5

~ 10 c (J 5

0

M

8

r~ 1 A

)1J A M J J A S 0

10 ~ Ii1

I(

I I I

A M J J A SO

25----------------- 7sect 20 0 15 E

0 ~ 10 c IIgt

l(J

5

M A M J J A S 0

25-----------------

9

M A M J J A S 0

42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 7:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

00 cgtTable 1 Continued

Collection Height (m) No Range of

Taxon Collected x plusmn SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 100 13 May Plagiognathus nigronitens Knight I 100 23 June Plagiognathus politus Uhler 506 251 plusmn 0004 1-7 5 May-13 Oct Psallus strobicola Knight 5 540 plusmn 081 3-7 27 May-23 June Pseudatomoscelis seriatus (Reuter) Reuteroscopus ornatus (Reuter) Rhinocapsus rubricans (Provancher) Spanagonicus albofasciatus (Reuter)

23 2673

2 30

191 plusmn 033 458 plusmn 0001 450 plusmn 250 423 plusmn 039

1-7 1-7 2-7 1-7

2 June-29 27 May-13 Oct 16 June-28 July 29 July-13 Oct

r m Cl

mp DERAEOCORINAE

Deraeocoris nebulosus (Uhler) 612 286 plusmn 0002 1-7 I rp

Deraeocoris nigrituius Knight Eustictus grossus (Uhler) Hyaliodes harti Knight Hyaliodes vitripennis (Say)

3 2 7 5

533 plusmn 500 243 plusmn 160 plusmn

088

069 060

4-7

1-5 1-4

13 June 22 Sept 30 June-l Sept 24 June-29 Sept

~ m ()

m Z 0

DICYPHINAE Dicyphus famelicus (Uhler) Dicyphus prob rhododendri Dolling

100 700

15 April 3 June

~ 0 r 0 Cl

BRYOCORINAE til

Sixeonotus tenebrosus (Distant) 3 333 plusmn 088 2-5 3 June-7 Aug

CYLAPINAE Fulvius imbecilis (Say) Fulvius slateri Wheeler

2 8

400 plusmn 450 plusmn

300 033

1-7 3-6

29 Julys Sept 24 June-25 Aug

-lt 0 a L IV

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

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42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

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O lOb 45 44

490gb 44 S

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050a I 0

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5 6 OOOb

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6 42 490S b

3 36

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49 50 050a

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46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 8:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ETOMOLOGIST 39

(Say) hawthorn lace bug C cydon[ae (Fitch) and greenhouse whitefly Triaeurodes aporariorum (Westwood) In the Harrisburg area of Pennsylvania it overwinters as adults and is trivoltine (Wheeler et al 1975)

In the present study D nebuoslis adults were found from early April to late September (Table I) They were collected at all seven flying heights with about 60 captured at 2-3 m (Fig I)

This species overwintered as adults but the number of generations per year is unclear from the data available (Fig 6) If as Wheeler et al (1975) reported for Harrisburg this species is also trivoltine near Asheville then the overwintered adults emerged in early April and reproduced during the spring Flight activity of their offspring began in early June generally increased from mid-June through July peaked in eady August and declined sharply during the second half of August the length of this flight activity period (ie during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals

K sillphurea has been collected on Ambrosia sp Chenopodium abum L Symphorshyicarpos orbicuatlls Moench (Knight 1941) and Sida spinosa L (Knight 1927 1941 1966) No information has been published on its life cycle other than its reported breeding on S spinosa in early September in Georgia (Knight 1927)

In the present study K suphurea adults were found from late May to early October Cfable I) They were collected at all seven flying heights with about 85 collected at I m (Fig 2)

This species probably overwintered as eggs this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (ie late May) (Fig 7) Also if by breeding in early September Knight (1927) meant he observed copulating pairs at that time then fertilized femlaes would have had sufficient time to oviposit before the end of the season If this species does overwinter as eggs then the flight data suggest three generations adults of the first generation were present from late May to about mid-July those of the second from mid-July to late August and those of the third from early September to the end of the season

L lineaaris (tarnished plant bug) feeds on numerous plants including alfalfa apple apricot aster bean beet blackbeny cabbage carnation carrot celery cherry chrysanshythemum clover cotton cucumber currant dahlia grape lettuce marigold pea peach pear peony plum potato quince raspberry rose strawberry tobacco and turnip (Kelton 1975) However it also feeds on certain arthropods including alfalfa plant bug Adephocoris lin eoallis (Goeze) potato leafhopper Empoasca fabae (Harris) pea aphid Acynhasiphon pisum (Harris) Colorado potato beetle Leptinotarsa decemJineata (Say) alfalfa weevil Hypera postica (Gyllenhal) species of Noctuidae Peuraprllcha insusaria (Guem6e) alfalfa blotch leafminer Agramyza frantella (Rondani) Aphidius ervi pulcher Baker A smithi Sharma and Subba Rao Praan sp species of Formicidae and the harvestman Phaangillm opilio L (Wheeler 1976)

This species overwinters as adults (Guppy 1958 Kelton 1975 Ridgway and Gyrisco 1960 Stewart and Khoury 1976) and has two (Guppy 1958 Kelton 1975) or perhaps three (Ridgshyway and Gyrisco 1960 Stewart and Khoury 1976) generations per year

Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet They found that L lineoaris flew fairly close to the ground of 323 adults collected 300 were captured within 6 ft of the ground and only one as high as 15-18 feet

In the present study L lineoaris adults were found from late March to mid-October (Table I) They were collected at all seven flying heights with about 60 captured at 1-2 m and about 3 at 7 m (Fig 3) this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960)

This species overwintered as adults and is apparently bivoltine near Asheville (Fig 8) Adults began to emerge from overwintering sites in late March Their adult offspring (summer generation) occurred from about late May to mid-August Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

) I

~ ~ 5 0

M A M J J A S 0

25

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0 10c

(J 5 0

M

sect 20 15 5

~ 10 c (J 5

0

M

8

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)1J A M J J A S 0

10 ~ Ii1

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A M J J A SO

25----------------- 7sect 20 0 15 E

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25-----------------

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42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

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v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 9:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

40 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

100-------------- 100------------------r 1 2 ~ ~ 80~ 80 0

6 60 ~ 605 0 40 ~ 40 cC

()

0 20~ 20 Qn

2 3 4 561 2 3 4 561 Meters Meters

100~------------------- 100-------------------~

3 4 ~ 80~ 80 0II gt ~ 60

0

~ 60

o 4040 CC

()

a 20i 20 Qn

2 3 4 587 Meters

234 5 e 7 Meters

100

5 60 I

6 60 5

0 40 C ~ 20 n

2 3 456 7 Meters

Figs 1-5 Flying height distributions of five mirid species during 1977-78 in a North Carolina black walnut plantation (I) Deraeocoris nebulosus (2) Keltonia sulphurea (3) Lygus lineoluris (4) Plagiognathus politus (5) Reuteroscopus ornatus

p poUtus has been collected from several plants including Pyrus (Leonard 1915 Knight 1941) Ambrosia sp Betula sp Carya sp Corylus sp Juniperus virginiana L Pinus strobus L Quercus sp Robinia pseudoacacia L Salix sp Solidago sp Symphoricarpos orbiculatus Moench Taxodium distichum (L) (Knight 1941) Erigeron and Verbascum (Froeschner 1949) It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine) (Froeschner 1949 Knight 1941) In Missouri adults of the first generashytion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949)

In the present study P paUlUS adults were found from early May to mid-October (Table I) They were collected at all seven flying heights with almost 50 captured at 1 m (Fig 4)

This species apparently overwintered as eggs thus agreeing with the findings of Leonard (1915) Supporting this conclusion were the large number of flying adults in the fall and the

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

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M A M J J A S 0

25

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0

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10 ~ Ii1

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42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

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46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 10:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 41

Figs 6-10 Seasonal flight activities of five mirid species during 1977-78 in a r-iorth Carolina black walnut plantation (6) Deraeocoris nebulosus (7) Keltonia suphurea (8) Lygus linealaris (9) Plagiognathus politus (10) Reulerascopus omalus

extended period in the spring before the first adults were collected (ie early May) (Fig 9) It is apparently bivoltine near Asheville adults ofthe first (summer) generation were present from early May to late July and those of the second (overwintering) generation from early or mid-August to the end of the season

R ornatus has been collected from Ambrosia (Kelton 1964 Knight 1941) and Chenoshypodium album L (Knight 1941) No information has been published on its life cycle

In the present study R ornatus adults were found from late May to mid-October (Table 1) They were collected at all seven flying heights with more than 55 captured at 5-7 m (Fig 5)

This species probably overwintered as eggs this based on the same reasons as those given above for P politus The number of generations per year is unclear from the data available (Fig 10) However it is likely that at least one generation is represented by the few adult

25 6 sect 20 5 15E

0 10~ 1c

(J

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25

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42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

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O lOb 45 44

490gb 44 S

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050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

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3 49

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47 47 OOOb

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46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

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Page 11:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

42 THE GREAT LAKES ENTOMOLOGIST Vo 16 No2

specimens collected between late May and mid-August and a later generation by the drashymatic increase in numbers (ie increase in flight activity) in September Interestingly of the 2673 adults collected during this study almost 93 were captured in September and almost 50 during the third week of September (Fig 10)

ACKNOWLEDGMENTS

We wish to thank Mr D Brenneman and the staff of the North Carolina Division of Forestry Morganton for their help in collecting data and maintaining the window traps This research was partially supported by the USDA Forest Service North Central Forest Experiment Station

LITERATURE CITED

Froeschner R C 1949 Contributions to a synopsis of the Hemiptera of Missouri Pt IV Hebridae Mesoveliidae Cimicidae Anthocoridae Cryptostemmatidae Isometopidae Meridae (sic) Amer Midland Natur 42 123-188

Guppy J C 1958 Insect surveys ofclovers alfalfa and birdsfoot trefoil in eastern Ontario Canadian Entomo 90523-531

Kelton L A 1964 Revision of the genus Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (HemipteraMiridae) Canadian Entomo 961421-1433

---- 1975 The lygus bugs (genus Lyglls Hahn) of North America (Heteroptera Miridae) Mem Entomo Soc Canada 95 1-101

Knight H H 1927 Notes on the distribution and host plants of some North American Miridae (Hemiptera) Canadian Entomo 5934-44

---- 1941 The plant bugs or Miridae of Illinois Illinois Natur Hist Surv Bull 221-234

--- 1966 Keltonia a new genus near Reuteroscopus Kirk with descriptions of new species (HemipteraMiridae) Canadian Entomo 98590-591

Leonard M D 1915 The immature stages of Plagiognathus politus Uhler and Campylshyomma verbasci Herrick-Schaeffer (sic) (Capsidae Hemiptera) J New York Entomo Soc 23193-197

McPherson J E and B C Weber 1980 Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 1 Pentatomoidea Great Lakes Entomo 13177-183

---- 1981a Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 2 Coreoidea Great Lakes Entomo 14 11-13

---- 1981b Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 3 Reduvioidea Great Lakes Entomo 14 15-17

1981c Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 4 Cimicoidea Great Lakes Entomo 1419--22

1981d Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 5 Lygaeoidea Great Lakes Entomo 14 133-136

---- 1981e Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation 6 Tingidae and Aradidae Great Lakes Entomol 14 137-140

Ridgway R L and G G Gyrisco 1960 Studies of the biology of the tarnished plant bug Lygus lineolaris J Econ Entomo 531063-1065

Stewart R K and H Khoury 1976 The biology of Lygus lineolaris (Palisot de Beauvois) (HemipteraMiridae) in Quebec Ann Entomo Soc Quebec 2152-63

Wheeler A G Jr 1976 Lygus bugs as facultative predators p 28-35 in D R Scott and L E OKeefe (eds) Lygus bug host-plant interactions Unlv Press Idaho Moscow

Wheeler A G Jr B R Stinner and T J Henry 1975 Biology and nymphal stages of Deraeocoris nebulosus (HemipteraMiridae) a predator of arthropod pests on ornamenshytals Ann Entomo Soc Amer 681063-1068

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 12:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON

ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA PENT ATOMIDAE)

J E McPherson IT E Vogt I and S M Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pushybescence could be produced individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fallspring morpho

Thyanta calceata (Say) ranges from New England south to Florida and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971) This phytophagous stink bug exhibits adult dimorphism McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic green adults with short pubescence (shorter than diameter of tibia) are found during the summer months and brown adults with long pubescence during the fall and spring Adult dimorphism results from developmental photoperiod (McPherson 1977b 1975a) with a threshold photoperiod ofabout 125L 115D involved in the dimorphic response (McPherson 1975b) animals reared in photophases above and below the threshold develop into the summer and fallspring morphs respectively

To determine if the photophase during each 24 h period had to be continuous (eg 16 h) or could be split (eg S h S h) and still produce the same morph McPherson and Paskewitz (l9S2) reared animals under SL 16D 8L4D8L4D and J6LSD photoperiods The SL4D8L4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight24 h They found that those reared under SL 16D and SL4D8L4D became the fallspring morph and those in 16L8D the summer morpho Thus during each 24 h period it is the length of each photophase rather than the combined lengths of all photoshyphases that determine the adult morpho Also since seotophases of 16 h and 4 h were involved in the production of the fallspring morph and 8 h the production of the summer morph it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h This raised another question What was the length of the scotophase below which the animals would no longer respond but instead develop into the summer morph The results of an experiment to determine this are presented here

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incushybator (239 11C) under a 24LOD photoperiod the stock was established with indishyviduals collected July 1982 in Poinsett County Arkansas They were maintained in mason jars (five of each sexjar) provided with cheesecloth as an oviposition site a paper toweling strip and filter paper and fed green snap beans (Phaseolus vulgaris L) as described by McPherson (1971)

IDepartment of Zoology Southern Illinois University Carbondale IL 62901 2Department of Entomology University of Georgia Athens GA 30602

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

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Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

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Page 13:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

gt-l r tI1 a 0 tI1 gtTable I Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light gt-l t

Color gt tI1 rnDorsal Ventral Pubescence tI1 Z

Photoperiod Sex Brown Green Prob Brown Green Prob Short Long Prob gt-l 0 s

8L4D8L4D 0 49 49 I 0 50 0 9L3D9L3D 49 076a 48 2 050a I 49 050a t

0 a 9L3D9L3D 0 49 I 48 2 I 49

Hrn gt-l10L2D 1OL2D 48 2 050a 48 2 069a 2 48 050a

1OL2D 10L2D 0 48 2 48 2 2 48 IlL ID IlL 1D 19 31 3546b 19 31 3546b 18 32 1406b -lt

2 IlL ID IlL 1D 0 19 31 19 31 18 32 I15L05D 115L05D I 49 1806b I 49 1806b 50 0 Oooa ~

Z 115L05D 115L05D 0 I 49 I 49 50 0 24LOD 3 47 031a 3 47 031a 49 I 050a N

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

8lJ3 b 49

3

OAlb 43 45

O lOb 45 44

490gb 44 S

4AOb 8 I

050a I 0

Oooa 43 0

I 47 8113 b

0 49

7 5 O09b

3 3

5 6 OOOb

3 3

6 42 490S b

3 36

42 49 4AOb

36 50

49 50 050a

50 49

7 50 Oooa

3 49

50 I oooa

47 47 OOOb

47 47 OOOb

47 14 4304b

14 0 Oooa

0 1 050a

47 I 8113 b

i

gt-lr til Cl l til

~ ~ - til rJl

til Z as 0

sect rJl gt-l

v

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

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Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

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This publication is available in microform

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Page 14:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

00 8L4D8L4D 0 49 I 24LOD 3 47

8L4D8L4D yen 43 7 9L3D9L3D 46 4

9L3D9L3D S 46 4 IOL2D lor 2D 44 6

10L2D JOL2D yen 44 6 ilL 1D ilL 1D 8 42

IlL 1D IlL 1D yen 8 42 115L05D 115L05D I 49

115L05D 115L05D yen I 49 24LOD 0 50

8L4D8L4D yen 43 7 24LOD 0 50

exact probability test x 2 x test for independent samples corrected for continuity

Significant at the 005 level of probability

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46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 15:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

46 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971) 8L4D8L4D 9L3D9L3D 10L2D IOL2D IlL ID IlL ID II5L05D 115L05D and 24LOD All individuals were reared In239 I 1degC and in about 260 ft-c during the light phases (Sylvania 15W Daylight FI5T8D)

Adult characters compared were color (green or brown) and pubescence (long or short) short hairs were defined as those shorter than the diameter of the tibia The 005 level of significance was chosen for all comparisons

RESULTS AND DISCUSSION

Rearing males or females in 8L4D8L4D 9L3D9L3D and IOL2D IOL2D produced similar results most individuals were brown (males 96--98 females 86--92) with long pubescence (males 96--100 females 94) (Table I) Rearing the two sexes in llLID IlL I D produced a marked increase in the percentage ofgreen adults (males 62 females 84) and adults with short pubescence (males 36 females 72) Rearing individuals in 115L05D 115L05D produced a second increase (98 green adults 100 short pushybescence both sexes) and these percentages were similar to those for individuals reared in constant light

These results show that the developmental photophase generally must be longer than 2 h for the fallspring morph (brown with long pubescence) to be produced Between 2 hand 12 h most individuals no longer respond but appear as though reared in constant light (ie develop into green adults with short pubescence) As the length of the scotophase is deshycreased females generally fail to respond before males (recall that at IlL ID ilL ID 84 of the females were green compared to only 62 of the males) Thus scotophase as shown in the earlier experiment (McPherson and Paskewitz 1982) does function to break the photo phase but can be overridden if the scotophase is not of sufficient duration

ACKNOWLEDGMENT

We wish to thank Dr H E Barton Department of Biological Sciences Arkansas State University State University for sending us the specimens used in establishing the laborashytory culture

LffERATURE CITED

Blatchley W S 1926 Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida Nature Pub Co Indianapolis

McPherson J E 1971 Laboratory rearing of Euschistus tristigmus tristigmus J Econ Entomo 64 1339-1340

---- 1977a Notes on the biology of Thyanta calceata (HemiptemPentatomidae) with information on adult seasonal dimorphism Ann Entomol Soc Amer 70370-372

1977b Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (HemipteraPentatomidae) with information on ability of adults to change color Ann Entomo Soc Arner 70373-376

---- 1978a Sensitivity of immature TJryanta calceata (HemipteraPentatomidae) to photoperiod as reflected by adult color and pubescence Great Lakes Entomo 1171-76

1978b Effects of various photoperiods on color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo II 155-158

McPherson 1 E and S M Paskewitz 1982 Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (HemipteraPentatomidae) Great Lakes Entomo 1597-98

Oetting R D and T R Yonke 1971 Biology of some Missouri stink bugs 1 Kansas Entomo Soc 44446--459

1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE CERAMBVCIDAE AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF

AGRILUS BILINEATUS (COlEOPrERA BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1

Roben A Haack 2 Daniel M Benjamin3 and Kevin D Haack4

ABSTRACT

The species of Buprestidae Cerambycidae and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks Quercus spp in Wisconsin were Chrysoshybothris femorata (Olivier) and Dicerca sp (Buprestidae) Amniscus macula (Say) Cyrtashyphorus verrucosus (Olivier) Euderces picipes (Fabricius) Graphisurusfasciatus (DeGeer) Neodytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) (Cerambycidae) and Monarthrum fasciatum (Say) Monarthrum mali (Fitch) Pseudopityophthorus minutissimus (Zimmerman) and Xylaterinus paUlUS (Say) (Scolytidae) In general weakened oaks were first attacked by A bilineatus and at times that same year by C femorata G fasdalus and P minutissimus Infestation by Mfascishyatum M mali and X poUtus began the season following first attack by A bilineatus With the exception of A bilineatus the above mentioned Buprestidae and Cerambycidae apshypeared to preferentially infest dead wood often those portions that had died the previous season

The twolined chestnut borer Agrilus bilineatus (Weber) (Coleoptera Buprestidae) is a major pest of weakened oaks (Quercus spp) throughout eastern Nonh America Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought ice-storm damage and fall cankerworm Alsophila pomelaria (Harris) (Lepidopshytera Geometridae) defoliation The biology of A bilineatus has recently been studied by Dunbar and Stephens (1975 [976) in Connecticut Cote and Allen (1980) in New York and Pennsylvania and Haack Benjamin and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin

In declining oaks A bilineatus is normally the first borer to attack (Dunbar and Stephens 1975) Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years Attacked trees usually die during August and September of the second or third year of infestation Occasionally tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982)

Once an oak is attacked or killed by A bilineatu$ it becomes a suitable host to several other wood borers inner bark (phloem) borers bark beetles and ambrosia beetles in the families Buprestidae Cerambycidae and Scolytidae Study of borer succession is of practishycal imponance when attempting to assess the impact of several insect species on a host Nevertheless few such studies have been done Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina and Cote and Allen (1980)

I Research supported by the School of Natural Resources College of Agricultural and Life Sciences University of Wisconsin-Madison and tne Wisconsin Department of Natural Resources

2Department of Entomology and Nematology University of Florida Gainesville FL 32611 3Department of Entomology University of Wisconsin Madison WI 53706 4Department of Entomology Texas AampM University College Station TX 77843

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 17:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

48 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

listed the associated borers they had reared from A bilineatus-infested oak bolts We present here observations on the Buprestidae Cerambycidae and Scolytidae associated with A bilineatus in oaks in various stages of decline recorded in Wisconsin in 1979

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (04-()7 m2l made of nylon mosquito netting stapled at ca breast height (13 m) on 51 oaks (20-70 cm db h) in four host-condition categories Trees were assigned to a category based on external symptoms and signs of A bilineatus attack during fall 1978 and spring 1979 Cateshygories were (I) apparently healthy ie having no evidence of 1978 A bilineatus attack (2) some 1978 crown attack (3) tree death in 1978 and (4) tree death in 1977 Traps were installed in late May 1979 on 10 white oaks (Q alba 1) 7 black oaks (Q velutina Lam) 2 red oaks (Q rubra L) and I bur oak (Q macrocarpa Michx) in an oak woodlot near Madison Dane County Wisconsin and on 10 white oaks 7 black oaks 2 red oaks and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest Jefferson County Wisconsin Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September Adults were identified at the University of Wisconshysin and at the Insect Identitication and Beneficial Introduction Institute USDA Beltsville Maryland

In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A bilineshyatus larvae larval galleries and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site During that study we also recorded the number and location of all other borers encountered Briefly 25 red and black oaks (5category 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979 when symptoms of current-year A bilineshyatus attack were most evident Categories were (I) apparently healthy ie having no symptoms of 1979 A bilineatus attack (II) 25-50 crown death in 1979 after one season of attack (III) tree death in 1979 after one season of attack (IV) tree death in 1979 after two seasons of attack and (V) tree death in 1978 after at least two seasons of attack After felling bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca 5 cm We recorded the length and diam (inside the bark) of each bolt All borers were recovered from the bark cambial region and wood by carefully splitting with hammer and chisel numbers were recorded per square metre of bolt area The Scolytidae were identified to species However the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level using the keys of Burke (1917) Craighead (1923) and Peterson (1960) some larvae were reared to adults

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps The Buprestidae were A bilineatus and Chrysobothris femorata (Olivier) The Cerambycidae were Amniscus macula (Say) Cyrtophorus verrucosus (Olivier) Euderces picipes Fabricius) Graphisurus fasciatus (DeGeer) Neoclytus acuminatus (Fabricius) Sarosesthes fulminans (Fabricius) and Xylotrechus colonus (Fabricius) Table 1 presents number collected collection period host trees and host tree condition for each borer species

Borers were collected only from traps on dead oaks (categories 3 and 4) none were collected from the healthy or crown-attacked oaks However later inspection of the crownshyattacked oaks revealed A bilineatus exit holes in crown branches and along the upper bole of each but none along the lower trunks where the traps had been placed The most commonly collected cerambycid in our study was G fasciatus it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980) At times A bilineatus Cjemorata and G fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982 Chittenden 1905 and Craighead

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 18:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1 Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (13 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison Dane Co Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest Jefferson Co Wisconsin) from 20 May through 14 September 1979

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q alba (1) Q rubra (1) Q velutina (1) Chrysobothris femorata 8 13 June- 4 July Q alba (2) Q rubra (2) Q velutina (12)

CERAMBYCIDAE

Amniscus macula 2 1 June- 8 June Q velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q alba (2) Q velutina (2) Euderces picipes 3 3 June-ll June Q velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q alba (2) Q rubra (2) Q velutina (12) Neoclytus acuminatus 4 17 June-26 June Q rubra (2) Q velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q velutina (2) Xylotrechus colonus 13 8 June- 4 July Q rubra (2) Q velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period of adult emergence 1 ~ oaks dead for ca 1 year (died fall 1978) 2 ~ oaks dead for ca 2 years (died fall 1977)

1923 respectively) On no occasion did we collect C femorata or G fasciatus from cateshygory 3 oaks without also collecting A bilineatus but A bilineatus was at times the only borer recovered from a given trap Savely (1939) also found the above three borers as well as X colonus together in oak logs sampled within a year of felling Craighead (1923) reported a one-year life cycle for X colonus However Gardiner (1960) stated that two years were required for X colon us to complete development in Quebec A one-year life cycle has also been reported for N acuminatus (Baker 1972) A macula (Craighead 1923) and C verrucosus (Duffy 1953) To our knowledge voltinism data are not available for E picipes and S fulminans However Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood often the season following tree death If this was the case in our study then E picipes and S fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death

Although category I oaks had appeared unattacked two of the five oaks had been atshytacked by A bilineatus in 1979 Nevertheless as Haack and Benjamin (1982) reported all recovered A bilineatus larvae (n = 83) had died as first or second instars Only one other borer was found a Chrysobothris larva This larva was found alive in the cambial region of a branch sample (7 cm dia) where three dead A bilineatus larvae were recovered Apshyparently A bilineatus seldom attacks relatively vigorous oaks but when it does larval development is usually not successful Borers other than A bilineatus seem to attack vigorshyous oaks even less often the above Chrysobothris larva probably arrived once A bilineatus larvae were already present based on the apparent precedence ofthe latters galleries

Category II oaks had 25-50 crown death after one season of A bilineatus attack We recovered larvae of A bilineatus and Chrysobothris and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae) the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II and their mean densities are presented in Figure I Over 90 of the A bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin However A bilineatus larvae feed and develop primarily in phloem (inner bark) Two of the

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 19:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

50 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

80

N

~ 60

a 40 W

ID

~

) 20 z

0

CATEGORY II

BILINEATUS B - BUPRE STI OA E A_A

s shy SCOLYTIOAE S

--

AI _--- 05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 1 Mean Agrillls bilineatus Buprestidae (Chrysobothris) and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50lt crown death in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Chrysobothris larvae were found constructing pupal cells in the sapwood the others were in the cambial region P minutissimus was recovered from a single gallery system in a 4-cmshydia branch sample Also working in Wisconsin McMullen et al (1955) recorded two P minutissimus generations per year with flight periods in May and August and with lastshyinstar larvae being the overwintering form In our study the relatively warm fall temperashytures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI (1955) study (larvae) In southern Ohio P minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969) No empty P minutissimus galleries (representing the first generation of 1979) were observed This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969) Therefore the category II oaks were probably not suitable for P minutissimus colonization during its May flight period However by August these oaks were suitable for P minutissimus as a result of the A bilineatus branch-girdling that had taken place that summer

Category III oaks had died the year of sampling after one season of A bilineatus attack We recovered larvae of A bilineatus Chrysabothris and Graphisllrus and brood of P minutissimus (Table II Fig 2) Larval densities of A bilineatus were highest in the upper clear bole major branching began between 7 and 9 m in the oaks of this study Over 90 of the A bilineatlls were recovered from pupal cells We found 24 Chlysobothris larvae conshystructing pupal cells in sapwood and 37 in the cambial region All Graphisurus larvae were collected from the same tree two of the 17 larvae were constructing pupal cells in the cambial region We collected last-instar larvae pupae and callow adults of P minutissimus from one branch sample (5 cm dia)

Category IV oaks had died in the year of sampling after two seasons of A biineatus attack We collected larvae of A bilineatus Chrysobothris Graphisurus Neaciytus and Xylotrechus and brood of P minutissimus and the ambrosia beetles (Scolytidae) Monarshythrum Jasciatum (Say) Monarthrum mali (Fitch) and Xyloterinus paUtus (Say) (Table II Fig 3) Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries not numbers of individuals No living A bilineatus larvae were found above the 65-m sampling height where it had attacked the year before the average lowest extent of A bilineatus exit holes was ca 75 m Current A bilineatus larval densities

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 20:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120

100 (j

SO

0 UJ 60 al

gt

40z

20

0

A

1 S-~

B- BUPRESTIDAE

A1C- CERAMBYCIDAE

5 - SCOLYTIDAE

B__B_B

--c-c- - B C-le~

05 45 S5 125

HEIGHT ABOVE GROUND (M)

Fig 2 Mean Agrilus bilineatus Buprestidae (Chrysobothris) Cerambycidae (GraphisllruS) and Scolyshytidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decemshyber 1979 that had died in September 1979 after one season of A bilineatus attack Kettle Moraine State Forest Jefferson Co Wisconsin

100 CATEGORY IV

A 1 BlllHEATUS80 OJ

e 60

0

W 40

e gt z 20

0

45 85 125

HEIGHT ABOVE GROUND 1M J

Fig 3 Mean Agrillis bilineatlts Buprestidae (Chrysobothris) Cerambycidae (Graphisurus Neocytus XyJotrechlls) and Scolytidae (Monarthrllm Pselldopityophthorlls Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A hilineatlls attack Kettle Moraine State Forest Jefferson Co Wisconsin

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 21:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

52 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

were highest in the 25-m samples lower densities at 45 and 65 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferenshytially) or a combination of both We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region Chrysobothris densities were highest in the upper trunk of category IV oaks where A bilineatus had attacked the previous year Graphisurus larval densities were highest in the region (65 and 85 m) that represented the lowest extent of A bilineatus infestation during the previous year In the cambial region a few Graphshyisurus larvae but no Xylotrechus larvae were preparing pupal cells Xylotrechus and Neoshyelytus larvae were recovered primarily in samples from the zone between previous-year and current-year A bilineatus attack The active galleries of M Jasciatum and X POlilUS conshytained mostly callow adults and some pupae M mali galleries had callow adults only In Indiana X politus overwinters as adults and is univoltine (MacLean and Giese 1967) In Missouri M Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974) In Wisconsin X POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J O Haanstad pers comm) 5 Flight period data are not available for the Monarthrum species in Wisconsin but they are probably similar to that of X politus because we found galleries ofX poUtus and Monarthrum spp of similar length and having similar life stages in the same oaks sampled in June 1979 Therefore since the A bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982) the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles We also noticed in these category IV oaks that A bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each amshybrosia-beetle gallery suggesting that the staining (and thus the ambrosia beetles) preceded A bilineatus The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study traps were installed the third week of May which was after their peak flight The P minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia) that had been infested the previous year by A bilineatus indicating that this scolytid attacks material both conshycurrently with and the year follOWing A bilineatus attack

Category V oaks had died the year prior to sampling after at least two seasons of A bilineatus attack We found larvae of Chrysobothris Dicerca (Buprestidae) Cyrlophorus Graphisurus Neoclytus Sarosesthes and Xyiotrechlls and brood of X poUtus (Table 2 Fig 4) Chrysobothris larvae were found only in lower-trunk samples indicating that they infest material killed the previous year but not material killed two seasons earlier seven larvae were in sapwood and 17 in the cambial region However the Dicerca larvae were only found in samples from branches that had died two seasons earlier Savely (1939) collected Dicerca larvae from oak logs sampled ca two years after felling The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year Nevertheless a few Graphisurus Neoclytus and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier It is uncertain if these larvae had developed from eggs laid in 1978 or 1979 However since (I) most larvae appeared fully grown and were in or constructing pupal cells (2) cerambycid exit holes were already present in some of these same samples and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle Sarosesthes larvae were most common at 25 m this species appeared to prefer the lower trunk of oaks having died the previous season Active X politlls galleries were found mostly in lower trunk samples Apparently this scolytid attacks oaks the year after death but not portions having died two seasons earlier We found several old galleries ofMfasciatum M mali and X poUtus that were active the previous season old P mifllltissimus galleries were also present

5Department of Entomology University of Wisconsin Madison WI 53706

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Page 22:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2 Percent of oak samples containing Buprestidae Cerambycidae and Scolytidae described by host condition category and height above ground (5 oakscategory Kettle Moraine State Forest Jefferson Co Wisconsin December 1979)

Sampling Heights (m)

Borer 05 25 45 65 85 105 125 145 (Na) Siteb

Category II Oaks

A bilineatus Chrysobothris P minutissimus

80 80 100 60

100 40

100 20

100

20

(186) (6) (8)

BCSW CSW C

Category III Oaks

A bilineatus Chrysobothris Graphisurus P minutissimus

100 60

100 40 20

100 40 20

100 20 20

100 20 20

100 20 20

100 60 20

100 20

20

(638) (61) (17) (14)

BCSW CSW C C

Category IV Oaks

A bilineatus Chrysobothris Graphisurus Neoclytus Xylotrechus M fasciatum M mali P minutissimus X poUtus

100 20

20

60

100 80 60

20 20

60

100 80 80

40

80

60 100 80 60 80

60

100 60 60 80

20

80

20

20

40 20

20

(230) (147) (137) (11) (14)

(3) (3)

(14) (42)

BCSW CSW C SW C SW SW C SW

Category V Oaks

Chrysobothris Dicerca Cyrtophorus Graphisurus Neoclytus Sarosesthes Xylotrechus X poUtus

60

100

60 80 80

80

20 100 20

100 100 80

40

20 100 60 60

100 80

20 100 40 20 60 20

20 20 20 40

80

20

20 40

(24) (2) (4)

(99) (11) (28) (48) (40)

CSW C SW C SW CSW C SW

aN the number of individuals (or gallery systems for tile ambrosia beetles M jasciatum M mali X politus) recovered from all samples for the indicated host condition category of oaks bSite refers to the area(s) in which each borer was recovered B outer bark C cambial region SW sapwood

SUMMARY

A successional pattern of Buprestidae Cerambycidae and Scolytidae can now be apshyproximated for a typical declining oak in Wisconsin In general A bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one At times A bilineatus is joined that first summer by small numbers of Chrysobothris

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

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Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

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Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

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This publication is available in microform

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Page 23:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

54 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

50

40 N

Ii

- 30 a w ro 20 Ii l

Z 10

0

CATEGORY V

Cc 8

C ~C C~ CERAMSYCIDAE

S- SCOLnC

C

5-i- ~ -5 Ca a- shy~Agt-C~---B~B

05 45 85 125

HEIGHT ABOVE GROUND (M)

Fig 4 Mean Buprestidae (Chrysobolhris Dicerca) Cerambycidae (Cyrlophorus Graphisums Neoshyelyus Saroseslhes Xylotrechus) and Scolytidae (Xyoteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA bilineaus attack Kettle Moraine State Forest Jefferson Co Wisconsin

Graphisurus and P minutissimus During spring of year two the lower trunk is colonized by ambrosia beetles such as M fasciatum M mali and X poUtus In the summer of year two A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries Also at this time Chrysobothris and Graphisurus may join A bilineatus in the lower trunk and also reattack the upper trunk and crown along with P minutissimus Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus In spring of year three X poUtus continues to colonize the lower trunk which had died in the fall of year two In the summer of year three following the exit of A bilineatus the lower trunk is now infested by Graphshyisurus Neociytus and Xylotrechus as well as Cyrtoplwrus and Sarosesthes The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation In this way the insect fauna in the crown branches upper trunk and lower trunk changes from year to year during the decline of an oak tree and also after its death

ACKNOWLEDGMENTS

We thank Dr Dave J Hall and Paul E Pingrey Wisconsin Department of Natural Reshysources for assistance in locating forest stands Tom Byrnes for access to his oak woodlot John O Haanstad for use of his Xyloterinus politus emergence data Dr D M Anderson Insect Identification and Beneficial Insect Introduction Institute USDA Beltsville MD for identification of the Scolytidae and Dr Douglas C Allen Dr David G Nielsen Dr James D Solomon and John O Haanstad for critical review ofthis manuscript

LITERATURE CITED

Baker W L 1972 Eastern forest insects USDA Misc Pub 1175 Burke H E 1917 Flat-headed borers affecting forest trees in the United States USDA

Bull 437 Chittenden F H 1905 The flat-headed apple tree borer (Chrysobothris femorata) USDA

Bur Entomo 32 rev

1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

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Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

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This publication is available in microform

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1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote W A III and D C Allen 1980 Biology of two-lined chestnut borer Agrilus bilineatus in Pennsylvania and New York Ann Entomol Soc Amer 73409-413

Craighead F C 1923 North american cerambycid larvae Canadian Dept Agric Entomol Branch Bull 27

Duffy E A J 1953 A monograph of the immature stages of British and imported timber beetles (Ceram bycidae) British Mus (Natur Hist) London

Dunbar D M and G R Stephens 1975 Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut For Sci 21 169-174

---- 1976 The bionomics of the two lined chestnut borer p 73--83 in I F Anderson and H K Kaya (eds) Perspectives in forest entomology Academic Press New York

Gardiner L M 1960 Description of immature forms and biology of Xylntrechus colonus (Fab) (Coleoptera Cerambycidae) Canadian Entomo 92820-S25

Haack R A and D M Benjamin 1982 The biology and ecology of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) on oaks Quercus spp in Wisconsin Canadian Entomo 114385-396

Haack R A D M Benjamin and B A Schuh 1981 Observations on the biology of Phasgonophora sulcata (Hymenoptera Chalcididae) a larval parasitoid of the twolined chestnut borer Agrilus bilineatus (Coleoptera Buprestidae) in Wisconsin Great Lakes Entomo 14 113-116

Knul J N 1946 The longhorned beetles of Ohio (Coleoptera Cerambycidae) Ohio BioI Surv Bull 39 133-354

MacLean D B and R L Giese 1967 The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera Scolytidae) Canadian Entomo 99285-299

McMullen L H bull E W King and R D Shenefelt 1955 The oak bark beetle Pseudoshypityophthorus minutissimus (Zimm) (Coleoptera Scolytidae) and its biology in Wisconshysin Canadian Enlomol 87755-757

Peterson A 1960 Larvae of insects Edwards Brothers Ann Arbor Rexrode C O 1969 Seasonal development and habits of Pseudopityophthorus spp

(Coleoptera Scolytidae) in southern Ohio Canadian Entomol 101306-313 Roling M P and W H Kearby 1974 Life stages and development of Monarthrum

fasciatum (Coleoptera Scolytidae) in dying and dead oak trees Canadian Entomo 106 1301-1308

Savely H E If 1939 Ecological relations of certain animals in dead pine and oak logs Ecol Monog 9321-385

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 25:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G Voss i and Richard E Riefner Jr 2

As early as 1912 mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid Habenaria obtusata (Pursh) Richardson in Reeses Bog a cedar swamp at the north end of Burt Lake Cheboygan County Michigan near the campus of the University of Michigan Biological Station (Dexter 1913) A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosquishytoes always females of the genus Aedes Details of mosquito behavior and the pollination process have recently been described with excellent pictures by Stoutamire (1968) Thien (1969) and Thien amp Utech (1970) These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia but no other moths or insects of other orders have been reported as pollinators

The blunt-leaf orchid is a circumpolar species ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St Clair County) Toward this southern limit of its range it favors coniferous swamps and forests where the soil is cold and often moist The inconspicuous plants are rarely over 20 cm tall with a single leaf and a few greenish flowers about 5---) mm long scarcely larger than the mosquitoes which pollinate them While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives they have no monopoly on pollination We now report as an apparent pollinator a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted)

Anageshna primordialis (Dyar) was originally placed (1906) in Geshna but was transshyferred by Munroe (1956) to a monotypic new genus It is now classified in the Pyralidae subfamily Pyraustinae (not Nymphulinae as originally thought) Forbes (1923 p 581) who had first questioned its taxonomic placement stated its occurrence as Very common in damp places in June and early July Indeed it seems to be a rather common moth in the cedar swamps of northern Michigan although (like most microlepidoptera) not previously reported from the state in any literature we have seen The wingspread is barely 15 mm and the wings are brownish with transverse bands of rather angular pale spots (Fig I) Specishymens in the collections of the University of Michigan Museum of Zoology and the Entoshymology Museum of the Department of Entomology Michigan State University indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula ie the entire range of the orchid in the state the moth has also been collected farther south in Berrien Livingston Oakland and Washtenaw counties Dates in the northern part of the state are 16 June-21 July None of the museum specimens bear pollinia perhaps because they were probably not collected in the habitat of this orchid

On 24 June 1976 the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reeses Bog where mosquitoes continue to be major pollinators Eventually the moth pulled itself free without removing the pollinium However on 25 June 1981 a moth similarly stuck removed the pollinium firmly attached to an eye the usual location for pollinia of this species to become cemented on insect visitors (Fig 2) One specimen captured in free flight bore two pollinia both attached to the right eye (Fig 3)

This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers after probing a flower it pushes its proboscis into the spur and feeds upon

iHerbarium The University of Michigan Ann Arbor MI 48109 27439 La Palma Ave Suite 206 Buena Park CA 90620

58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

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58 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

Fig I Anageshna primordialis from a flower of Habenuriu obusaa Reeses Bog Cheboygan Co Michigan 24 June 1976 Wingspan is 15 mm (Photo by D Bay)

Fig Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the orHu n Reeses Bog 25 June 1981 Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye (Photo by D Bay)

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 27:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig 3 Anageshna primordialis with two pollinia attached to the right eye collected in flight in Reeses Bog 25 June 1981 (Photo by R Riefner)

nectar before taking flight Moths bearing pollinia have been captured into the evening hours The moths are apparently not attracted to the flowers from a distance but as sugshygested by Stoutamire (1968) for mosquitoes move toward the flowers after coming to rest on the inflorescence This behavior seems to suggest attractants perceived close at hand The flowers tested UV- and have no scent perceptible to us

Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar especially from spurs only partly filled Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes Anageshna like the mosquitoes ranges well to the south of the orchid (into Florida Kimball 1965 p 219) and clearly is not closely dependent upon it In fact it appears to be a moth of quite catholic tastes having been reported to visit pig carrion in a state of advanced decay (Payne amp King 1969)

ACKNOWLEDGMENTS

We are grateful to Dr Eugene Munroe (Ottawa Ontario) for identifying the moths we collected to David Bay for photographic assistance to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station The specimens shown in Figures 1 and 2 are in the collection of the senior author that in Figure 3 will be placed in the collection of the Biological Station

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 28:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

60 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

LITERATURE CITED

Dexter John Smith 1913 Mosquitoes pollinating orchids Science 37867 Dyar Harrison G 1906 The North American Nymphu1inae and Scopariinae J New York

Entomol Soc 1477-107 Forbes William T M 1923 The Lepidoptera of New York and neighboring states Cornell

Vniv Agric Exp Sta Mem 68 729 p Kimball Charles P 1965 The Lepidoptera of Florida an annotated checklist Arthropods of

Florida and neighboring land areas Vol 1 Florida Dept Agric Gainesville 363 p + 26 pI

Munroe Eugene 1956 Geshna primordialis Dyar with descriptions of two new genera and two new subspecies (Lepidoptera Pyralidae) Canadian Entomol 88126-128

Payne Jerry A and Edwin W King 1969 Lepidoptera associated with pig carrion J Lepid Soc 23 191-195

Stoutamire Warren P 1968 Mosquito pollination of Habenaria obtusata (Orchidaceae) Michigan Bot 7203-212

Thien Leonard B 1969 Mosquito pollination of Habenaria obtusata (Orchidaceae) Amer J Bot 56232-237

Thien Leonard B and Frederick Utech 1970 The mode of pollination in Habenaria obtusata (Orchidaceae) Amer J Bot 57 1031-1035

Voss Edward G 1972 Michigan Flora Part I Gymnosperms and Monocots Cranbrook Inst Sci Bull 55 488 p

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 29:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA ARANEAE)

Brent D Opell l

ABSTRACT

Names synonyms and distributions ofu10borid spiders known from North Central and South America are provided

The first comprehensive revision of American Uloboridae was Muma and Gertschs (1964) treatment of the familys United States and Canadian members This was followed by Lehtinens (1967) reevaluation of the familys generic division Chickerings (1968) study of Panamanian and West Indian Miagrammopes and Opells (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes Studies by Lubin et al (1982) and Opell (1981 1982) broughtthe number of known American uloborid genera to 14 and the number of recognized species to 71

The purpose ofthis list is to provide ready access to the names synonyms nomenclatural changes and general distribution of the American Uloboridae As a revision of Miagramshymopes sensu lato (Huanacauria Miagrammopes Mumaia Miagramopsidis and Ranguma the latter two not represented in the New World) nearing completion will render obsolete any listing of these species they are only summarized Included are all other known taxa except the following which largely due to the unavailability of type material are of uncertain status Uloborus minutus Mello-Leimo U tetramaculatus Mello-Leimo U formosus Marx (in Banks) U ater Mello-Leimo Petrunkevitchia venusta Mello-Leitao and P pusilla Mello-Leitao Genera are arranged according to Opells (1979) phylogenetic hypothesis Within each genus species are arranged alphabetically the only exception being in Philoponella where species are listed under three groups The type species of each genus if not included in a generic synonym or listed below the genus name is marked by an asterisk A colon following a name denotes the first transfer of the species to the cited genus

Five genera (Uloborus Zosis and Philoponella Miagrammopes and the closely related Mumaia) and four species (U campestratus U segregatus Z geniculatus and P semiplumosa) are found in North Central and South America Uloborus trilineatus P republicana and P tingena are found in both Central and South America and Mumaia mexican a in both North and Central America As Figure 1 shows the greatest number and percent of endemic genera and species are found in South America and the least in North America Because this evaluation may be biased by the number of named Miagrammopes species totals exclusive of these species are also provided

ACKNOWLEDGMENTS

Collections made by William G Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species Preparation of this list was supported in part by National Science Foundation grant DEB-8011713

1 Department of Biology Virginia Polytechnic Institute and State University Blacksburg V A 24061

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 30:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

62 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA

(South Qf Panama)(Canada Uniteltl (5 Mexico through States ~ Mexico) Panama [ I Indies)

Coni faber Sybota [IDEMIC ~ENERA Ariston I-kianacauria

Orinomana Ponella

TOTAL NLMlER 6 10

OF GENERA

EI()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER

Of SPECIES 16 (15) 31 (12) 36 (22)

Fig I Distribution and number of known American uloborid taxa Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria Miagrammopes and Mumaia

ARISTON O Pickard-Cambridge 1896 3 species

A albicans O Pickard-Cambridge 1896 SE Mexico and Honduras A arisus OpeII 1979 Panama A mazolus Opell1979 SW Mexico

HYPTIOTES Walckenaer 1837 4 species

Mithras (preoccupied) C L Koch 1834 Type-M paradoxlls C Koch 1834 Vptiotes WaIckenaer 1837 Type-V anceps Walckenaer 1837 Hyptiotes Erickson 1845 emendation Cylfopodia Hentz 1847 Type-C cavaus Hentz 1847

H cavaus (Hentz) 1847 E half of US and Canada H gerschi Chamberlin amp Ivie 1935 W US and Canada H pllebla Muma amp Gertsch 1964 SW US H tehama Muma amp Gertsch 1964 Central W US

SIRATOBA Opell 1979 2 species

Adson (in part) O Pickard-Cambridge 1896

S referena (Muma amp Gertsch) 1964 SW US and N Mexico S sira Opell 1979 Central Mexico

HUANACAURIA Lehtinen 1967 Type-Miagrammopes bambusicola Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

3 species Brazil (2) Venezuela (I)

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 31:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O Pickard-Cambridge 1869 Type-M thwaitesii O Pickard-Cambridge 1869

22 species Antilles (I) Brazil (6) Cuba (2) Guiana (I) Jamaica (3) Panama (7) Puerto Rico (I) Venezuela (I) Virgin Islands (I)

MUMAIA Lehtinen 1967 Type-Miagrammopes corticeus Simon 1892

Miagrammopes (in part) O Pickard-Cambridge 1869

6 species Antiles (I) Argentina (I) Mexico (I) Panama (I) Puerto Rico (2) St Vincent (I) US (I) Venezuela (I) Virgin Islands (I)

SYBOTA Simon 1892 3 species

Sylvia (preoccupied) Nicolet 1849 Sybota Simon 1892 replacement name for Sylvia

S abdominalis (Nicolet) 1849 Chile Sylvia abdominalis Nicolet 1849 Sylvia ater Nicolet 1849 Sylvia similis Nicolet 1849 Sylvia rubiginosa Nicolet 1849 Sylvia vittata Nicolet 1849 Uloborus abdominalis Simon 1887 Sybota abdominalis Simon 1892

S mendozae Opell 1979 Argentina S osornis Opell 1979 Chile

ORINOMANA Strand 1934 2 species

Orinomus (preoccupied) Chamberlin 1916 Orinomana Strand 1934 replacement name for Orinomus

O bituberculata (Keyserling) 1882 Peru Uloborus bituberculatus Keyserling 1882 Orinomus lamprus Chamberlin 1916 Orinomana lampra Strand 1934

O mana Opell 1979 Chile

ULOBORUS Latreille 1806 8 species

Uloborus Latreille 1806 Type-Uloborus walckenaerius Latreille 1806 Phillyra Hentz 1850 Type-Phillyra mammeata Hentz 1850 Veleda Blackwall 1859 Type-Veleda lineata Blackwall 1859 Philoponus Thorell 1887 Type-Philoponus pteropus Thorell 1887

U campestratus Simon 1893 SE US through N South America U cinereus O Pickard-Cambridge 1898 U spernax O Pickard-Cambridge 1898

U diversus Marx in Banks 1898 W US and NW Mexico U albineus Marx in Banks 1898 U californicus Banks 1904 U utahensis Chamberlin 1919 U saphes Chamberlin 1924 U crepedinis Chamberlin 1924

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 32:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

64 THE GREAT LAKES ENTOMOLOGIST Vol 16 No2

U eberhardi Opel 1981 Costa Rica U eongatus Open 1982 Argentina U glomosus (Walckenaer) 1847 (1842) E half of US amp Canada N Mexico

Epeire gomosa Walckenaer 1837 (1842) Phillyra mammeata Hentz 1850 Phillyra riparia Hentz 1850 Uloboms plumipes Emerton 1888 Uloborus americanus Comstock 1912 (1913) Uloborus glomosus Chamberlin amp Ivie 1944

U metae Opell 1981 E central Colombia U segregatus Gertsch 1936 Texas through N South America U trilineatus Keyserling 1882 S Mexico through South America

U penicillatus Simon 1891 U aegrotus Simon 1893 U maniculatus Simon 1893 U bucki Mello-Leitao 1943 U plumipes Mello-Leitao 1947 U plumipedatus Roewer 1951 replacement name for

U plumipes MelJo-Leitao preoccupied by U plumipes Lucas 1846

ZOSIS Walckenaer 1837 2 species

Uloborus (in part) Latreille 1806 Zosis Walckenaer 1837 Type-Zosis caraibe Walckenaer 1837 Orithyia Blackwall 1858 Type-Orithyia williamsii Blackwall 1858

Z geniculatus (Olivier) 1789 pantropical Aranea geniculata Olivier 1789 Zosis caraibe Walckenaer 1837 Uloborus zosis Walckenaer 1841 Uloborus latreillei Thorell 1858 Orithyia williamsii B1ackwall 1858 Uloborus domesticus Doleschall 1859 Orithyia luteola Blackwell 1865 Uloborus williamsii O Pickard-Cambridge 1871 Uloborus geniculatus Thorell 1890 Ulobarus luteolus Roewer 1954 Zosis geniculatus Lehtinen 1967

Z peruvianus (Keyserling) 1882 E central Colombia to N central Brazil to N central Argentina Uobrus peruanus Keyserling 1882 Uloborus peruvianus Bonnet 1959 emendation Zosis peruvianus Opell 1981

OCTONOBA Opell 1979 1 species

Uloborus (in part) Latreille 1806

0 octonaria (Muma) 1945 Central and SE US Uloborus octonarius Muma 1945 Octonoba octonaria OpeU 1979

CONIFABER Opel in Lubin et al 1982 1 species

Conifaber panus Opell in Lubin et al 1982 E central Colombia

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 33:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel 1979 1 species

Uloborus (in part) Latreille 1806

P lactescena (Melo-Leitao) 1947 Paraguay and S BraziL Uloborus lactescens Mello-Leitao 1947 Panella lactescena OpeU 1979

PHILOPONELLA MelomiddotLeitao 1917 15 species

Uloborus (in part) Latreille 1806 Philopanella MelomiddotLeitao 1917

Philoponella republicana species group Opell 1979

P divisa Opell 1979 N South America P republicana (Simon) 1891 Panama amp South America

Uloborus republicana Simon 1891 Uloborus cuminamensis MellomiddotLeitiio 1930 Uloborus mundior Chamberlin amp Ivie 1936 Zosis mundior Lehtinen 1967 Philopenella republicana Opell 1979

P signatella (Roewer) 1951 SE Mexico through Honduras Uloboms signatus O PickardmiddotCambridge 1898 Uloborus signatellus Roewer 1951 replacement name for preoccupied

U signatus O PickardmiddotCambridge 1898 Philoponella signatella Opel 1979

P tingena (Chamberlin amp Ivie) 1936 Costa Rica through Colombia Uloborus tingnes Chamberlin amp Ivie 1936 Philoponella tingena Opell 1979

P viGina (0 Pickard-Cambridge) 1898 S Mexico through Central America Uloborus vicinus O PickardmiddotCambridge 1898 Philoponella vicina Opell 1979

Philoponella semiplumosa species group Opel 1979

P arizonica (Gertsch) 1936 Arizona Uloborus arizonicus Gertsch 1936 Philoponella arizonica Opell 1979

P oweni (Chamberlin) 1924 SW US Uloboms oweni Chamberlin 1924 Philoponella oweni Opell 1979

P semiplumosa (Simon) 1893 Texas through N South America Uloborus semiplumosus Simon 1893 Uloborus variegatus O PickardmiddotCambridge 1898 Uloborus abstrusus Gertsch amp Davis 1942 Philoponella semiplumosa OpeU 1979

P subvittata Opell 1981 Guyana P viltala (Keyserling) 1882 South America

Uloborus vittatus Keyserling 1882 Uloborus servulus Simon 1892 Uloborus semiargenteus Simon 1893 Uloborus amazonicus Mello-Leitao 1949 Philoponella vittata Opell 1979

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 34:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

66 THE GREAT LAKES ENTOMOLOGIST Vol 16 No 2

Philoponella Jasciata species group Opel 1979

P bella Opell 1979 NE Colombia P collina (Keyserling) 1882 Peru

Uloborus collin us Keyserling 1882 Philoponella collina Opel 1981

P Jasciata Mello-Leitao 1917 SE Brazil amp Paraguay Uloborus Jasciatus Mello-Leimo 1917 Philoponella Jasciata Opel 1979

P para Opell 1979 Paraguay

LITERATURE CITED

Chickering A M 1968 The genus Miagrammopes (Araneae Uloboridae) in Panama and the West Indies Breviora 2891-28

Lehtinen P T 17 Classification of the cribellate spiders and some allied families with notes on the evolution of the suborder Araneomorpha Ann Zool Fennici 4 199-468

Lubin Y D B D Open W G Eberhard and H W Levi 1982 Orb plus conewebs in Uloboridae (Araneae) with a description of a new genus and four new species Psyche 8929-M

Muma M M and W 1 Gertsch 1964 The spider family Uloboridae in North America north of Mexico Amer Mus Novitates 211-43

Opell B D 1979 Revision of the genera and tropical American species of the spider family Uloboridae Bull Mus Compo Zool 148443-549

---- 1981 New Central and South American Uloboridae (Arachnida Araneac) Bull Amer Mus Nat Hist 170219-228

---- 1982 A new Uloborus Latreille species from Argentina (Arachnida Araneae Uloboridae) Proc BioI Soc Washington 95552-554

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England

Page 35:  · THE GREAT LAKES ENTOMOLOGIST . Published by the Michigan Entomological Society . Volume 16 No.2 . ISSN 0090-0222 . TABLE OF CONTENTS . Seasonal Flight Patterns of Hemiptera in

INFORMATION FOR AUTHORS

Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomologist Appropriate subjects are those of interest to amateur and proshyfessional entomologists in the North Central States and Canada as well as general papers and revisions directed to a larger audience while retaining an interest to readers in our geographical area

Manuscripts must be typed double-spaced with wide margins on white 8~ x 11 or equivalent size paper and submitted in duplicate Footnotes legends and captions for illustrations should be typed on separate sheets of paper Titles should be concise identifyshying the order and family discussed The author of each species mentioned must be given fully at least once in the text A common name for each species or group should be given at least once when such a name exists The format of references should follow that used in recent issues Photographs should be glossy Drawings charts graphs and maps must be scaled to permit proper reduction without loss of detail Contributors should follow the recommendashytions of the Council of Biology Editors Style Manual 4th ed

Papers published in The Great Lakes Entomologist are subject to a page charge of $3000 per published page Members of the Society who are authors without funds available from grants institutions or industry and who are unable to pay costs from personal funds may apply to the Society for financial assistance

Authors will receive page proof together with an order blank for separates Short papers on new ecological information distribution records and similar topics are

welcomed for publication in the Entomological Notes section All manuscripts for The Great Lakes Entomologist should be sent to the Editor Dr D C

L Gosling 69063 Wallowa Road White Pigeon Michigan 49099 USA Other correshyspondence should be directed to the Executive Secretary (see inside front cover)

This publication is available in microform

University Microfilms International

300 North Zeeb Road 30-32 Mortimer Street Dept PR Dept PR Ann Arbor Mi 48106 London WIN 7RA USA England