The I GREAT LAKES ENTOMOLOGIST2 THE GREAT LAKES ENTOMOLOGIST Vol. 9, No. 1 southern edge, from 4:00 p.m. to 6:OOp.m. on the west edge, and from 6:OOp.m. to 8:00 p.m. on the northern

The

I GREAT L A K E S E N T O M O L O G I S T

Vol. 9, No. 1 Spring 1976

Volume 9

THE GREAT LAKES ENTOMOLOGIST

Published by the Michigan EntomoIogical Society

No. 1

TABLE OF CONTENTS

Keys and Notes on the Buprestidae (Coleoptera) of Michigan Stanley G. Wellso, Gary V. Manley, and John A. Jackman . . . . . . . . . . . . 1

Indiana Ensifera (Orthoptera) W. P. McCafferty and J. L. Stein . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Insects Associated with Michigan Bumblebees (Bombus spp.) Robert W. Husband and Thomas M. Brown. . . . . . . . . . . . . . . . . . . . . . 57

Neurocordulia (Platycordulia) xanthosoma (Williamson) in Texas (Odonata: Libellulidae: Corduliinae)

Curtis E. Williams. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

The Subcolonization and Buildup of Tetrastichus julis (Hymenoptera: Eulophidae), a Larval Parasitoid of the Cereal Leaf Beetle (Coleoptera: C~Q-somelidae). in the Lower Peninsula of Michigan

P. A. Logan, F. W. Stehr, and R. J. Sauer . . . . . . . . . . . . . . . . . . . . . . 75

Description of the Lama of Dioedus punctatus LeConte (Coleoptera: Tenebrionidae)

Daniel K. Young . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

COVER ILLUSTRATION

Scanning electron micrograph of Brachys ovatus (Weber) (Coleoptera: Buprestidae) 14X

Vol. 8, No. 4 of The Great Lakes Entomologist was mailed on March 1: 1976.

THE MICHIGAN ENTOMOLOGICAL SOCIETY

1975-76 OFFICERS

President President-Elect Executive Secretary Editor

David C. L. Gosling Donald C. Cress

M. C. Nielsen D. C. L. Gosling

The Michigan Entontological Society traces its origins to the old Detroit Entomo- logical Society and was organized on 4 November 1954 to ". . . promote the science of entomology 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. Mentbership in the Society, which serves the North Central States and adjacent Canada, is open to all persons interested in enton~ology. There are four paying classes of membership:

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SURSCRIPTION INFORMATION

lnstitutions and organizations, as well as individuals not desiring the benefits of ~nentbership, may subscribe to The Great Lakes Enton~ologist at the rate of $10.00 per volume. The journal is published quarterly; subscriptions are accepted only on a volume (4 issue) basis. Single copies of The Great Lakes Entorr~ologist are available at $2.75 each, with a 20 per cent discount for 25 or more copies sent to a single address.

MICROFILM EDITION: Positive microfilm copies of the current volume of The Great Lakes Entomologist will be available at nominal cost, to members and bona fide sub- scribers of the paper edition only, at the end of each volunle year. Please address all orders and inquiries to University Microfilms, Inc., 300 North Zeeb Road, Ann Arbor, Michigan 48106, USA.

Inquiries about back numbers, subscriptions and Society business should be directed to the Executive Secretary, Michigan Enton~ological 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 @ 1976, The Michigan Entomological Society


KEYS AND NOTES ON THE BUPRESTIDAE (COLEOPTERA) OF MICHIGANI

Stanley G. Wellso,2 Gary V. Manley,3 and John A. Jackman4

ABSTRACT

The d i s ~ b u t i o n and dates of adult activity for Michigan buprestids are discussed. Keys to the genera and species, as well as host information are presented for 116 species and one subspecies. Information on collecting techniques, illustrations of genitalia of 14 species, and scanning electron micrographs of certain structures useful in species identification are presented and discussed. In addition, Pachyschelus confusus, a new species, is described from bush clover.

INTRODUCTION

Recent North American regional papers on buprestids are those by Knull (1925) for Pennsylvania, Franklin and Lund (1956) for Georgia, and Ban (1971) for the Pacific Northwest. In this paper, we present information on general buprestid collecting techniques, keys, a new species description, distribution, collecting dates and hosts.

The data have been transcribed from specimens in the authors' collections (SGWC, GVMC, JAJC) and those of the following individuals and institutions: Michigan State University (MSUC), Purdue University (PURC), University of Michigan (UMMZ), Univer- sity of Wisconsin (UWEM), Josef N. Knull (JNKC), Dr. Gayle Nelson (GHNC), and Daniel Young (DYCC). In addition, information previously presented on Michigan species of buprestids by Hubbard and Schwarz (1878) who reported 38 species from the Lower Peninsula and 30 species from the Upper Peninsula, Townsend (1898) who reported three species, and Andrews (1921) who reported 32 species from the Upper Peninsula, has been noted; however, because of synonymy, not all of these records refer to valid species.

COLLECTION TECHNIQUES

Buprestid adults in collections generally do not reflect the abundance of these beetles in our fauna Because the adult life span approximates a month in duration the host must be known to assure efficient collecting. The following suggestions are presented to enable collectors to improve their collections of adult buprestids.

(1) The most important method is to cage larval host wood so that more information is accumulated about the site of adult oviposition and the host range of the species. Often adults deemed 'unusual' or 'rare' can be reared in this manner. For example, it is difficult to capture adult Agrilus charnplaini Frost on its host hop hornbeam, Ostyra virginiana; however, this species can be readily obtained by collecting its galls from hop-hornbeam and rearing the adults in the laboratory.

(2) More specimens, especially species of Agrilus, Brachys, Anthaxia and Chryso- bothris, can be swept from the edge of dense deciduous woods than from the interior.

(3) One should concentrate collecting at temperatures above 20°C. Also more specimens are usually taken on clear rather than cloudy days and on still rather than windy days.

(4) Collecting in full sunlight seems to be important; thus, from 10:OO a,m. to noon collect at the southeastern edge of a wood lot, from noon to 3:00 p.m. collect at the

]Michigan Agricultural Experiment Station Journal Article 7179. Part of a cooperative project between the Agricultural Research Service, USDA, and the Department of Entomology, Michigan State University, East Lansing, Michigan 48824.

2 ~ ~ ~ ~ , Agricultural Research Service, Department of Entomology, Michigan State University, East Lansing 48824.

37907 Rustic Forest, San Antonio, Texas 78239. 4~epar tment of Entomology, T e x a ~ A. & M. University, College Station, Texas 77844.

2 THE GREAT LAKES ENTOMOLOGIST Vol. 9, No. 1

southern edge, from 4:00 p.m. to 6:OOp.m. on the west edge, and from 6:OOp.m. to 8:00 p.m. on the northern edge in June. A net with a handle of 25 feet in length is useful, since a larger area of leaves exposed to full sunlight may be swept (Fig. 1).

(5) Other sites that warrant observation are trees and shrubs that have been injured or killed, preferably within a few months prior to adult activity. These trees and shrubs are

Fig. 1. A net with a 25-foot aluminum handle enable's a collector to sweep a much larger vertical area.

1976 THE GREAT LAKES ENTOMOLOGIST 3

attractive as oviposition sites, and beetles can be captured by hand or net from them. Tree dumps often seem very unproductive, if the collector concentrates only on the dead logs; however, buprestids emerging from the dead trees may be abundant on adjacent vegetation To improve collecting, note the prevailing winds and collect on downwind logs or vegetation. In this situation, it is possible to collect many adults from atypical plants which should not be listed as hosts (e.g., 25 species of buprestids were collected from willow, Sulix sp., in a tree dump).

GENERAL INFORMATION

Eighteen of the 42 North American genera of buprestids including 116 of the approximately 760 species have been collected in Michigan. For convenience and because of floral differences, Michigan has been divided into three zones (Fig. 2). The number of

SOUTHERN LOWER

UPPER

NORTHERN

Fig. 2. Michigan's three zones: Upper (U), Northern Lower (NL), and Southern Lower (SL).

THE GREAT LAKES ENTOMOLOGIST Vol. 9, No. 1


Michigan buprestid species from the three zones are: 45 species from the Upper (U) Zone: 71 species from the Northern Lower (NL) Zone; and 100 species from the Southern Loner (SL) Zone. The following number of species have been collected only in the zone listed: 5 (U), 0 (NL), and 41 (SL).

The seasonal occurrence of adult buprestid activity within each of the zones is presented in Fig. 3. The SL zone with its more diverse flora and more intense collecting shows a e a t e r number of species than the other two zones. Figure 3 does not indicate those species that overwinter either as free-living adults or within their pupal cells; these may be found earlier or later in the year. For example, adult Taphrocents gracilis Say were collected from the bulrush, Scirpus cyperinus, as late as November 17, 1973, at Okemos, Michigan, when the temperature reached 20.6'C. All Taphrocerus species in Michigan overwinter as free-living adults. The following genera are also known to have some species that overwinter as adults: Acmaeodera, Buprestis, Dicerca, Ptosima, Pachyschelus.

Keys to the genera and species of Michigan adult buprestids follow. The male genitalia are illustrated for Michigan species of Chalcophora, Anthaxia, Pachyschelus and for Agrilus pseudofallax. The complete recorded data are listed only for those species that have two or fewer collection records. Host data are for larval and/or adult collection sites. Chalcophora lacustris LeConte is synonymized with C. virginiensis (Drury), and Pachyschelus confusus, new species, is described.

KEY TO GENERA OF MICHIGAN BUPRESTIDAE

Scutellum absent, elytra fused along median suture. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acmaeodera Eschscholtz (p. 12)

Scutellum present, elytra separate . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Mesal width of metacoxal plates of uniform width or less than lateral width. . 3 Mesa1 width of metacoxal plates greater than lateral width . . . . . . . . . . . . 8 Pronotum truncate at base . . . . . . . . . . . . . . . . . . Ptosima Solier ~(p. 13) Pronotum sinuate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Head and prosternum without antenna1 groove . . . . . . . . . . . . . . . . . . . 5 Head and prosternum with antennal groove . . . . . . . . . . . . . . . . . . . . . 6 Basal metatarsal segment as long as the next three segments together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrilus Curtis (p. 13)

Basal metatarsal segment slightly longer than the next segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eupristocerus Deyrolle (p. 13)

Scutellum small, 116 or less than width of beetle . . . . . . . . . . . . . . . . . . I Scutellum large, 113 width of beetle . . . . . . . . . .I Pachyschelus Solier (p. 17) Body elongate; prosternum without a pit between procoxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taphrocerus Solier (p. 20)

Body ovate; prosternum with a deep pit between procoxae . . . . . . . . . . . . . Brachys Solier (p. 20)

Head with deep median longitudinal groove; specimens usually longer than . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18mm 9

Head without deep median longitudinal groove; specimens usually shorter than 18mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Pronotum grooved medially; apical third of lateral margin of elytra strongly serrate; Host: Hardwoods . . . . . . . . . . . . . Chalcophorella Kerremans (p. 6) Pronotum elevated medially; apical third of lateral margin of elytra entire or finely serrate; Host: Conifers . . . . . . . . . . . . . . . Chalcophora Solier (p. 6) Posternal process acutely angulate laterally posterior to procoxae . . . . . . . .ll Prosternal process obtuse laterally posterior to procoxae. . . . . . . . . . . . . .12 Lateral Iobes of third metatarsal segment about equal in length to the second metatarsal segment . . . . . . . . . . . . . . . . . Chrysobothris Eschscholtz (p. 11) Lateral lobes of third metatarsal segment at least twice length of second metatarsal segrient . . . . . . . . . . . . . . . . . . . . . . Actenodes Dejean (p. 12) Mentum somewhat translucent . . . . . . . . . . . Mehnophila Eschscholtz (p. 9)

6 THE GREAT LAKES ENTOMOLOGlST Vol. 9, No. 1

12'. Mentum dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13 13(12'). Pronotum with a median longitudinal ridge and elytral apices prolonged and

more reddish than the remainder of the elytra. . Pocilontoa Eschscholtz (p. 8) 13'. Pronotum without a median longitudinal ridge and elytral apices prolonged . .14 14(13'). Prothorax truncate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15 14'. Prothorax sinuate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 lS(14). Body broad and short; clypeus broad and short. . . Anthaxia Eschscholtz (p. 9) 15'. Body long and narrow; clypeus long and narrow . . Agrilaxia Kerremans ,(p. 11) 16(14'). Mesosternum and metastemum fused, suture obsolete . . . . . . . . . . . . . . .17 16'. Mesosternum separated from metasternum by a distinct suture. . . . . . . . . . .

Cinyra Laporte and Gory (p. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17(16). Apical antennal segments laterally truncate . . . . . . Buprestis Linnaeus ( p . 8)

. . . . . . . 1 7 Apical antennal segments laterally serrate. Dicerca Eschscholtz (p. 6 )

Subfamily BUPRESTlNAE Tribe BUPRESTlNl

Genus CHALCOPHORA Solier

These species can be collected from decadent or stressed conifers and occasionally from logs or other surfaces in full sun. Based on comparisons of the type of Chalcophora lacustris LeConte and its genitalia, we synonymize Chalcophora lacustris LeConte with Chalcophora virginiensis (Drury).

KEY TO THE SPEClES OF CHALCOPHORA 1. Subsutural groove of elytra interrupted at basal 115 by a raised area; size:

22-32 mm (Male genitalia, Fig. 4A) . . . . . . . . . . . . . . . virginiensis (Drury) 1'. Subsutural groove continuing to elytral base although sometimes shallow. . . . 2 2(lt). Elytral depressions finely and densely punctured; elytra tapering behind the

posterior thud; elytral apex truncate; size: 19-24 mm (Male genitalia, Fig. 4B). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . liberta (Germar)

2'. Elytral depressions densely punctured and very rough; elytra tapering a little behind the middle to apex; elytral apex moderately obtuse; size: 24-32 mm (Male genitalia, Fig. 4C) . . . . . . . . . . . . . . . . . . . . . . . . . fortis LeConte

Genus CHALCOPHORELLA Kerremans A single Michigan species, campestris (Say), recorded by Hubbard and Schwarz (1878)

(specimen not seen) and one specimen labeled Michigan (MSUC). Host: Hardwoods.

Genus DICERCA Eschscholtz

KEY TO THE SPEClES OF DZCERCA (after Nelson, 1975)

. . . . . . . . . . . . . . . 1. Tip of elytron entire, truncate or produced at suture 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . 1'. Tip of elytron strongly bidentate 9

2(1). Second segment of antenna distinctly shorter than third . . . . . . . . . . . . . 3 2'. Second segment of antenna subequal to the third. . . . . . . . . . . . . . . . . . 6 3(2). Mesotibia of male toothed laterally; last sternite of female tridentate, head with

. . . . . . irregular raised area on vertex; Host: Pine, Pinus sp. tenebrosa (Kirby) . . . . . . . . . . . . . . 3'. Mesotibia of male simple; last sternite of female entire. 4

4(3'). Smooth areas of elytra conspicuous; elytra brassy-brown, cupreous or green . . . . . . . . . . . . . . . . . above; Host: Conifers tuberculata (LaPorte & Gory)

Single record: Alcona County, 20-VII-1920, A. W. Andrews (MSUC). 4'. Smooth areas of elytra inconspicuous; dark brown or black or with a cupreous

tinge above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5(4'). Pronotal side margin strongly expanded; elytral tips moderately produced; Host:

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Maple, Acer sp. lugubris LeConte 5'. Pronotal side margin not or weakly expanded; elytral tips weakly produced;

Host: Pine . . . . . . . . . . . . . . . . . . . . . . . . . . . . punctulata (Schonherr)


6(2'j. Median goove of pronotum well developed, punctures of upper surface generally coarse laterally and more rugose . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6'. Median groove of pronotum faintly indicated; punctures of upper surface mdera te Iaterally and less rugose . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7(6). Mesotibia of male simple or with a slight apical dilation; Host: Aspen, Populus sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . callosa Casey

7'. Mesotibia of male with well developed lateral tooth; Host: Aspen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tenebrica (Kirby)

Fig. 4. Male genitalia: dorsal view (left) and ventral view (right). ( A ) Chalcophora virginiensis, ( B ) C. liberta, ( C ) C. fortis, and ( D ) Agrilus pseudofallax.


8(6'). Pronotum widest at middle; elytral striae finely but usually distinctly indicated; lateral lobes of male genitalia rather abruptly tapering to apex; Host: Hardwoods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . divaricata (Say)

8'. Pronotum widest at base and feebly rounded to apex (sometimes subparallel towards base); elytral striae but faintly indicated; lateral lobes of male genitalia tapering gradually to apex; Host: Alder, Alnus sp. . . . . . . . . caudata LeConte

9 1 Elytral tips distinctly prolonged; Host: Witchhazel, Hamamelis virginiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pugionata (Germar)

9'. Elytral tips not or faintly prolonged . . . . . . . . . . . . . . . . . . . . . . . . .10 lO(9'). Raised smooth areas of pronotum and elytra distinctly indicated; Host: Oak,

Quercus sp. . . . . . . . . . . . . . . . . . . . . . . . . asperata (LaPorte and Gory) 10'. Raised smooth areas of pronotum and elytra feebly indicated. . . . . . . . . . . l l ll(lO1). Hind coxal plate notched with tooth on outer side of notch; pronotal margins

narrowed from base to apex; Host: Hardwoods . . . . . . . . obscura (Fabricius) 11'. Hind coxal plate indistinctly notched without tooth; pronotal margins subparallel

to beyond middle, then converging to apex; Host: Hickory, Carya sp. . . . . . . lurida (Fabricius) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Genus POECILONOTA Eschscholtz

KEY TO THE SPECIES OF POECILONOTA 1. Apex of elytra usually reddish and elongate . . . . . . . . , . . . . cyanipes (Say)

This species has been swept from aspen, and reared from the gall of Agrilus criddlei on willow. I t has been associated with the galls of the longhorn beetles, Saperda calcarata Say (Hofer, 1921) and S. concolor LeConte (Knull, 1920) in Populus tremuloides and P. deltoides, respectively.

1'. Apex of elytra broadly truncate and not reddish; recorded from Keweenaw Co. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ferrea (Melsheimer)

Genus CINYRA LaPorte and Gory

A single Michigan species, gracilipes (Melsheimer) has been collected on: oak; hawthorn, Crataegus sp.; and on the flowers of goldenrod, Solidago sp.

Genus BUPRESTIS Linnaeus

KEY TO THE SPECIES OF BUPRESTIS 1. Elytra with yellow or orange markings . . . . . . . . . . . . . . . . . . . . . . . . 2 1'. Elytra without yellow or orange markings . . . . . . . . . . . . . . . . . . . . . . 5 2(1). Elytral color green . . . . . . . . . . . . . . . . . . . . . . . . . . 3

. . . . . . . . . . . . . . . . . . . . . . . . . . 2'. Elytral color brown, never greenish 4 3(2). Elytral apices strongly bidentate . . . . . . . . . . . . . . . . . . . fasciata Fabricius 3'. Elytral apices pointedg Host: Cottonwood, Populus deltoides . . . ronfluenta Say 4(2'). Pronotum evenly convex, unicolorous; Host: Conifers . . . . . meculipennis Gory

. . . . 4'. Pronotum with five longitudinally raised areas, bicolored; Host: Conifers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . consuhrisGory

5(11). Elytra with raised costae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5'. Elytra without raised costae; Host: Pine . . . . . . . . . . . salisburyensis (Herbst) 6(5). Elytra bronze or black; lateral abdominal sternites maculated with orange or

. . . . . . . . . . . . . . . . . . . yellow spots; Host: Conifers rnaculativentris Say 6'. Elytra greenish or bluish; lateral abdominal sternites not maculated with orange

or yellow spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7(6'). Apical half of elytra with dense punctures between costae; elytra usually brilliant

green to blue with sutural and lateral margins cupreous; anterior angles of pronotum not visible from above . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7'. Apical half of elytra with transverse ridges connecting costae; elytra copper- brown, often with greenish or bluish punctures; anterior angles of pronotum

. . . . . . . . . . . . . . . . . . visible from above; Host: Pine sulcicollis (LeConte) Thirtyeight females but no males were collected on bleached driftwood logs on the shore of Lake Superior adjacent to Whitefish Bay in July, 1971.


8(7). E w a l costae strongly convex and with summits nearly impunctate; Western N. America; Host: Conifers. . . . . . . . . . . . . . . . . . . . . . . aurulenta Linnaeus Often emerge indoors, larval life cycle reported as long as 5 1 years; this species imported in wood from the west.

8'. Elytral costae not strongly convex and with summits rather thickly punctate; Host: Conifers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . striata Fabricius

Genus MELANOPHILA Eschscholtz

KEY TO THE SPECIES OF MELANOPHILA

1. Elytra without yellow spots and with acute apices; Host: Conifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . acuminata (DeGeer)

1'. Elytra with or without yellow spots and with broadly rounded and usually serrate apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 Length less than 7 mm; without yellow elytral spots; Host: Pine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aeneola Melsheimer

2'. Length greater than 7 mm; usually with yellow elytral spots . . . . . . . . . . . 3 . . . . 3(2'). Disc of pronotum rough and with rows of large punctures; Host: Conifers

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . fulvoguttata (Harris) 3'. Disc of pronotum striolate; Host: Conifers. . . . drummondi nicolayi Obenberger

Genus ANTHAXIA Eschscholtz

At least two color forms are found i61 each of the following four Michigan species of Anthaxia: cyanella Gory, fisheri Obenberger, quercata (Fabricius), and viridifrons Gory. Since detailed biological studies have not been undertaken, it is unknown whether any of these complexes are represented by more than one species. For example, the thorax and elytra of male cyanella individuals range from a bronzegreen to a green-blue and all of the females are entirely blue; these color forms occur sympatrically. Thus, some males of cyanella and quercicola can easily be confused and should be separated by their genitalic differences. Both sexes of fisheri are represented by a larger dark purple form and a smaller bronze-green form; they often are collected together from American plum, Prunus americana, and have similar genitalia. Anthaxia quercata has two size forms with large males being greener and small males more blue-green. The forms are from widely separated localities but have similar genitalia and both occur on oak. Both sexes of viridifrons are represented by having either bronze or blue elytra forms; they have emerged from American elm, Ulmus americana, and males have similar genitalia.

KEY TO THE SPECIES OF ANTHAXIA . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Tarsal claws with tooth a t base. 2

. . . . . . . . . . . . . . . . . . . . . . . . . . 1'. Tarsal claws without tooth at base. 9 2(1). Elytra purple-black, tapered strongly at apex; Host: American plum (Male

genitalia, Fig. 5A). . . . . . . . . . . . . . . . . . . . . . . . . . . fisheri Obenberger . . . . . . . . . . . . . . 2'. Elytra green, blue or bronze, tapered gradually at apex 3

3(2'). Prothorax uniformly green for males and blue for females; female with frons not . . . . . . . . . entirely blue; Host: Oak (Male genitalia, Fig. 5B) quercicola Wellso

. . . . . . . . . . . . . . . . . . . . . . 3'. Prothorax bronze or more than one color 4 4(3'). Male: frons green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4'. Female: from not green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 5(4). Ventral metathoracic sclerite and abdominal sclerites bronze; Host: American

. . . . . . . . . . . . . . . . . . . plum Hak ,- Fig. 5.4) fisheri Obenberger . . . . . . . . 5'. Ventral metat%oracic sclerite -geen, abdominal sclerites not green 6

. . 6(5'). Apical 113 of el>- unicolorous; Host: on Hickory (Male genitalia, Fig. 5C). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cyanella Gory

. . . . . . . 6'. Apical 113 of elytra bronze and blue-geen (Male genitalia, Fig. 5D). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . quercata (Fabricius)

. . . . . . . . . . . . . . . . . . . . 7(43 Eh'tra bhe: front of head blue cyanella Gory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7'. Eh'tra not b lue . 8

. . . . . . . . . . . 8(7'). Pronotal disc bronze; Host: American plum fisheri Obenberger


8'. Pronotal disc bronze with 2 green areas on either side of the midline and along basal margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . quercata (Fabricius)

9(11). Pronotum with 4 depressions; 2 on either side of midline located centrally on the disc, and 2 near the humeral angIes causing the adjacent margin to appear flat; color dorsally bronze (Male genitalia, Fig. 5E); Host: Conifers; emerged from Black spruce, Picea mariana . . . . . . . . . . . . . . . . . . expansa LeConte

9' Pronotum not as described above . . . . . . . . . . . . . . . . . . . . . . . . . . .10 lO(9'). Lateral pronotal margins each for 114 width of thorax differing from the disc

color; elytra dark blackishpurple; Host: Willow (Male genitalia, Fig. SF)!. . . . . viridicornis (Say) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

10'. Lateral pronotal margins each more broadly pigmented anteriorly than posteriorly and differing in color from disc; elytra uniformly bronze, blue or dark purplz; Host: Elm, Ulmus sp. and Hickory (Male genitalia, Fig. 5G) . . . . . . . .

viridifrons Gory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Fig. 5 . Anthaxia male genitalia: dorsal view (left) and ventral view (right). (A) fisheri, (B) quercicola, (C) cyanella, (D) quercata, ( E ) expansa, ( F ) viridicornis, and ( G ) viridifrons.


Genus AGRILAXIA Kerremans

Three specimens of a single Michigan species, flavimana (Gory), were collected in a mixed hardwoods at Romulus, Wayne County, on 16-VII-1974 by Richard S. Taylor.

Tribe CHRYSOBOTHRINI Genus CHRYSOBOTHRIS Eschscholtz

KEY TO THE SPECIES OF CHR YSOBOTHRIS 1. Lateral margin of last abdominal sternite serrate. . . . . . . . . . . . . . . . . . . 2 1' Lateral margin of last abdominal sternite not serrate; rarely interrupted. . . . .15 2(1). Disc of pronotum uneven, longitudinally sulcate at middle, and frequently with

elevated callosities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2'. Disc of pronotum even, without a distinct median depression or elevated

callosities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 3(2). Clypeus transversely truncate or slightly sinuate in front; Host: Pine . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .floricola Gory 3'. Clypeus emarginate in front. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4(3'). Anterior tibia of male with several small spines on its inner margin; 8th

abdominal tergite of female medially carinate . . . . . . . . . . . . . . . . . . . . 5 4'. Anterior tibia of male with a single apical spine and dilated near the apex on the

inner margin; 8th abdominal tergite of female not medially carinate. . . . . . . 9 5(4). C l j p u s acutely notched at middle but not semicircularly rounded on each side

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5'. Clypeus semicircularly rounded on each side of notched middle (Fig. 6) . . . . 7 6(5). Hale: front of head bright green; female: underside greenish medially; Host:

Hickory. . . . . . . . . . . . . . . . . . . . . . . . . . adelpha Gemminger & Harold 6'. Male: front of head, greenish-bronze; female: underside purplish-bronze; Host:

American plum . . . . . . . . . . . . . . . . . . . . . . . . sloicola Manley & Wellso 7(5'). Antenna gradually narrowed to apex, the last segment not distinctly quadrate

and narrower than the tenth segment . . . . . . . . . . . . . . . . . . . . . . . . . 8 7'. Antenna not narrowed to apex, the last segment quadrate and as wide as the

tenth segment; Host: Hardwoods (Fig. 6). . . . . . . . . . . rugosiceps Melsheimer 8(7). Male antennal segments pale yellow on outer margin; elytral disc and apex

unicolored; posterior pair of foveae separated by a longitudinal costa; Host: Hardwoods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . viridiceps Melsheimer

8'. Wale antenna1 segments bronzy-green and usually reddish-cupreous toward apex; e l y t d disc and apex bicolored with the apex reddish; posterior pair of foveae usually confluent, the bisecting costa interrupted; Host: Hardwoods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . femorata (Olivier)

9(43. Body above green to violaceous blue or purple; Host: Pine . . . . harrisi (Hentz) 9'. Bod:- abave bronze, black or brownish-cupreous . . . . . . . . . . . . . . . . . .10

lO(9'). -+ntennal scgrnents 4 to 11 yellow on outer margin; Host: Pine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . dentipes (Germar)

10'. . b e sqments 4 to 11 not yellow on outer margin . . . . . . . . . . . . . . l l 1 l(103. Prosternum with an anterior median lobe. . . . . . . . . . . . . . . . . . . . . . -12 11'. Pronwum aithout an anterior median lobe . . . . . . . . . . . . . . . . . . . .13 12(113. apem hallowly emarginate in front; length: smaller than 10 mm; Host:

Conifen . . . . . . . . . . . . . . . . . . . . . . . . . . . . pusilla Castelnau & Gory 12'. Qpeas deeply emaregate in front; length: larger than 10 mm; Host: Conifers .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . blanchardi Horn 13(111j. w r speck 1417 mm; Host: Pine. . . . . . . . . . . . . . . . . . or0110 Front 13'. SmaIk=rrpecies:8-13mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14(13'j. Male: posterior tibia arcuate; female: abdomen broadly arcuately emarginate

renually at apex: Host: Conifers. . . . . . . . . . . scabripennis Castelnau & Gory 14'. -Mate: possenor tibia of male straight; female: abdomen triangularly emarginate

mua@- at apex: Host: Conifers. . . . . . . . . . . . . . . . . . . trinervia (Kirby) 15111. EIytra uithout longitudinal costae; Host: Oak. . . . . . . . . concinnula LeConte


Fig. 6. Head of Chrysobothris rugosiceps. Arrow pointing to the semicircularly emarginate clypeus.

One record: Livingston County, 2 mi. N. of Whitmore Lake, 4-VII-1956, G. H. Nelson.

15'. Elytra with costae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 16(1S1). Body above dark bronzy-brown; Host: Hardwoods and Conifers; emerged from

Black spruce. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sexsignata (Say) 16'. Body above violaceous, blue or cupreous; Host: Oak. . . . . . . azurea LeConte 17(2'). Elytra with distinct foveae; Host: Conifers . . . . . . . . pusilla Castelnau & Gory 17'. Elytra without distinct foveae; Host: Pine . . . . . . . . . . . . . . harrisi (Hentz)

Genus ACTENODES Dejean

KEY TO THE SPECIES OF ACTENODES 1. Fourth antennal segment strongly triangular, at widest part nearly twice as wide

as third, and the following segments transverse; Host: Maple . . . . acornis (Say) 1'. Fourth antennal segment slightly wider than third, the following segments not

transverse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . simi Fisher Two records: Berrien County, 4-VII (UMMZ); Manistee County, 5-VII-1952, R. Dreisbach (MSUC).

Subfamily ACMAEODERINAE Tribe ACMAEODERINI

Genus ACMAEODERA Eschscholtz

KEY TO THE SPECIES OF ACMAEODERA

1. Elytra dark with isolated spots; last ventral sternite without subapical plate, procoxae with branched setae (Fig. 7); Host: Hardwoods . . . htbulus (Fabricius)


Fig. 7. Branched setae on fore coxa of Acmaeodera tubulus.

Frequently found feeding on the petals of wild geranium, Geranium maculatum, and occasionally found ovipositing on red oak, Quercus borealis.

1'. Elytra with elongate yellow markings; last ventral sternite with a subapical plate; pronotum usually with a lateral yellow spot; prowxae without branched setac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pulchella (Herbst)

Genus PTOSIMA Solier

A dark blue and yellow species, gibbicollis (Say), inhabits redbud, Cercis canadensis, in southern Michigan.

Subfamily AGRILINAE Tribe AGRILINI

Genus EUPRISTOCERUS Deyrolle

A single robust species, cogitani (Weber), with a copper colored thorax and black elytra with faint white pubescent markings, is often collected on speckled alder, Alnus rugosa.

Genus AGRILUS Curtis

KEY TO THE SPECIES OF AGRILUS 1. Elytral apices prolonged, terminating in a large spine. . . . . . . . . . ferrisi Dury

One record: Three pupae collected from hackberry, Celtis occidentalis, from Ingham County, Okemos, 15-VI-1972

1'. EIytral apices not prolonged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2


2(1'). Antennae serrate beginning with the fourth segment . . . . . . . . . . . . . . . . 3 2'. Antennae serrate beginning with the fifth segment . . . . . . . . . . . . . . . . .37 3(2). Tarsal claws bifid with the inner tips nearly touching . . . . . . . . . . . . . . . 4 3'. Tarsal claws bifid with the inner tips widely separated. . . . . . . . . . . . . . .18 4(3). Pygidium with a projecting carina . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4'. Pygidium without a projecting carina . . . . . . . . . . . . . . . . . . . . . . . . . 6 5(4). Body above uniformly black; Host: Hop hornbeam. . . . . . . . champlaini Frost

Most easily collected by rearing adults from hop hornbeam galls. 5'. Head and thorax cupreous (occasionally with a bluish tinge), elytra black; Host:

Stem galls on blackberry, raspberry, and dewberry, Rubus sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ruficollis (Fabricius)

6(4'). Abdominal segments with pubescent spots laterally; Host: Honey locust, Gleditsia triacanthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . difficilis Gory One record: Port Huron (Hubbard and Schwarz, 1878). No specimens seen.

6'. Abdominal segments without pubescent spots laterally. . . . . . . . . . . . . . . 7 7(6'). Male: Metatibi.de without a distinct apical spine on the inner margin . . . . . . 8 7'. Male: Metatibiae with a distinct apical spine on the inner margin . . . . . . . . 9 8(7). Metatarsi shorter than tibiae; segment one as long as the combined length of the

next three segments; Host: Hardwoods . . . . . . . . . . . . . . . . armatus (Say) 8'. Metatarsi as long or longer than the tibiae; segment one as long as next four

segments; Host: Boxelder, Acer negundo, and hop hornbeam. . masculinus Horn 9(7'). Antennal segments with long white setae beneath; Host: Oak. . crinicornis Horn 9'. Antenna] segments without long pubcscence beneath . . . . . . . . . . . . . . .10

lO(9'). Male: last ventral abdominal segment fimbriate at apex; Host: Oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . defectus LeConte

10'. Male: last ventral abdominal segment not fimbriate at apex . . . . . . . . . . . .ll 1 l.(lOf).Male: prosternum conspicuously pubescent. . . . . . . . . . . . . . . . . . . . . .12 11'. Male: prosternum not conspicuously pubescent . . . . . . . . . . . . . . . . . . .15 12(11). Prosternum deeply emarginate; Host: Black walnut, Juglans nigra (reared

specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cliftoni Knull 12'. Prosternum truncate or slightly emarginate . . . . . . . . . . . . . . . . . . . . . .13 13(12'). Prehumeral carina of pronotum very indistinct; male genitalia with parameres

parallel; Host: Oak . . . . . . . . . . . . . . . . . . . . . . . . . . . geminatus (Say) 13'. Prehumeral carina of pronotum very distinct; male genitalia with parameres

expanded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 14(13'). Frons with dense pubescence nearly concealing the surface behind the epistoma;

pronotum bicolored; Host: Hickory . . . . . . . . . . . . . . . . . . . . otiosus Say 14'. Frons with pubescence but not concealing the surface of the epistoma;

pronotum unicolored dorsally; Host: Hazelnut, Corylus americana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atricornis Fisher

15(111). Male: second abdominal segment transversely concave; Host: Black walnut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . transimpressus Fall

15'. Male: second abdominal segment not transversely concave . . . . . . . . . . . .16 16(15'). Male genitalia with protruding flcshy lobes at apex of parameres; frons usually

bluish; Host: Hop hornbeam . . . . . . . . . . . . . . . . . . . . . . . osburni Knull 16'. Male genitalia without protruding fleshy lobes at apex of parameres; frons bluish

or greenish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17 17(16'). Male genitalia expanded near middle and parameres with tips parallel; Host:

. . . . . . . . . . . . . . . . . . . . . . . Butternut, Juglans cinerea juglandis Knull 17'. Male genitalia expanded near middle and parameres arced to apex; Host: Oak. .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . frosti Knull . . . . . . . . . . . . . . . . . . . . . . . . . . 18(3'). Pygidium with a projecting carina .19

18'. Pygidium without a projecting carina . . . . . . . . . . . . . . . . . . . . . . . . .30 19(18). Pronotum with a densely pubescent median longitudinal groove; Host: Haw-

. . . . . . . . . . thorn; Apple, Malus pumila; and Serviceberry, Amelanchier sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vittaticollis (Randall)

. . . . . . 19'. Pronotum without a densely pubescent median longitudinal groove .20


20(19'). Pronotum without, or with only feebly indicated, prehumeral carinae. . . . . .21 20'. Pronotum with distinct prehumeral carinae. . . . . . . . . . . . . . . . . . . . . .25 21(20). Elytra with pubescent spots or pubescent vittae. . . . . . . . . . . . . . . . . . .22 21'. Elytra without pubescent spots or pubescent vittae. . . . . . . . . . . . . . . . .23 22(21). Elytra black with a distinct whitish or yellowish pubescent vitta on each elytron;

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Host: Oak bilineatus (Weber) 22'. Elytra blue or bluish-black with an indistinct pubescent vitta in the basal

. . . . depressions of each elytron; Host: Hop hornbeam bilineatus carpini Knull 23(211). Vertical aspect of the abdominal segments densely pubescent (except sometimes

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . the second) .24 23'. Vertical aspect of abdominal segments not conspicuously pubescent; Host:

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Willow criddlei Frost This species forms distinct galls (Fig. 8) that are often split open revealing the underlying cambium.

24(23). Elytra blue or blue-black; vertical surface of second abdominal segment glabrous, all others segment with white pubescence; Host: American hazel, Hop hornbeam, Oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . acutipennisMannerheim

Fig. 8. Willow stem gall of Agrilus criddlei showing typical fissures in bark exposing the underlying cambium.


24'. Elytra olivaceous bronze to black; vertical surfaces of all abdominal segments with white or gold pubescence; Host: Oak . . . . . . . . . quadriimpressus Ziegler

25(201). Vertical surface of abdominal segments (except sometimes the second) conspicuously pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26

25'. Vertical surface of all abdominal segments uniformly, but not conspicuously pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27

26(25). Frons densely pubescent, the pubescence nearly obscuring the surface; elytral apex broadly rounded; Host: Lombardy poplar, Populus nigra L. var. italica . .

granulatus (Say) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26'. Frons not densely pubescent, the surface distinctly visible; elytral apex acute or

narrowly rounded; Host: Willow . . . . . . . . . . . . . . . . . quadriguttatus Gory 27(25')L Lateral margins of pronotum with a reddish tinge; elytra greenish-bronze to

reddish-cupreous with a faint pubescent spot at apical third, often running . . . . . indistinctly along the sutural margin to apex; Host: Alder. pensus Horn

The color of 45 Michigan specimens reared from alder was more variable than Fisher (1928) noted; also, Michigan specimens as reported previously by Carlson (1969) typically have an indistinct apical elytral spot.

27'. Above olivaceous black or'blue-gray to bronze . . . . . . . . . . . . . . . . . . .28 28(27'). Vertical surface of abdominal segments obscured :by an even distribution of

pubescence; Host: Shoots of Quaking aspen, Populus tremuloides, or Big tooth aspen, P. grandidentata (Nord et al., 197 I)\. . . . . . . . . . . . . horni Kerremans

28'. Vertical surface of abdominal segments with some pubescence but not obscuring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . thesurfa ce 29

29(28'). Elytra without pubescent spots; male genitalia ventrally with the bevel of the inner margin of each pararnere wide; female with projecting pygidial spine about equal in length and width, elliptical in cross section at apex; Host: Birch, Betula sp. . . . . . . . . . . . . . . . . ; . . . . . . . . . . . . . . . . . . . . . anxius Gory

29'. Elytra often with a pair of indistinct spots at basal third; male genitalia ventrally with the bevel of the inner margin of each paramere narrow; female with projecting pygidial spine longer than wide, rounded in cross section at apex; Host: Aspen . . . . . . . . . . . . . . . . . . . . . . . . . . liragus Barter & Brown

30(18'). Elytra with pubescent spots or lines. . . . . . . . . . . . . . . . . . . . . . . . . .31 30'. Elytra without pubescent spots or lines. . . . . . . . . . . . . . . . . . . . . . . .32 31(30). Prosternal lobe deeply emarginate in front; Host: Hardwoods. . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . obsoletoguttatus Gory 31'. Prosternal lobe not deeply emarginate in front; Host: Honey locust and

Hackberry . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . fallax Say 32(30r). Outer antennal segments (7-11) distinctly wider than long . . . . . . . . . . . .33 32'. Outer antennal segments (7-11) not distinctly wider than long . . . . . . . . . .34 33(32). Prosternal lobe broadly rounded in front; Host: Stem galls on Raspberry, Rubus

sp. and Rose, Rosa sp. . . . . . . . . . . . . . . . . . . . . . . . . rubicola Abeille 33'. Prosternal lobe emarginate in front; Host: Willow . . . . . . . . . . politus (Say) 34(32'). Dorsally dark blue to greenish-blue; prosternal lobe deeply emarginate in front;

. . . . . . . . . . Host: Tartarian honeysuckle, Lonicera tartarica coeruleus (Rossi) 34'. Dorsally not blueish; prosternal lobe shallowly emarginate in front, elytra bronze

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . or purple-reddish .35 35(34'). Elytra blackish-bronze to olivaceous; setae on lower fourth of face more dense,

obscuring surface; Host: Dogwood, Cornus sp. . . . . . . . . . cephalicusLeConte 35'. Elytra usually reddish at apex or with purple tinge; setae on lower fourth of face

not obscuring surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36 36(35'). Body above unicolored; pronotum and elytra the same color; Host: Hawthorn. .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crataegei Frost 36'. Body above bicolored; pronotum cupreous and elytra purplish or piceous; Host:

Seniceberry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . amelanchieri Knull 37(2'). Elytra with pubescent spots, vittae, or irregular designs . . . . . . . . . . . . . .38

. . . . . . . . . . . 37'. Elytra sometimes uniformly pubescent but never patterned .41 . . . . . . . . . . . . . . . . . . . . . . . . . 38(37). Pronotum with prehumeral carinae .39


. . . . . . . . 38'. Pronotum without prehumeral carinae; Host: Privet, Ligustmm sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subcinctus Gory

39(38). Elytra with pubescent spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 39'. Elytra with irregular pubescent designs; Host: Hackberry. . . . lecontei Saunders 40(39). Elytral pubescent spots distinct, medial spots rounded; [Male genitalia refigured

. . . . . . . from Fisher (1928) (Fig. 4D)] ; Host: Honey locust pseudofallax Frost 40'. Elytral pubescent spots indistinct, medial spots elongate; Host: Honey locust . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . egeniformis Champlain & Knull 41(37?. Pronoturn with prehumeral carinae . . . . . . . . . . . . . . . . . . . . . . . . . .42 41'. Pronotum without prehumeral carinae; Host: Maple . . . . . . . . . . putillus Say 42(41). Surface above bicolored; Host: False indigo, Amorpha fruticosa . . pusillus (Say) 42'. Surface above unicolored . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . i43 43(42'). Elytra uniformly sparsely clothed with distinct white hairs; prostemum deeply

emarginate in front; Host: Hackberry . . . . . . . . olentangyi Champlain & Knull One record: Lenawee County, Morenci, 13-VI-1974, S. G. Wellso

43'. Elytra not uniformly cIothed with distinct hairs, although an indistinct pubescent vitta may be present on each elytron; prosternal lobe truncate or slightly emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .44

44(43'). Each elytron with a very indistinct often interrupted sutural vitta; apex of aedeagus blunt; Host: Hackberry. . . . . . . . . . . . . . . . . . . . paracelti Knull

44'. Elytra with distinct pubescence only in humeral depression . . . . . . . . . . . .45 . . . 45(44'). Male genitalia with iides of the parameres nearly parallel; Host: Hackberry.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . celtiKnull 45'. Male genitalia with sides of the parameres very strongly, arcuately expanded;

Host: Black locust, Robinia pseudoacacia . . . . . . . . . . . . . . . egenus Gory

Genus PACHYSCHELUS Solier

KEY TO THE SPECIES OF PACHYSCHELUS 1. Elytra purple with white pubescent markings; male genitalia (Fig. 9A); Host:

Wild geranium purpureus (Say) . . . . . . . . . . . . . . . . . . 1'. Elytra black without white pubescent markings 2

2(11). Male genitalia (Fig. 9B); Host: Desmodium sp. . . . . . . . . . . . . laevigatus Say 2'. Male genitalia (Fig. 9C, D); Host: Lespedeza sp.. . . . . . . . . . . . confums n.sp.

Pachyschelus confusus Wellso and Manley, NEW SPECIES

HOLOTYPE MALE: Form and size of P. laevigatus (Say); ovate; color uniformly shining black.

Head convex, slightly bronze, with longitudinal depression in front; surface with a few scattered punctures and setae; surface between punctures finely granulose.

Pronotum widest at base, narrowest at apex, sides arcuate from the apex to base, hind angles slightly produced; surface with scattered punctures, each puncture bearing a single recumbent seta; surface between punctures finely granulose; scutellum trapezoid, irn- punctate, and glabrous.

Elytra almost as broad as pronotum, widest at base, narrowest at apex, sides arcuate from apex to base, tips rounded and entire lateral apical margins serrulate; disk convex, each elytron with a lateral excavation behind umbone; surface with punctures, each puncture bearing a single recumbent white seta (Fig. 10A).

Ventral surface (Fig. 10B) concolorous with dorsal; underside of thorax with scattered punctures, medially with scattered, moderately long white setae; hind tibia with a concave area about midway on outer margin containing a row of stout setae (Fig. 10C); abdomen minutely granulate and bearing scattered short white setae; last abdominal sternite with scattered small tubercles, setae denser and longer than on other abdominal segments, posteriorly obtusely rounded with a small spine at apex. Size: length 2.13 mm; width, 1.27 mm.

ALLOTYPE FEMALE: Differs from the male by having the last abdominal sternite depressed, with two sets of four teeth on either side of the apex (Fig. 10D). Size: length, 2.19 mm; width, 1.48 mm.




TYPES: Holotype male: Michigan: Shiawassee County, T5N R1E Sec. 21, 21-VIII-1974, G. V. Manley, sweeping bush clover, Lespedeza capitata Michx. Allotype female: Same collection data as holotype. Both types deposited in MSUC. Paratypes: 460 males and 380 females from type locality in late August; 13 males and 7 females collected by Daniel Young at the type locality on 4-V1-1975; and 25 males and 16 females from Oakland County, T2N R8E Sec. 21, 8-W-1974, G. V. Manley. Paratypes deposited in authors' collections, MSUC, UMMZ, UWEM, JNKC, GHNC, DYCC, USNM, MCZC, AMNH, CASC, CHAH [hbbreviations listed in Arnett and Samuelson (196911.

VARIATION: No variation noted other than the range in size. Males: length, 1.92-2.33 mm; width, 1.14-1.31 mm. Females: length, 2.13-2.51 mm; width, 1.31-1.49 mm.

COMPARISON: This species is very similar to laevigatus Say and is undoubtedly placed with it in collections. The two species can be separated by the male genitalia (Fig. 9B, C). In Michigan, confusus is associated with Lespedeza sp. (specifically capitata), and laevigatus with Desmodium sp. (specifically canadense).

Genus BRACHYS Solier

KEY TO THE SPECIES OF BRACHYS 1. Last ventral segment with long hairs along the emargination; usually larger than

5.5 mm; Host Oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . ovatus (Weber) Apparently parthenogenic in Michigan, since no males were represented in over 100 specimens examined.

1'. Last ventral segment without long hairs along the emargination; usually smaller than 5.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 1 ' ) Elytra with a purple, blue, or green luster especially in the humeral region; apical elytral setae predominately gold; length, 3-5.75 mm; Host: Hardwoods, usually oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aerosa Melsheimer

2'. Elytra dark brassy; apical elytral setae light gold to silver; length, 3-3.75 mm; Host: Oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aenrginosus Gory

Genus TAPHROCERUS Solier All of these, ;species have been collected on bulrushes.

KEY TO THE SPECIES OF TAPHROCERUS

1. Elytra with pubescent markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1'. Elytra without pubescent markings . . . . . . . . . . . schaefferi Nicolay & Weiss 2(1). Smaller species: less than 3 mm in length. . . . . . . . . . . . nicolayi Obenberger 2'. Larger species: usually greater than 3 mm in length. . . . . . . . . . . gracilis Say

We speculate that the forward pointing thoracic sendla may be used to give the beetle the information to ensure that it does not become so tightly lodged between the bases of two adjacent bulrush blades that its legs could not be extended (Fig. 11).

ADDITIONAL SPECIES AND STUDIES

Certain species, although not available during the preparation of this paper, probably occur in Michigan. These include: Acmaeodera ornatoides Barr, Chrysobothris chryseola Illiger, C. verdigripennis Frost, Poecilinota thureura Say, and Ptosima walshii LeConte.

Additional studies are warranted on Anthaxia sp., as well as on the Agrilus anxius, A. acutipennis, A. arcuatus, A. politus, and Chrysobothris femorata complexes. Biological studies are necessary to better define the species limits, and separate or combine subspecies and color morphs.

ACKNOWLEDGMENTS

We thank Dr. Roland L. Fisher for his counsel during the course of this work, and Marian Mahler for her help with the tables and illustrating the genitalia. We appreciate


Fig. 11. Taphrocems gracilis: (A) head and pronoturn, and (B) pronotal "belt buckle" sensory organs whose setae point anterad.

receiving the loan of holotypes of Ckalcophora lacustris LeConte and Agrilus pseudofaltax Frost from Janice C. Scott a t the MCZC, and the loan of specimens from Dr. Irving Cantrall at the UMMZ. We express our appreciation to William S. McAfee and Arthur Ackennan for their technical assistance in obtaining the scanning electron micrographs.

LITERATURE CITED

Andrews, A. W. 1921. The Coleoptera of the Shiras Expedition to Whitefish Point, Chippewa County, Michigan. Pap. Mich Acad. Sci. Arts. Lett. 1:293-390.

Arnett, R. H, and G. A. Samuelson. 1969. Directory of Coleoptera Collections of North America (Canada through Panama). Cushing-Malloy, Inc. Ann Arbor, Michigan. 123 p.

Barr, W. F. 1971. Family Buprestidae, p. 55-89. In M. H. Hatch, The beetles of the Pacific Northwest Univ. of Washington Press, Seattle and London.

Carlson, R. W., and F. B. Knight. 1969. Biology, taxonomy, and evolution of four sympatric Agrilus beetles (Coleoptera: Buprestidae). Contrib. Am. EntomoI. Inst. (Ann Arbor) 4: 1-105.


Fisher, W. S. 1928. A revision of the North American species of buprestid beetles belonging to the genus Agrilus. U.S. Natl. Mus. Bull. 145. 347 p.

Franklin, R. T., and H. 0. Lund. 1956. The Buprestidae (Coleoptera) of Georgia. Ga. Agric. Expt. Stn. Tech. Bull. (N.S.) 3,48 p.

Hofer, G. 1921. The aspen borer and how to control it. U.S. Dept. Agric. Farmers Bull. 1134, 9p.

Hubbard, H. G., and E. A. Schwarz. 1878. The Coleoptera of Michigan. Proc. Am. Philos. SOC. 17:593-666.

KnuU, I. N. 1920. Notes on Buprestidae with descriptions of new species. Entomol. News 21:s.

KnuU, I. N. 1925. Buprestidae of Pennsylvania. Ohio State Univ. Stud. 2:l-71. Nelson, G. H. 1975. A revision of the genus Dicerca in North America (Coleoptera:

Buprestidae). Entomol. Arb. Mus. Frey, Munich, W. Germany. 26:87-180. Nord, I. C., F. B. Knight, and G. B. Vogt. 1971. Identity and biology of an aspen root

girdler, Agrilus horni. For. Sci. 11: 33-41. Townsend, T. 1889. Contribution to a list of the Coleoptera of the Lower Peninsula of

Michigan. Psyche 5: 231-235.

INDIANA ENSIFERA (ORTHOPERA)

and

J . L. Stein

Department of Entomology Purdue University

West Lafayette, Indiana 47907


INDIANA ENSIFERA (ORTHOPERA)'

W. P. McCafferty and J . L. Stein2

A total of 67 species of long-horned grasshoppers and crickets were reported to occur in Indiana by Blatchley (1903) in his "Orthoptera of Indiana." Distributional information concerning thek species was sparse and has not been significantly supplemented since that time. Subsequent works which have dealt either heavily or exclusively with the Indiana fauna include Fox (1915), Blatchley (1920), Cantrall and Young (1954), and Young and Cantrall(1956).

The study reported herein was instigated in order to synthesize and augment our knowledge of the ensiferan fauna of the state of Indiana, including a complete and current record of the geographic, temporal, and ecological distribution of the species. The results, moreover, incorporate considerable taxonomic changes since Blatchley's time, substantiate 10 new state records increasing the known number of state species to 83, implicate an additional eight species as possibly occurring in Indiana, and establish 400 new county records for 61 of the 83 Indiana species. Identification tables for all of these species were presented by Stein and McCafferty (1975).

The data were derived primarily from the study of over 3,800 museum specimens housed at Purdue University's Laboratory of Insect Diversity, the University of Michigan Museum of Zoology, and the Illinois Natural History Survey. Specimens were sexed and identitied or verified, and the label data were recorded.

The style of presentation used is similar to that of Cantrall (1968). The annotated checklist includes for each species, and subspecies when appropriate, the complete taxonomic name, the common name, and brief statements concerning the seasonality of adults and ecological associations. Species verified by us as new state records are marked with an asterisk. Geographic distribution of each species in the form of plotted counties is indicated on individual Indiana maps which accompany each specific treatment. Open circles signify that there are previously existent county records from the literature. Solid circles signify that specimens taken from the county have been studied by the authors and are present in one of the collections mentioned above. A solid circle, alone, would therefore indicate a new county record. Readers are referred to Figure 1 for a complete labelling of county names.

For the purpose of attempting to draw correlations between specific distributions and natural features of the environment, the readers are referred to Lindsey (1966). This work dealt in depth for Indiana with such things as the historical geology, bio~limate, vegetation and floral areas, soil types, speleology, and mammalian distribution, all of which may possibly have either a direct or indirect bearing on the distribution of the ensiferan fauna In addition, Chandler (1966) divided the state into regions and biotic zones based on certain elements of the insect fauna including the Hymenoptera, Coleoptera, Odonata, and Orthoptera in part. The "zones" are strongly correlated with floral areas and the Pleistocene geology of Indiana. A striking difference in fauna is sometimes apparent between a southern unglaciated area of Indiana and the remaining glaciated area of the state.

Blatchley (1903) had earlier estimated life zones in the state based primarily on the distribution of Orthoptera. Modifying Merriam's (1898) concepts, Blatchley concluded that the "Transition Zone" (Alleghanian fauna) overlapped the northern fourth of the state, the "Lower Austral Zone" (Austroriparian fauna) covered the greater part of the southern third of the state, and the "Upper Austral Zone" (Carolinian fauna) covered essentially the entire state and mingled with the latter two faunas. Our data indicates that the number of species of Ensifera in the state that might be considered Alleghanian or

l~ublished with the approval of the Director of the Purdue Agricultural Experiment Station as Journal Series No. 5910.

2~epar tmen t of Entomology, Purdue University, West Lafayette, Indiana 47907.


Fig. I . The counties of the State of Indiana.


Austroriparian is much smaller than Blatchley originally thought, due to many factors such as synonymy, original misidentification of species, and our much expanded distributional data Nevertheless, there remains evidence that much of the current southern ensiferan fauna in Indiana has been derived from a previously more widespread preglacial fauna. Also, on the basis of distributional data, supposed northern extensions of the predominantly southern species are associated in most cases with large river valleys such as the Wabash and White. It could, furthermore, be inferred from this, that either postglacial dispersal has been more general but evidence of such obliterated by recent agricultural development, or that recent dispersal has indeed been via river valleys.

The biotic units of Indiana as proposed by Chandler (1966) do not appear to be applicable to the Ensifera because of either considerable overlap in species' distribution among zones or a wide disparity in collecting records. Hopefully, the raw distributional data concerning Indiana Ensifera are now complete enough for those who would wish to undertake a more thorough examination of the geographic affinities of each species. We would caution, however, that the total range of any species along with all available information concerning evolutionary origins are additional critical considerations in any comprehensive study of zoogeography.

Hebard (1934) and Cantrall (1968) published faunal accounts of Orthoptera species for the neighboring states of Illinois and Michigan respectively. These and various other treatments containing collecting records of Ensifera from areas in proximity to Indiana in adjacent states have indicated that eight species and one subspecies may eventually be found in Indiana A list of these follows with the adjacent state distribution indicated parenthetically for each:

Scudderia fasciata Beutenmiiller (Michigan and Ohio) Diastremmena apicalis Brunner (Adventive: Illinois) Ceuthophilus pallidipes E. M. Walker (Ohio and Michigan) Ceuthophilus silvestris Bruner (Michigan) Ceuthophilus guttulosus nigricans Scudder (Kentucky) Gryllus rubens Scudder (Illinois) Oecanthus celerinicfus T. J . Walker (Illinois and Kentucky) Oecanthus laricis T. J . Walker (Michigan and Ohio) Cyrtoxipha columbiana Caudell (Illinois, Kentucky, and Ohio)

Family TETTIGONIIDAE (The Katydids)

Subfamily PHANEROPTERINAE (Bush and Roundheaded Katydids)

Genus SCUDDERIA Stal (Bush Katydids)

*septentrionalis (Serville) 1839. Northern Bush Katydid (Fig. 2). Records for the state are from August. The species is presumed to be much more widespread than is indicated but is rare in collections. It has been taken in deciduous wooded areas.

pistillata (Brumer) 1878. Broadwinged Bush Katydid (Fig. 3). Adults have been taken from late sumnier to early fall in the lakes region of northern Indiana. It has been taken primarily in low bushes and other plants associated with ponds and bogs.

cwicauda curvicauda (DeGeer) 1773. Curvetailed Bush Katydid (Fig. 4). Adults have been taken in late summer and early fall. Populations are present throughout Indiana and are usually found in association with marshy habitats.

texensis Saussure and Pictet 1897. Texas Bush Katydid (Fig. 5). Adults of this rather widespread species have been collected in late summer and early fall. It is known from marshes, swamps, bogs, and low marginal vegetation.


Scudderia septentr ionaZis

Scudderia curvicauda curvicauda

Scudderia p i s t i ZZata

Scudderia t exens i s


Scudderia furcata furcata

furcata furcata Brunner 1878. Forktailed Bush Katydid (Fig. 6). Adults are known from late summer to early fall. This bush katydid is known more from marginal bushes and trees in the hilly areas of southern Indiana and from marshy to prairie habitats in the more northern areas.

Genus INSCUDDERIA Caudell

taxodii Caudell 1921. Cypress Katydid (Fig. 7). This species has been taken in ea11y October in the state and is known only from the extreme southwestern comer. This species is associated with cypress swamps and so occurs only in this restricted ecological area of Indiana

Genus AMBLYCORYPHA Stal (The Roundheaded Katydids)

oblongifolia (DeGeer) 1773. Oblongwinged Katydid (Fig. 8). Adults have been taken from late summer to early fall. This species should eventually be known from throughout the state. Its habitat ranges from marshy areas to marginal shrubbery and weedy fields

uhleri Stal 1876. Uhler's Katydid (Fig. 9). Adults of this species are present in late summer and early fall. I t has been taken only in the southern half of the state, associated with sedges and willows in the Wabash River Valley and grasses and oaks in other areas.

rotundifolia rotundifolia (Scudder) 1862. Roundwinged Katydid (Fig. 10). Adults have been taken in late summer and early fall throughout the state. It is commonly found on or close to the ground among grasses and weedy vegetation in open woodlands or damp ravines.

3 0 THE GREAT LAKES ENTOMOLOGIST Vol. 9, No. 1

Amb Zycorypha ob Zongifo Zia

Amb Zycorypha rotundifoZia rotundifoZia

Amb Zycorypha uh ter i

Microcentrum rhombifoZium

1976 THE GREAT LAKES ENTOMOLOGIST 3 1

Microcentrum ret inerve Pterophy l l a came ZZifo Zia camel l i fo l ia

Genus MICROCENTRUM Scudder (Bush Katydids)

rhombifolium (Saussure) 1859. Broadwinged Katydid (Fig. 11). Adults are known from late summer to mid fall. It evidently is a more central and southern species in Indiana, never having been taken in the northern counties. It is found associated with bushes and deciduous trees.

retinerve (~urmeister) 1838. Angularwinged Katydid (Fig. 12). Adults are present from late summer to early fall. The species is known more from the southern counties in Indiana where it has been taken on the ground or in low branches of shrubbery.

Subfamily PSEUDOPHYLLINAE (The True Katydids)

Genus PTEROPHYLLA Kirby

camellifolia camellifolia (Fabricius) 1775. Northern True Katydid (Fig. 13). This common and widespread katydid is known from mid summer to early fall. It is found in the crowns of deciduous trees.

Subfamily COPIPHORINAE (The Coneheaded Katydids)

Genus NEOCONOCEPHALUS Karny (I'he Coneheaded Katydids)

exiliscanorus (Davis) 1887. Slightly Musical Conehead (Fig. 14). Adults have been taken in late summer and early fall. In Indiana it is known only from heavy thickets and grasses along the Ohio River.


nebrascensis (Brunner) 1891. Nebraska Conehead (Fig. 15). This species has been taken in late summer and early fall. It most likely will eventually be taken throughout most of the state, and has been found in tall grassy habitats.

*ensiger (Harris) 1841. Swordbearing Conehead (Fig. 16). Adults have been taken from mid summer to early fall. This species is not known from southern Indiana. It is evidently associated with open fields and grasslands.

robustus (Scudder) 1862. Robust Conehead (Fig. 17). Adults are present from late summer to early fall. This species has been taken in grassy areas near water and appears to be more common to the river valleys of west central Indiana. In the north it is found in secondary field growth.

*bivocatus Walker, Witesell, and Alexander 1973. Twovoiced Conehead (Fig. 18). Adults have been taken in October in sandy, grassy areas of the dunes area of extreme northwestern Indiana.

palustris (Blatchley) 1893. Marsh Conehead (Fig. 19). Adults are present from late summer to early fall. The species is locally abundant in marshy areas and pond margins in west central Indiana.

retusus (Scudder) 1879. Roundtipped Conehead (Fig. 20). Adults have been taken in late summer and early fall in marshy to more xeric fields throughout the southern half of Indiana.

Subfamily CONOCEPHALINAE (The Meadow Katydids)

Genus ORCHELIMUM Sewille

dvaticum McNeill 1891. Longspuned Meadow Katydid (Fig. 21). Adults have been taken in Iate summer and early fall. This meadow katydid appears to be primarily a

NeoconocephaZus exCZiscmzoms NeoconocephaZus nebrascensis


NeoconocephaZus ensiger

NeocomcephaZus bivocatus NeoconocephaZus paZustris


NeoconocephaZus retusus Orche Z i m m siZvaticum

southern Indiana species since more northern records in the state cannot be confirmed. It is found in open deciduous woodlands on all types of vegetation.

vulgare Harris 1841. Common Meadow Katydid (Fig. 22). A widespread species taken throughout the state from late summer to early fall. It is common in upland habitats along fence rows and in fields of grass, clover, or mixed weeds.

gladiator (Bruner) 1891. Gladiator Meadow Katydid (Fig. 23). Adults are present in late summer and early fall. This species occurs only in northern Indiana where it has been taken in association with marshy and grassy meadows.

bullatum Rehn and Hebard 1915. Stout Meadow Katydid (Fig. 24). Adults occur in late summer and early fall. Specimens have been taken in isolated marshes in Tippecanoe County.

nigripes Scudder 1875. Blacklegged Meadow Katydid (Fig. 25). Adults are present in late summer and early fall. The species is common throughout Indiana in tall grasses and smartweed surrounding lakes, ponds, and marshes.

concinnum Scudder 1862. Stripefaced Meadow Katydid (Fig. 26). This species is found from late summer to early fall and is apparently restricted mostly to marsh or bog areas of the northern part of the state.

campestre Blatchley 1893. Duskyfaced' Meadow Katydid (Fig. 27). This species is present from mid summer to early fall. Verified records are from marshy areas of northern Indiana. It reportedly has also been taken in areas of the Wabash and White River valleys.

deIicatum Bruner 1892. Delicate Meadow Katydid (Fig..28). Adults are present from late summer to early fall. This species is known only from the northern part of the state in the Kankakee Sand Ridge area.


Orche limum vulgare

Orche limum b u l l a t m Orche limum nigripes


Orche lim concinnum

Orchelimum delicatwn

O r c h e l i m campes t r e

Orchelimwn volantwn


volantum McNeill 1891. Nimble Meadow Katydid (Fig. 29). Adults are present from late summer to early fall. The species frequents emergent vegetation along ponds and streams.

Genus CONOCEPHALUS Thunberg

fasciatus fasciatus (DeGeer) 1773. Slender Meadow Katydid (Fig. 30). Adults of this common species are present from mid summer to mid fall. Throughout Indiana it may be found in timothy and clover pastures and bluegrass meadows, especially around water and in low areas.

brevipennis (Scudder) 1862. Shortwinged Meadow Katydid (Fig. 31). This species is common from late summer to mid fall. I t is taken throughout Indiana in habitats similar to that of C. fasciatus.

nemoralis (Scudder) 1875. Woodland Meadow Katydid (Fig. 32). Adults are present from late summer to early fall. This species has been taken throughout the state but is somewhat uncommon in the northern part. It inhabits roadside vegetation, fence row and marginal wooded areas.

strictus (Scudder) 1875. Straightlanced Meadow Katydid (Fig. 33). Adults are present from late summer to early fall throughout Indiana. I t is taken primarily in open field situations

nigropleurum mruner) 1891. Blacksided Meadow Katydid (Fig. 34). Adults are present from mid summer to early fall. This species can be found over most of the state, but records are sparse for southern Indiana. I t inhabits grasses and sedges in areas in proximity to water and also shrubbery in marshy habitats.

attenuatus (Scudder) 1869. Longtailed Meadow Katydid (Fig. 35). Adults are present from late summer to early fall. This species is found around ponds, swamps, and marshes. In southern Indiana it has been taken in the Wabash River Valley.

ConocephaZus fasciatus faseiatus ConocephaZus brevipennis

-


ConocephaZus nemoraZis

Cono cephaZus nigrop Zeurm

ConocephaZus s t r i c t u s

ConocephaZus attenuatus


-

ConocephaZus sa Ztms A t Zanticus testaceus

saltans (Scudder) 1872. Prairie Meadow Katydid (Fig. 36). This species has been taken in late summer and early fall in central and northern Indiana. It has been taken in habitats ranging from open fields to cattail marshes.

Subfamily DECTICINAE (The Shieldbearing Katydids)

Genus ATLANTICUS Scudder testaceus (Scudder) 1900. Shortlegged Shield Bearer (Fig. 37). Adults of this species can

be taken most of the summer. The species is taken primarily in the low marginal shrubbery or leaf litter of woodlands.

davisi Rehn and Hebard 1916. Davis' Shield Bearer (Fig. 38). Adults are present throughout the summer and early fall. This species has been taken only in the southern half of the state and. mostly from sparsely wooded ravines.

Family GRYLLACRIDIDAE SUBFAMILY GRYLLACRlDINAE

Genus CAhfETONOTUS Uhler carolinensis Gerstaecker 1860. Carolina Leaf Roller (Fig. 39). Adults of this species have

been taken in late summer and early fall. I t is known from the southern two-thirds of the state in deciduous woodlands and associated shrubbery.

Subfamily RHAPHIDIPHORINAE (The Camel Crickets)

Genus TACHYCINES Adelung *asynamoms Adelung 1902. Greenhouse Stone Cricket (Fig. 40). This adventive species

may be taken year round where it has become established in greenhouses. It has been taken from only two greenhouses in the state.


Tachycines asynamorus

Camptonotus carozinensis

Udeopsy ZZa robusta


Genus UDEOPSY LLA Scudder

*robusta (Haldeman) 1850. Robust Camel Cricket (Fig. 41). Adults are present from late spring to early summer. This rather striking species has evidently become established in the western part of Indiana since 1920, and Indiana may be its present eastern-most distribution. In Lafayette, Indiana, specimens have been taken in gardens, under boards and porches, and in old stumps.

Genus CEUTHOPHILUS Scudder

brevipes Scuddbr 1862. Shortlegged Camel Cricket (Fig. 42). Adults are present from late spring through early fall. Known only from southern Indiana, this species inhabits caves and moist woodlands.

graclipes (Haldeman) 1850. Slenderlegged Camel Cricket (Fig. 43). Adults are known to occur throughout the summer and early fall. In Indiana it is known only from southern counties. I t inhabits woodlands and can be found in natural and man-made shelters.

stygius (Scudder) 1861. Cave Camel Cricket (Fig. 44). Adults are present from mid to late summer. Known only from the southern half of the state, this species often frequents wet caves and rocky embankments.

meridionalis Scudder 1894. Striped Camel Cricket (Fig. 45). Adults have been taken from mid to late summer. It inhabits wooded areas throughout the state.

latens Scudder 1862. Blacksided Camel Cricket (Fig. 46). Adults are present from mid summer to early fall. This species is known from wooded areas throughout the state.

maculatus Harris 1941. \Spotted Camel Cricket (Fig. 47). Adults are present throughout the summer and early fall. This species prefers xeric wooded areas. It has not been taken in northern Indiana.

I Ceuthophi lus brevipes I Ceuthophi lus graci Zipes


Ceuthophi Zus Zatens

Ceuthophi Zus meridionaZis

Ceuthophi Lus macu Latus


tenebrarum Scudder 1894. Shaded Camel Cricket (Fig. 48). Adults of this species are present throughout the summer. It is most common in wooded areas and is restricted to southern Indiana.

seclusus Scudder 1894. Secluded Camel Cricket (Fig. 49). This species is present throughout the summer. This species is known from only one county in Indiana and probably does not occur in the northern half of the state. It has been taken in dry wooded areas.

uhleri Scudder 1862. Uhler's Camel Cricket. (Fig. 50). Adults have been taken in Indiana in late summer and early fall. The species is common throughout the state in wooded areas.

divergens Scudder 1862. Divergent Camel Cricket (Fig. 51). This species has been taken throughout the summer. It evidently prefers'drier wooded areas.

guttulosus guttulosus T. Walker 1869. Leaflitter Camel C~icket (Fig. 52). Adults of this species have been reported from early spring to mid fall. The species is associated with leaf litter in forested areas under which the soil contains rodent burrows.

guttulosus thomasi Hubbell 1936. Thomas' Camel Cricket (Fig. 53). This subspecies apparently merges with C. g. guttulosus in Indiana.

elegans Hubbell 1934. Elegant Camel Cricket (Fig. 54). Although adults apparently may be present throughout the year (Hubbell, 1936), Indiana specimens have been taken in the spring. The species has been taken in corn fields and wooded areas.

Family GRYLLOTALPIDAE (The Mole Crickets)

Subfamily GRYLLOTALPINAE Genus NEOCURTILLA Kirby

a hexadactyla (Perty) 1832. Northern Mole Cricket (Fig. 55). Adults have been taken from mid summer through mid fall. This species is known throughout the state where it inhabits mud and sand banks of streams, ponds, and lakes.

I CeuthophiZus tenebrarum Ceu thophi Zus see Zusus


Ceuthophi Zus uhZeri

CeuthophiZus guttuZosus guttuZosus

I Ceuthophi Zus divergens

I CeuthophiZus guttuZosus thomasi I


Ceuthophi Zus eZegans

Family GRYLLIDAE (The Crickets)

Subfamily MYRMECOPHILINAE (The Antloving Crickets)

Genus MYRMECOPHILA LatreilIe

pergandei Bruner 1884. Eastern Antloving Cricket (Fig. 56). Adults are present from late spring to early fall. This species is common in southern Indiana, but to the north it has been taken only in Tippecanoe County. The species occurs in ant nests, often of Camponotus melleus Say.

Subfamily GRYLLINAE (The House and Field Crickets)

Genus MIOGRYLLUS Saussure (Lesser Field Crickets)

verticalis (Serville) 1839. Stripeheaded Field Cricket (Fig. 57). Adults are present from late spring through early fall. This species is restricted to southern Indiana where it may be found most frequently under logs and rocks on wooded hillsides.

Genus GRYLLUS Linnaeus (Field Crickets)

pennsylvanicus Burmeister 1838. Fall Field Cricket (Fig. 58). Adults present from mid summer to mid fall. This common cricket can be found throughout Indiana in open fieId environments.


Giy ZZus pennsy Zvanicus

Miogryl lus vert icaZis

&y ZZus vernalis

1976 THE GREAT LAKES ENTOMOLOGlST 47

vernalis Blatchley 1920. Smaller Spring Field Cricket (Fig. 59). Adults of this species are present in spring and early summer. Otherwise, its ecology and distribution are similar to that of G. pennsylvanicus, being almost ubiquitous.

*fultoni (Alexander) 1957. Fulton's Field Cricket (Fig. 60). Adults are known to occur in late spring and early summer. In Indiana this species is known only from Crawford County in the south. Alexander (1957) stated that this species was expected to occur on dry, Andropogon covered slopes. Crawford County provides such habitats.

*veletis (Alexander and Bigelow) 1960. Larger Spring Field Cricket (Fig. 61). Adults are present from late spring to early summer. This cricket has been found throughout the state and its habitat is similar to G. pennsylvanicus.

Genus ACHETA Linnaeus

domesticus (Linnaeus) 1758. House Cricket (Fig. 62). Adults can be found throughout the year. This species is adventive and domestic, and probably occurs in every county of the state.

Subfamily NEMOBIINAE (The Ground Crickets)

Genus ALLONEMOBIUS Hebard

fasciatus (DeGeer) 1773. Striped Ground Cricket (Fig. 63). Adults are present from mid summer to mid fall. This relatively common species may be taken in most moist habitats near marshes, ponds, lakes, and streams.

'tinnulus (Fulton) 1931. Tinkling Ground Cricket (Fig. 64). Adults are present in late summer and early fall. This species has been found over the state in deciduous wooded areas.

Cry ZZus fuZtoni G r y ZZus ve Zetis I


Acheta domesticus

AZZonemobius tinnuzus

- -

AZZonemobius fasciatus

A Zlonemobius aZ Zardi


*allardi (Alexander and Thomas) 1959. AUard's Ground Cricket (Fig. 65). Adults are present from late summer to mid fall. This species has been taken throughout the state in open field habitats.

maculatus (Blatchley) 1900. Larger Spotted Ground Cricket (Fig. 66). Adults are present from late summer to mid fall. This widespread species is known to occur in open woodlands.

griseus griseus (E. M. Walker) 1904. Gray Ground Cricket (Fig. 67). This species has been taken in late summer and early fall. Records of this ground cricket are relatively scarce and so far it has not been collected in northern Indiana, although it is expected to occur there. It occurs in upland, sandy soil environments.

Genus NEONEMOBIUS Hebard

variegatus (Bruner) 1893. Smaller Spotted Ground Cricket (Fig. 68). Adults are present from late summer to early fall. The species occurs only in southern and central Indiana. I t is known to inhabit grassy areas along streams and fence rows in open woods.

palustris (Blatchley) 1900. Marsh Ground Cricket (Fig. 69). Adults are present from mid summer to early fall. This species is known to inhabit sphagnum bogs (Johnstone and Vickery, 1970), and thus has a discontinuous distribution over the state.

Genus EUNEMOBIUS Hebard

carolinus (Scudder) 1877. Carolina Ground Cricket (Fig. 70). Adults have been taken from mid summer through mid fall. This species is known from scattered localities over most parts of the state and can be found in wet environments as in marshes or around the margins of ponds and streams.

AZZonemobius macuZatus AZZonemobius griseus griseus


Neonemobius variegatus

Eunemobius caroZinus

Neonemobius palustr is

Eunemobius me kdi;cs


*melodius (Thomas and Alexander) 1957. Melodious Ground Cricket (Fig. 71). Adults are known from late summer to early fall. Scattered samples of this species have been taken in both northern and southern Indiana. I t is found in marshy habitats.

confusus (Blatchley) 1903. Confusing Ground Cricket (Fig. 72). Adults are present from late summer to early fall. It has been taken in scattered localities throughout the state. Blatchley (1920) adequately described its habitat as low, damp woods, usually near water, and in leaf litter.

Subfamily OECANTHINAE (The Tree Crickets)

Genus NEOXABEA Kirby

bipunctata (DeGeer) 1773. Twospotted Tree Cricket (Fig. 73). Adults are present in late summer and early fall. This species should occur throughout the state. It is most commonly taken on vaious deciduous trees and sometimes in vines and shrubs.

Genus OECANTHUS Serville

niveus (DeGeer) 1773. Narrowwinged Tree Cricket (Fig. 74). Adults are known from late summer to mid fall. This species frequents the crowns of several species of deciduous trees and also undergrowth.

fultoni T. J. Walker 1962. Snowy Tree Cricket (Fig. 75). Adults are known from mid summer to early fall. In Indiana this species has only been taken in the central region. Walker (1962) states that it may occur in shrubbery, vines, fruit trees, and fence rows.

exclamationis Davis 1907. Davis' Tree Cricket (Fig. 76). Adults are present in the late summer and early fall. This species, which is known niainly from central and southern Indiana, may be found in deciduous trees and undergrowth.

Eunemobius c o n f u s u s Neoxabea b i p u n c t a t a


Oecanthus n i v e u s

Oecan thus e x c t a m a t i o n i s

Oecan thus fu Z t o n i

Oecan thus Z n t C ~ e ~ n t s


latipennis Riley 1881. Broadwinged Tree Cricket (Fig. 77). Adults are known from late summer and early fall. This species is known from much the same areas as 0. exclamationis in the state, but occurs in low growths of vegetation such as shrubbery, along fence rows, and in open fields.

nigricornis F. Walker 1869. Blackhorned Tree Cricket (Fig. 78). Adults are present from mid summer to early fall. This common species may be found throughout Indiana in open fields and weedy environments.

argentinus Saussure 1874. Argentina Tree Cricket (Fig. 79). There are evidently two generations a year in this species (Walker, 1963). In Indiana, we have taken adults throughout the summer and early fall. Known mainly from the southern part of the state, it is commonly found in weedy growths and lower portions of trees.

quadripunctatus Beutenmiiller 1894. Fourspotted Tree Cricket (Fig. 80). This wide spread species may be found in late summer and early fall. Its habitat is similar to that of 0. argentinus.

pini Beutenmiiller 1894. Pine Tree Cricket (Fig. 81). Dr. R. D. Alexander (pers. comm.) has indicated to us that he has taken specimens in Parke County at Turkey Kun State Park. This would represent a newly reported state record for the species. The species is normally associated with pines.

Subfamily TRIGONIDIINAE (The Winged Bush Crickets)

Genus ANAXIPHA Saussure

exigua (Say) 1825. Say's Bush Cricket (Fig. 82). Adults are present from mid summer to early fall. This small cricket has been taken from much of the state where it frequents dense thickets and low moist habitats.

Oecanthus n ~ i g r i c o r n i s Oecanthus a r g e n t i n u s


A n a x i p h a e x i g u a - . ., Phy ZZopa:?xa : X , C C S _ - X S


H a p i t h u s a g i t a t o r O r o c h a r i s s a Z t a t o r

Genus PHYLLOPALPUS Uhler

pulchellus Uhler 1864. Handsome Bush Cricket (Fig. 83). Adults are present in late summer and early fall. This striking cricket is known mainly from the southern half of the state, the northern most record being from Tippecanoe County. It is found in undergrowth, grasses, and small trees, especially near water.

Subfamily ENEOETERINAE (The Larger Brown Bush Crickets)

Genus HAPITHUS Uhler

agitator Uhler 1864. Restless Bush Cricket (Fig. 84). Adults are present in late summer and early fall. The species is known only from the southern two-thirds of the state where it may be found in a great many habitats ranging from upland woods to open fields and shrubs.

Genus OROCHARIS Uhler

saltator Uhler 1864. Jumping Bush Cricket (Fig. 85). Adults are present in late summer and early fall. This species is known only from the southern two-thuds of the state where it occurs in deciduous trees, herbaceous undergrowth, and shrubs.

LITERATURE CITED

Alexander, R. D. 1957. The taxonomy of the field crickets of the eastern United States (Orthoptera: Gryllidae: Acheta). Ann. Entomol. Soc. Amer. 50:584-602.

Alexander, R. D., A. E. Pace, and D. Otte. 1972. The singing insects of Michigan. Great Lakes Entomol. 5: 33-69.


Blatchley, W. S. 1903. The Orthoptera of Indiana. 27th Ann. Rept., Dept. Geol. Nat. Res. Indiana. 123-47 1.

Blatchley, W. S. 1920. Orthoptera of northeastern America. The Nature Publ. Co. Indianapolis. 784 p.

Cantrall, I. J. 1968. An annotated list of the Dermaptera, Dictyoptera, Phasmatoptera, and Orthoptera of Michigan. Mich. Entomol. 1: 299-346.

Cantrall, I. J. and F. N. Young. 1954. Contrasts in the orpthopteran faunas of grasslands, forests, and transitional areas in southern Indiana. Proc. Ind. Acad. Sci, 1953. 63: 157-162.

Chandler, L. 1966. The origin and composition of the insect fauna. p. 345-361. In: Natural Features of Indiana. A. A. Lindsey, ed. Ind. Acad. Sci. Indianapolis.

Fox, H. 1915. Notes on the Orthoptera and Orthoptera habitats in the vicinity of Lafayette, Ind. Proc. Ind. Acad. Sci., 1914:287-321.

Hebard, M. 1934. The Dermaptera and Orthopera of Illinois. Bull. Ill. Nat. Hist. Surv. 20: 125-279.

Hubbell, T. H. 1936. A monographic revision of the genus Ceuthophilus (Orthoptera: Gryllacrididae: Rhaphidiphorinae). Univ. Fla. Biol. Sci. Ser. Publ. 2:l-551.

Johnstone, D. E. and V. R. Vickery. 1970. Notes on the palustris-cubensis complex of the genus Neonemobius Hebard (Orthoptera: GryUidae: Nemobiinae). J. Georgia Entomol. Soc. 5: 233-241.

Lindsey, A. A. (ed.). 1966. Natural Features of Indiana Ind. Acad. Sci Indianapolis. xxix + 600 p.

Merriam, C. H. 1898. Life zones and crop zones of the United States. U.S. Dept. AD., Div. BioL Surv., Bull. No. 10: 1-79.

Stein, J. L. and W. P. McCafferty. 1975. Diagnostic tables to the long-horned grasshoppers and crickets of Indiana. Purdue Univ. Agr. Exp. Sta Res Bull. No. 921:l-20.

Walker, T. J. 1962. The taxonomy and calling songs of the United States kee crickets (Orthoptera: Gryllidae: Oecanthinae) I. The genus Neoxabea and the niveus and varicornis groups of the genus Oecanthus. Ann. Entomol. Soc. Amer. 55:303-322.

Walker, T. 1. 1963. Taxonomy 11. The nigricornis group of the genus Oecanthus. Ann. Entomol. Soc. Amer. 56:772-789.'

Young, F. N. and I. J. Cantrall. 1956. Orthoptera of relict prairie fragments in Greene County, Indiana. Proc. Ind. Acad. Sci., 1955. 65: 11 1-1 15.


INSECTS ASSOCIATED WITH MICHIGAN BUMBLEBEES (BOMBUS SPP.)l

Robert W. Husband2 and Thomas M. Brown3

Studies of insect associates of bumblebees are not new. For example, Tuck (1896, 1897) reported over 50 species of insects associated with nests of British bumblebees. Sladen (1912) discussed nest associates and parasites of European bumblebees, and Plath (1934) published similar data for American bumblebees. Postner (1952) published more detailed data. He listed over 60 taxa of insects associated with bumblebees near Erlangen, Germany.

Michigan bumblebees may be divided into three groups of species which have northern, southern and cosmopolitan distributions. In an attempt to determine why many species have ranges which terminate within Michigan, a variety of studies, including studies of associations of bumblebees and other organisms, were initiated in the laboratory of T. Wayne Porter at Michigan State University and continued at Adrian College.

The purpose of this paper is to give a preliminary list of the insects found in association with Michigan bumblebees of the genus Bombus, This study follows, in part, the format used by Postner in reporting bumblebee associates. More than 50 taxa found in the nests of, on, or as internal parasites of ten species of bumblebees are included. The term taxa is used rather than species because it was not always possible to determine larvae and nymphs to species level. Additional studies of selected species of insects and associated bumblebees are planned.

METHODS

Several collecting methods were used over a period of 10 years. More than 60 nests were examined, most of which were collected at night. About half of the nests were placed in modified Berlese funnels and insects were collected below in 70% ethyl alcohol. Some nests were taken apart piece by piece, and insects encountered were collected and placed in ethyl alcohol. All nests were active immediately prior to examination. Bees were dissected for internal parasites. Insects were determined to the nearest taxa and forwarded for substantiation to the authorities listed in the acknowledgments.

RESULTS

In the following list of taxonomic categories, the Roman numeral behind the name indicates the number of nests in which the particular insect was found. The Arabic numeral indicates the number of specimens obtained. When a number is not present, insects were removed from bumblebees not associated with nests. The bumblebees associated with a particular insect are listed after the numbers following the name of the insect associate.

COLLEMBOLA

ENTOMOBRYIDAE, Entomobrya assuta Folson (II,20) Bombus fervidus, B. americanorum

E. griseoolivata (Packard) (IV,71) B. affinis, B. fervidus, B. americanorum E. multifasciata Tullberg (IV,6) B. bimaculatus, B. fervidus, B. americanorum Pseudosinella violenta (Folsom) (I,2) B. americanorum P. sexoculata Schott (11,134) B. bimaculatus, B. americanorum

l~ontr ibut ion No. 209, W. K. Kellogg Biological Station, Michigan State University, Hickory Comers, Michigan 49060.

2 ~ d r i a n College. 3 ~ i c h i ~ a n State University.


Wiflowsia buski Lubbock (111,309) B. americanorum W. buski distincta (II,24) B. americanorum W. buski nigrornaculata (III,12) B. americanorum

HYPOGASTRURIDAE, SchoteNa glasgowi Folsom (I,37) B. bimaculatus ISOTOMIDAE, Isotomu albeh Packard (1,184) B. bimaculatus TOMOCERIDAE, Tomocerus flavescens Tullberg (I,20) B. americanorurn

ORTHOPTERA

BLATELLIDAE, Parcoblatta pennsylvanica (DeGeer) (1,l) B. bimaculatus GRYLLLDAE (Nemobinae), Allonemobius sp. (1,l) B. bimaculatus

PSOCOPTERA

LIPOSCELLIDAE, Liposcelis sp. (VIII, c 30) B. affinis, B. americanorum, B. bimaculatus, B. fervidus, B. grisceocollis, B. vagans

HOMOPTERA

CICADELLIDAE, Erythroneura sp. (1,l) B. bimaculatus

HEMIPTERA

ANTHOCORIDAE, Xylocoris galactinus (Fieber) (1,l) B. bimaculatus

THYSANOPTERA

THRIPIDAE, Frankliniella tritici (Fitch), Bomhus nevadensis PALAEOTHRIPIDAE, Haplothrips leucanthemi (Scpank) (1,l) B. birnaculatus

COLEOPTERA

ANTHICIDAE, Anthicus sp. (I,4) B. fervidus CARABID AE, Pasimachus sp. (1,l) B. americanorum CRYPTOPHAGIDAE, Antherophagus ochraceus Melsheirner (X,35) B. affinis, B. ameri-

canorum, B. fervidus, B. vagans Cryptophagus valens Casey (1,101 B. affinis Cryptophagus sp. (1,l) B. americanorum Cryptophagus sp. (?) or Antherophagus sp. (?) (XI,683) B. affinis, B. americanorum,

B. bimaculatus, B. fervidus B. grisceocollis Genus unknown (1 , l ) B. americanorum

CUCUJIDAE, Ahasverus advena (Waltl) (III,8) B. americanorum, B. griseocollis DERMESTIDAE, Attagenus sp. (IV,10) B. americanorum, B. fervidus

Genus unknown, B. terricola HISTERIDAE, Dendrophilus sp. (II,2) B. americanorum LATHRIDIIDAE, Enicmus consimilis (Mannerheim) (1,l) B. americanorum

Corticaria sp. (1,l) B. bimuculatus Corticaria sp. (?) (1,l) B. bimaculatus

MELOIDAE, Genus unknown, B. affinis, B. griseocollis MYCETOPHAGIDAE, Typhaea stercorea (Linnaeus) (III,9) B. americanorum. B. fervidus,

B. griseocollis NITIDULIDAE, Carpophilus sp. (I,2) B. bimaculatus ORTHOPERIDAE, Sericoderus sp. (I,4) B. bimaculatus

Sericoderus sp. (?) (I,13) B. bimaculatus STAPHYLINIDAE, Aleocharinae (?) (1,l) B. americanorum

Subfamily and genus unknown (II,7) B. bimaculatus, B. vagans TENEBRIONIDAE, Cynaeus angustus (LeConte) (I,2) B. arnericanorum

Neatus tenebrioides (Palisot) (IV,50) B. americanorum, B. fervidus Tenebrio molitor Linnaeus (III,24) B. americanorum

NEUROPTERA

CHRYSOPIDAE, Chrysopa sp. prob. C. carnea Stephens, Bombus sp. Chrysopa sp. (?) Bombus affinis

1976 THE GREAT LAKES ENTOMOLOGIST

LEPIDOPTERA

NOCTUIDAE, Epizeuxis americalis (Geunee) (11,s) B. fervidus, B. griseocollis HERMINIINAE, genus unknown (I,3) B. americanorum PYRALIDAE, Vitula edmandsae (Packard) (X,1297) B. affinis, B. americanorum, B.

fervidus Pyralis farinalis Linnaeus (IV,23) B. americanorum, B. fervidus Hypsopygia costalis (Fabricius) (1,l) B. fervidus

TINEIDAE, Acedes fuscipunctella (Haworth) (V,11) B. affinis, B. americanorum, B. fervidus

DIPTERA

CALLIPHORIDAE (1,l) B. vagans CECIDOMYIIDAE, Lestodiplosis sp. (1,l) B. vagans CONOPIDAE, Conopinae, 32 larvae in abdominal cavities of B. affinis, B. bimaculatus, B.

fervidus, B. ternarius, B. terricola Myopinae, 1 larva in abdominal cavity of B. bimaculatus

MILICHIIDAE, Leptometopa sp. (II,16) B. americanorum, B. vagans MUSCIDAE, Fannia canicularis (Linnaeus) (111,194) B. americanorum, B. fervidus

Fannia sp. (111,120) B. affinis, B. americanorum, B. fervidus Muscina sp. (I,4) B. vagans

PHORIDAE, Dohrniphora sp. (II,2) B. americanorum PSYCHODIDAE, Psychoda sp. (1,l) B. americanorum SARCOPHAGIDAE (III,14) B. americanorum, B. bimaculatus, B. vagans. Ten sarcophagid

pupae heavily parasitized by Melittobia sp. Four larvae were taken from the abdominal cavity of B. terricola

Miltogramminae (1,l) B. bimaculatus SCATOPSIDAE, Scatopse fuscipes (Meigen) (1,l) B. fervidus

Genus unknown (1,s) B. americanorum SCIARIDAE, Bradysia sp. (II,2) B. bimaculatus, B. vagans

SIPHONAPTERA

CERATOPHYLLIDAE, Orchopeas leucopus (Baker) (II,2) B. bimaculatus, B. fervidus HYSTRICHOPSYLLIDAE, Epitedia wenmanni (Rothchild) (I,2) B. americanorum

HYMENOPTERA

BRACONIDAE, Apanteles mephoptericis (Packard) (V,23) B. affinis, B. bimaculatus, B. fervidus, B. griseocollis

EULOPHIDAE, Melittobiz chalybii Ashmead (I, over 500) from B. vagans pupa Melittobiz sp. (I, over 1000) from sarcophagid pupa in nest of B. vagans Pediobius sp., near P. williamsoni Girault (1,l) B. bimaculatus

FORMICIDAE Monomorium pharaonis ( ~ i n n a e u s ) ~ (I,95) B. americanorum Solenopsis molesta (Say) (I,9) B. americanorum Tapinoma sessile (Say) (I,21) B. americanorum Tetramorium caespitum (Linnaeus) (1,l) B. americanorum

VESPIDAE, Polistes fuscatus (Fabricius) (1,l) B. fervidus

DISCUSSION

Studies of insect associates of American bumblebees have been sparse in comparison to European studies. Plath (1934) dealt with insect associates more than Hobbs (1965, 1967) or Janzen (1971). These references considered bumblebees from Massachusetts, Western Canada and Central America. In Sweden, Hasselrot (1960) observed such insects

4 ~ h i s species is established in the Natural Science Building at Michigan State University and entered the nest after it was placed indoors.


as Aphomia sociella Linnaeus, Brachyoma devio Fallen, Volucella bombylans Linnaeus, Fannia sp., Antherophagus nigricomis Fabricus, and Lasius niger. Hasselrot, Sladen (1912) and Cumber (1949) discussed the relationships of insects associated with bumblebees. Skou, Holm and Haas (1963), and Free and Butler (1959) also discussed European bumblebee associates. Our study is most comparable to the work of Postner (1952) in Germany. In both cases, the majority of insect associates were recorded from 30 bumblebee nests. We include data from 30 additional nests which were studied for temperature regulation and other purposes.

The most common groups of insects found in association with German and Michigan bumblebees were the six orders Coleoptera, Lepidoptera, Diptera, Collembola, Hymenoptera and Psocoptera. These frequently associated orders include parasitic insects, scavengers, predaceous insects and a large number of insects for which the food habits are unknown. Six orders found less frequently were Orthoptera, Siphonaptera, Neuroptera, Homoptera, Hemiptera and Thysanoptera. We speculate that Thysanura, Diplura and Strepsiptera will be found in association with some Michigan bumblebees or their nests. The less frequently collected orders may have an accidental or casual relationship to bumblebees. Siphonaptera may be present in nests due to visits by buds or mammals or former utilization of the bumblebee nest as a bud or mammal nest. Thysanoptera and Neuropteralmay be distributed by bumblebees but their role in ,bumblebee nests is unknown. It is conceivable that Orthoptera might be sufficiently numerous to become pests but they were rarely found in our studies.

Collembola have been discussed by Snider and Husband (1966). Although no laboratory studies have been done, Collembola have been reported to be commensals (Postner 1952). Three genera which were found as nest associates in Germany, Entomobrya, Pseudosinella and Tomoceros, were also found in Michigan. The genus Willowsia, not found by Postner, was present in eight nests of Bombus americanorurn Of the nine insects found in three or more nests, only Willowsia buski and T. molitor were found with only one species of bee. Psocoptera were nearly as abundant as Collembola in bumblebee nests. The genus Liposcelis was found in Germany and Michigan. Psocoptera may be scavengers.

Coleoptera were very common in nests in Germany and Michigan. Plath (1922) reported Antherophagus sp. from "almost every one of 50 nests which I encountered, and in one case over 20 beetles were found." Cryptophagidae, including Antherophagus ochraceus Mclsheimer, were ithe most numerous beetles encountered in Michigan bumblebee nests. Antherophagus nigricomis Fabricius was one of the most common beetles reported in German bumblebee nests by Postner. Postner cited 15 families of Coleoptera found with German bumblebees while we found 13 families. Typhaea stercorea (Linnaeus) (Mycetophagidae) and Tenebrio molitor Linnaeus were common to Michigan and Germany.

Lepidoptera were common in several Michigan bumblebee nests. Among the five species found, the pyralids, Vitula edrnandsae (Packard) and firalis farinalis Linnaeus, and the tineid, Acedes fuscipunctella (Haworth), were most common. Bumblebee nests which had V. edmandsae produced more cocoons than nests in which V. edmandsae were lacking (Hobbs et al., 1960). Plath (1934) considered V. edmandsae "comparatively harmless." We found no Galleria melonella (Linnaeus) although Janzen (1971) found this harmful associate in a nest of Bombus pullatus in Costa Rica. The European wax moth, Aphomia sociella Linnaeus, was considered by Hasselrot (1960) to be a more serious pest.

Diptera were very common in bumblebee nests with 10 families found in this study. Fannia canicularis (Linnaeus) was most numerous. Although Hasselrot (1960) and Plath (1934) found Volucella sp. (Syrphidae) numerous in some nests, Postner (1952) did not list this family. Hobbs (1967) reported V. bombycolans (Linnaeus) attacking live larvae and pupae. Hasselrot (1960) reported that the tachinid Brachycoma devia Fallen consumes or parasitizes cocoons of bumblebees. We found no specimens of Volucella. Hobbs (1965) reported that Physocephala texana (Syrphidae) killed a B. rufocinctus queen. Sarcophagidae were removed from the abdominal cavities of several bumblebees. Conopidae were even more numerous and were found in five different species. Scatopse fuscipes Meigen (Scatopsidae) was rare but reported from both Germany and Michigan.


Hymenoptera have been mentioned as serious pests of bumblebees by several American and European scientists. Braconidae, Ichneumo~dae and Eulophidae were relatively uncommon. However, in a given nest, numbers could be very high. Hobbs (1965) and Plath (1934) indicated that ants are major pests of some bumblebee nests. Since most of our nests were collected at night and since ants do not tend to remain in nests for long periods of time, ants may be under-represented in this study.

We have not included Psithyms spp., social parasites of bumblebees, since we did not encounter them in nests examined and the distribution data of field collected Psithyms spp. will be published later. Plath (1934) reported Dasyllis grossa (Asilidae) attacking Boinbus fewidus. We have not yet observed this. We did not find Brachycoma sp. (Tachinidae) found by Plath. We found no Dermaptera, although Holm (personal commu~cation) indicated that they are common in Denmark. We did not find Protura although Postner (1952) reported this group from bumblebee nests in Germany.

There is insufficient evidence to speculate on many of the associations of bumblebees and other insects. Several Diptera and Hymenoptera are internal parasites. Most of the insects that appear to be scavengers in a nest may serve a useful role in removing wastes and debris. Some predaceous insects may function to control populations of scavenger insects. Some insects may serve to keep fungus from overgrowing the nest. Relative humidity is usually more than 60% and the temperature is usually more than 30°C.

We are unaware of insects which are species specific to bumblebees and no other taxonomic group, as are three species of Acarina, Kuzinin laevis, Pneumolaelaps spp. and Locustacarus buchneri. None of these acarine species survive the winter in nests in Michigan. K. laevis deutonymphs and Pneumoloelaps spp. females attach to hibernating queens. Insects have not been found on hibernating queen bumblebees. Larval Diptera and Hymenoptera which are internal parasites of bumblebees do not appear restricted to a bumblebee host.

If associations between bumblebees and other insects were close, one would expect these insects to occur wherever bumblebees are found. This may be the case if some parasitic insects are host restricted to bumblebees as is the parasitic mite Locustacarus buchneri. K. laevis, a fungus, pollen and nematode feeding mite which survives only on hibernating queens, is found from above the Arctic Circle to the equator. Insects which are only casually associated with bumblebees in temperate regions would be absent or reduced in high arctic regions where the climate is less favorable. In contrast, the few species of bumblebees that exist in the tropics of Brazil would have more casually associated insects on the bees and in their nests. Additional observations and experiments with tropical bumblebees are needed to confum this hypothesis. It is still not possible to give a clear explanation of why there are 17 species of bumblebees in Michigan and six species in Brazil, but the balance of obligate and casual insect associates in and out of the nest may be significant to bumblebee distribution.

ACKNOWLEDGMENTS

We are very grateful to the foIlowing authorities at the U.S. National Museum of Natural History and the Entomological Research Institute in Ottawa, Canada, for aid in identification and for other assistance: D. M. Anderson, E. C. Becker, W. A. Connell, K. C. Emerson, T. L. Erwin, 0. S. Flint, Richard H. Foote, R. J. Gagne, G. Gordh, J. L. Herring, J. M. Kingsolver, Lloyd Knutson, J. P. Kramer, M. MaKay, P. M. Marsh, J. E. H. Martin, W. Mason, A. S. Menke, E. L. Mockford, E. G. Momoe, K. O'Neill, Reece I. Sailer, D. R. Smith, T. J. Spilman, G. Steyskal, D. M. Weisman, W. W. Wirth and C. M. Yashimoto. Richard Snider, Michigan State University, was very helpful with identification of and comments on Collembola. Irving Cantrall, University of Michigan, provided identifications of Orthoptera. We are particularly grateful to Dr. T. Wayne Porter, Michigan State University, for encouragement over a long period of time, for suggestions, and for the use of his laboratory.


LITERATURE CITED

Cumber, R. A. 1949. Bumblebee parasites and commensals found within a thirty mile radius of London. Proc. Roy. Entomol. Soc. London. Ser. A24:119-127.

Free, John B. and Colin G. Butler. 1959. Bumblebees. Collins, London. 208 p. Hasselrot, T. B. 1960. Studies on Swedish bumblebees (Genus Bombus Latr.). Opu. Sc.

Entomol. Suppl. 17: 1-192. Hobbs, G. A. 1965. Ecology of species of Bombus Latr. (Hymenoptera: Apidae) in

southern Alberta. 111. subgenus Cullumanobombus Vogt. canad. ~n tomol . 97: 1293-1302.

Hobbs, G. A. 1967. Ecology of species of Bombus (Hymenoptera: Apidae) in southern Alberta. VI. Subgenus Pyrobombus. Canad. Entomol. 99: 1271-1292.

Hobbs, G. A., J. F. Virostek and W. 0. Nummi. 1960. Establishment of Bombus spp. (Hymenoptera: Apidae) in artificial domiciles in southern Alberta. Canad. Entomol. 92:868-872.

Janzen, Daniel H. 1971. The ecological significance of an arboreal nest of Bombus pullatus in Costa Rica. J. Kansas Entomol. Soc. 44(2):210-216.

Plath, 0. E. 1922. Notes on the nesting habits of several North American bumblebees. Psyche. 29(5-6):189-202.

Plath, 0. E. 1934. Bumblebees and their ways. Macmillan Co. 201 p. Postner, M. 1952. Biologisch-okologische untersuchungen an hummeln and ihren nestern.

Veroff. Ubersee Museum, Bremen. Reihe A 2:45-86. Skou, J. P., S. V. Holm and H. Haas. 1963. Preliminary investigations on disease of

bumblebees. Roy. Vet. and Ag. College Yearbook. pp. 2741. Sladen, F. W. L. 1912. The humble-bee, its .life history and how to domesticate it.

Macmillan. London. 283 p. Snider, Richard J. and Robert W. Husband. 1966. Collembola found in bumblebee nests.

Trans. Amer. Micros. Soc 85(3):473475. Tuck, W. H. 1896. Inquiline and other inhabitants in nests of aculeate Hymenoptera.

Entomol. Mon. Mag. 7: 153-155. Tuck, W. H. 1897. Coleoptera etc., in the nests of aculeate Hymenoptera. Entomol. Mon.

Mag. 8:58-60.


NEUROCORDULIA (PLA TYCORDULIA) XANTHOSOMA (WILLIAMSON) IN TEXAS (ODONATA: LIBELLLII-I DAE: CORDULIINAE)

Curtis E. Williams1

The rarity of a species is usually judged by the number of specimens in collections, both private and institutional, and the number of places where it has been found or seen. On this basis, Neurocordulia xanthosoma (Williamson) is rare. Its crepuscular habits, its brown coloration, and its habitat requirements, coupled with the habits of most collectors, have been its protection. Nevertheless, because of its apparent special requirements and man's modification of, and encroachment on, its environment, it may soon be a species of the past.

It was first collected in 1907, when Oklahoma was still Indian Territory, near Wister, now of Le Flore County, by Mr. E. B. Williamson on June 4, and on August 2 by Frank Collins, a boy helper living in the area. Mr. Williamson described it in 1908 as Platycordulia xanthosoma, n.gen., n.sp. Since then it has been reported from two counties in southeastern Kansas (Kennedy, 1917), seven additional counties in Oklahoma (Ortenburger, 1927; Bird, 1932; Pritchard, 1936; Byers, 1937; Bick, 1951; Bick & Bick, 1957), and from Marion County in northeastern Texas (Harwell, 1951). The nymphs from the Mississippi River at Alma, Wisconsin, and Fairport, Iowa, described as of this species by supposition (Byers, 1937) have proven to be Neurocordulia molesta Walsh (Needham & Westfall, 1955:355). Although Needham & Westfall (1955:355,360) accepted the opinion of R. S. Hodges, who made an extensive study (unpublished) of the genus Neurocordulia, and treated Platycordulia as a subgenus, some odonatologists still question the change in status because of marked differences in the wing venation as first pointed out by Williamson.

My f i s t introduction to Neurocordulia (Platycordulia) xanthosoma was on June 20, 1971, in McLennan County on Lake Creek, a spring-fed stream flowing from a reservoir by the same name. A large thunder-shower was approaching, casting its dark shadow over the area, and had caused the dragonflies to cease flying, Then I saw one flitting in and out among exposed tree roots in the shadows of the vertical bluff-bank of the creek. It flew up and down, in and out of a maze of roots, thus slowly and completely covering a section of the bank. When I netted it I thought it was a teneral male of some common species and was not impressed with my "catch." However, I placed it in an envelope with my usual stick in the bottom to prevent the eyes from being crushed, then in my cooler to keep it alive and inactive. Later, while photographing it, I realized that it was not teneral and was different from any other dragonfly in my collection.

Upon learning how little was known of the habits and life history of this species, I began paying special attention to it during the remainder of the 1971 season and of each of the following years through 1975.

COLLECTION AND OBSERVATION SITES

Since my original discovery I have observed many individuals and taken additional specimens in Texas as follows: McLENNAN Co.-Lake Creek (stream): June 21 (several observed before sunup) 1971; June 29 (Id) 1972; June 15 (observed one for several hours), June 23 (one observed), August 3 (IS'), August 9 (two seen) 1973. Lake Waw (cove and island): May 14-16 (15 exuviae), May 19 (28,,7?), May 21 (29 + 1 exuvia), May 23 (2d + 3 exuviae). June 4 (39 with exuviae), June 24 (many foraging and 1 mating pair) 1975. Hog Creek: May 27 (19 observed ovipositing), June 10 (3 seen), July 3 (many foraging at dawn) 1974; May 13 (6d 1 9 and more seen) 1975. South Bosque

- -

1704 Foster St., Marlin, Texas 76661.


River: June 19 ( 2 4 1972; May 24 ( 2 4 , June 11 ( 3 4 , June 12 ( 3 4 , June 14 ( 2 4 , and June 26 (19 and several more individuals seen) 1973; June 15 (8 exuviae) 1974 with S. Dunkle. FALLS Co.-Brazos River, near Marlin: May 15, (1 teneral d) 1975.

Some of these specimens have been deposited in the Florida State Collection of Arthropods, Gainesville, Florida, in the E. B. Williamson Collection, Museum of Zoology, University of Michigan, Ann Arbor, Michigan, and in the Baylor University Collection, Waco, Texas.

SITE 1. McLennan County, Lake Creek. The stream originates at the spillway of Lake Creek Reservoir on Farm to Market Road No. 1860, 5 miles west of Riesel.

SITE 2. McLennan County, South Bosque River. Six miles west of Waco city limits on Highway No. 84. I t empties into the southwestern arm of Lake Waco. Generally the banks of the river are steep and heavily overgrown with shrubs, trees and weeds.

SITE 3. McLennan County, Hog Creek. This stream flows into the west side of Lake Waco north of the mouth of South Bosque River.

Fig. 1. Map of the vicinity of Waco, Texas. Collecting sites 1-4 in McLennan County: (1) Lake Creek; (2) South Bosque River; (3) Hog Creek; (4) Lake Waco-cove and island.


SITE 4. McLennan County, Lake Waco. Cove and island on the west side of the lake about a mile northeast of the mouth of Hog Creek. The bottom of the cove is gravel with a heavy overlay of mud and debris.

SITE 5. (not shown on map, about 10 mi. south of Site No. 1) Falls County, Brazos River. Four miles west of Marlin where it is crossed by Highway No. 7.

Each of the streams at Sites 1-3 has a rock and gravel bottom except in the deeper pools where the current is not swift and mud collects. At Site 5 the Brazos River has a mud bottom.

In almost every case where the xanthosoma were located, a vertical bank was involved. It is my belief that these banks are normally one of their habitats. On the days that the river was bankfull, they were driven from these places by the water, and I happened to f i id a place where they re-located along steep banks which were not flooded. In looking for xanthosoma, I tried to find them under all kinds of weather conditions, and just after the huge rain with the river full had to be an unusual condition. But, that was the only time I found them in any quantity. Many other times I have looked for xanthosoma in vain.

FLIGHT AND FORAGING

The time largest numbers of N. xanthosoma can be seen is at dawn and at dusk. The males and females fly together at these periods.

During the dawn and evening flights they are usually unmolested while foraging. Especially in the early morning, large numbers of small insects are on the wing. On several mornings before and until shortly after sunrise, the air above Hog Creek and South Bosque River was dense with swarming males of a tiny mayfly, Choroterpes nanita Traver. N xanthosoma were flying in numbers, apparently haphazardly, in and out of the clouds of these insects and feeding upon them. At a later date I observed them feeding among a mating swarm, aU males, of a slightly larger mayfly, Tricolythodes explicatus (Eaton), over a gravel road which runs along the lake shore and within 40 feet of the cove. This site was at least half a mile from Hog Creek, the nearest stream. Surely with the air so dense with these mayflies and with no competition from other dragonflies, the xanthosoma could eat their fiil in the short time they foraged.

At times, the xanthosoma encounter competition or themselves become the prey. I have seen them at the first light of dawn and also at 8:30 in the evening flying low over the water of a small stream flowing into the South Bosque River. They appeared to be foraging and were making trips up and down Ithe stream for a distance of 50 yards or more. Several A n w junius were also hawking low over the water, but a little higher than the xanthosoma, and were observed catching or trying to catch them.

Between the dawn and dusk flights the xanthosoma may continue their foraging but more cautiously and inconspicuously. At Lake Creek on June 15, 1973, I observed one from 2:00 to 6:00 p.m. Across the narrow stream from where I sat, the bank was low and vertical, and above it was a thick growth of weeds, tau grass and shrubs. Clouds were moving across the sky, so conditions were alternating between sunshine and shadow. When a large cloud obscured the sun for as long as one to three minutes, xanthosoma would come out of some grass and weeds growing beneath a shrub and start foraging at the edge of the weeds and grass. Sometimes the foraging would include the vertical bank of the stream. As it dipped in and out of the weeds, it was also flying sideways, and a distance of 20-30 feet at the edge of the weeds or vertical bank would be covered in 10 to 15 seconds. As soon as the sun came out, it immediately returned to cover beneath the shrub. Several times there was only a second or two of foraging before the sun came out again and cover was sought immediately. Here at Lake Creek there were many larger dragonflies, including some Epicordulia princeps, patrolling their territories. On two occasions, a xanthosoma was observed trying to fly down to the water, or to the vertical bank above the water. It was attacked immediately by two or three of the patrolling dragonflies, and driven away. On the South Bosque River where the xanthosoma occurred, there were very few larger species, and the xanthosoma were not molested

66 THE GREAT LAKES ENTOMOLOGIST Vol. 9. No. 1

while flying in the shadows, even though the sun was shining. Perhaps for protection, those at Lake Creek have adopted a different manner of foraging because as soon as the sun disappeared behind a cloud, the other dragonflies stopped flying and offered no threat.

Occasionally during a period of foraging an individual will be seen very briefly perching much like a libelluline, but most of the time they stay on the a- until they retreat to cover in dense stands of tall grass, weeds, or under the lower limbs of trees (some with grapevines growing over them), but usually in the more densely-leaved shrubs Here they hang-up in a vertical position in much the same manner as species of the Aeshnidae. They are very difficult to locate. Often to flush one out, the shrub must be shaken rather vigorously, and once flushed out it seeks new cover with as little flying as is necessary. They are very reluctant to fly when the sun is high, and one can be approached within an inch before it takes off.

MATING AND OVIPOSITION

The fust mating pair I observed flew from the direction of a stream and lit near me. The only thing I noted that was different from the usual behavior of libellulines was that they alighted beneath a branch of a densely-leaved shrub which was in the shadow of a large tree. This occurred a little after sunrise. On June 24, 1975, at Lake Waco, I saw another mating pair flying over the water of the cove as the pair approached the island. They lit under the limb of a small hackberry tree. This was at 6:30a..m., just after the dawn flight. When they flew away after being photographed at a r q o e of 12-14 inches, I lost sight of them.

Many times I have observed females ovipositing in the streams, but so far, not in the lake. However, at about 9:30 one morning 1 saw a female ovipositiq in a large pond-like area of the creek. She flew hurriedly all over the "pond," pausing vei-y briefly here and there to deposit eggs. Normally, oviposition occurs thirty minutes to an hour after the dawn flight. Harwell (1951:205) reported that at Caddo Lake in Marion County, "The female deposited eggs while attended by the male." All the ovipositkg females I have seen were alone, not even protected, guarded, or disturbed by a male. The males do not patrol an area nor do they seem to establish a territory.

NYMPHS AND EXUVIAE

Over the years since the species was described, several collectors have searched in vain for nymphs or exuviae about which there would be no doubt of their identity as xanthosoma. After locating two habitats of the adults in McLennan County, I too looked many times and many hours for nymphs, and later for the exuviae near the water and as far as 40 to 50 feet from water, on weeds, rocks, small shrubs, and on tree trunks, without success from 1972 through 1974. 1 had assumed that .ranthosoma was strictly a stream dweller. In the winter of 1974 and spring of 1975, I worked very hard searching the two streams, South Bosque River and Hog Creek, which flow into Lake Waco, without finding either nymphs or exuviae.

On June 15, 1974, Sidney W. Dunkle and I searched on foot, then by canoe, along the banks of South Bosque River. On the trunk of a large ash tree leaning from the bank above a place where the water was deepest and pond-like conditions existed, we found four of the coveted exuviae. Later, another was found on a twig in the debris at the base of this tree, and two on the underside of leaves of a poison ivy vine growing up the trunk. At another site, one was found on a dead twig near the edge of the bank about three feet above water. A fuller account of the finding of these exuviae and a description of them will be submitted for publication (Williams & Dunkle, in manuscript).

I took seven adults and saw more on May 13, 1975, a new early record, at Hog Creek. I renewed my efforts to find exuviae, but found none along the streams. Then it occurred to me that they must be emerging mostly in the lake area and later flying up the streams. On May 14, 15 and 16, I examined an area in the general vicinity of the mouths of the river and creek. In a small cove (Site No. 4) measuring about 50 X 25 yards, bounded by the lake shore on two adjacent sides and an island on the third side, I


found 15 xanthosom exuviae: Generally the nymphs had favored the larger tree trunks for emergence, but two exuviae were on plant stems of about oneeighth inch in diameter. All were on plants or trees at the edge of the water or near the shore-line. Most of the exuviae appeared to be a week or more old. After finding these exuviae, I checked the area for several nights about midnight and before dawn but found no tenerals emerging. However, I subsequently found exuviae in the daytime. On May 21, just after the fust light of dawn, I found an emerged female still clinging to its exuvia, another very teneral female, and two exuviae. On May 23, I searched the area again at dawn and at 5:30 a.m., but did not discover any emerging until 6: 10 a.m. and 7:00 a.m. At 7:00 a.m., the moulting occurred on a stem, next to the leaves of a small hackberry tree. On June 4, at 6:00 a.m., I located two tenerals, each very near its exuvia, and at 7: 15 a.m., one in the process of emerging from its nymphal skin. Each was on a leaf, 3 4 feet above ground, of a small tree 2-5 feet from the water. The three were photographed. To me, the above data on the hours of emergence indicate that xanthosoma nymphs do not leave the water until long past midnight, and teneral adults emerge a little before dawn or shortly thereafter. I thought the emergence season had ended in early June but I collected very teneral adults and fresh exuviae as late as June 21.

Emergence often occurs with the nymph in a horizontal position with the ventral side up (Fig. 2), and at heights of 3 inches to 12 feet above ground. On two occasions an exuvia was found about 3 inches above ground on the underside of an exposed horizontal section of the rpot of a small tree.

Some of the adults taken May 13 on Hog Creek were not teneral. Allowing seven days for the maturing of those that were fully adult as the beginning of emergence for the season, and June 21 as the end, I estimate the period to be at least 46 days and maybe longer. Such a long period indicates that the degree of emergence synchronization is very low in this species. The latest date I saw an adult was August 9, 1973.

The places where the exuviae and emerging tenerals were found were all near areas of deeper water or deep pools. During the winter and spring there were downpours of rain

Fig. 2. Neurocordulia xanthosoma (Williamson) exuvia Horizontal position assumed on the underside of a tree limb before the emergence of the adult.


Fig. 3. Neurocordulia xanthosoma (Williamson), exuvia, dorsal view.

which made raging torrents of the streams entering the lake. Apparently the currents favored the particular cove where collecting of the immature stages of xanthosoma was most successful. The surface of the water was thickly covered with driftwood and debris that had washed down. Possibly nymphs clinging to the debris settled where the water was deepest. I t is also possible that nymphs in the later stages of development seek the deeper pools near the mouth of a stream, on the south side of the island, and at a site on the south shore of the cove under a sycamore log, one limb of which was submerged in the water.

These suppositions were verified when I visited the cove and island on December 20 and 22, 1975. The water-level was very low, and some of the logs which normally were completely submerged had one end visible near the shore and the other end extending out into the water. Many of the logs, 4 to 8 inches in diameter and 4 to 12 feet in length, were pulled ashore. I found that the bark was loose and could be removed easily. Seventeen xanthosoma nymphs were found clinging to the logs beneath the bark. They were black like the logs, and were clinging with legs outspread and body tightly pressed against the logs. I found five on the outer surface of logs and other debris. Another was taken from the under-surface of a large rock. Most were taken from the deeper water off the cove-side of the island, although about one-third the total were taken from the water off the opposite shore of the island. Three were taken along the lakeshore across the cove from the island.

The nymphs and some of the water-soaked bark were put in an aquarium in daylight, and soon the nymphs were clinging to the under surface of the bark which was resting on the bottom. Inspections after dark revealed that all nymphs were out crawling on the sandy bottom. The aquarium was in an unheated building, and the water temperature was 50 degrees F, about the same temperature as the water in the lake.

Measurements of length and width were taken of all nymphs. Length stated is from tip of folded labium to tip of anal appendages, and width is the width of abdomen at its widest point. Length and width of the smallest nymph was 6.5 and 5.0 mm, and the


Fig. 4. Island and cove, Lake Waco, where nymphs were fust found on December 20, 1975.

largest nymph was 23.0 and 11.0 mm. Wing-pads of the larger specimens were 6.06.5 mm in length and reached to slightly beyond the junction of abdominal segments 6 & 7. Wing-pads of the smallest specimens are scarcely discernible, and the dorsal hooks are like tiny dots, however the lateral spines are very well developed on the smallest specimen.

Because of the great variation found in the degree of development of these nymphs, I suspect that the larval development of xanthosoma requires at least two years, or that the long period of emerging adults and of egg laying may account for the range of size of the nymphs at the end of the breeding season. Some of the nymphs appeared to be in their final instar.

REDESCRIPTION OF THE FEMALE

The following description of the female is based on two specimens, and is. more detailed than that published for the allotype (Kennedy, 1917).

Face yellow, with labrum more brown than yellow. Vertex yellow, darker on front. Occiput is brighter yellow. Eyes pale, with dark brown line along rear margin.

Thorax a lighter brown than abdomen, a pubescence on dorsum and entire sides softens the color and details. A pale yellow spot surrounds the spiracle. Yellow middorsal carina similar to that of male.

Wing venation similar to that of male, except for a longer anal loop. Crossveins in midbasal spaces may be 0, 1, or 2. Number of cells in the triangle of front wing 3 in 3 wings, 5 in 1 wing, and of all hind wings 2. Brown spots on all antenodal and postnodal crossveins out to tips of wings, darker on postnodals. Crossvein in hind wing triangle heavily marked with brown. Brown spot on subnodus of each fore and hind wing. In hind wing, a brown spot between fourth antenodal crossvein and distal angle of triangle. A larger spot of brown at base of hind wing begins between fust paranal cell and rear end of membranule, leaving this paranal cell mostly clear. I t then extends fonvard to cubitus and outward to proximal angle of triangle. This spot also covers rear of


Fig. 5. Neurocordulia xanthosoma (WiUiamson), adult male, dorsal view.

Fig. 6. Neurocordulia xanttzosoma (Williamson), adult female, dorsal view.

THE GREAT LAKES ENTOMOLOGIST 71

Fig. 7. Neurocordulia xanthosorna (Williamson), adult male, lateral view.

Fig. 8. Neurocordulia xanthosoma (Williamson), adult female, lateral view.


membranule for one-third its length. Upper two-thirds of membranule, white. This white continues rearward along inner border of membranule to its junction with wing margin, bordering brown spot with a thin line of white. Color in wings is more brown than yellow, which causes wings to appear spotted where the color is concentrated.

Abdomen shining golden brown and almost bare of hairs. First three basal segments similar to those of male (Figs. 5,6), swollen most at junction of segments 2 and 3, narrowed posteriorly on 3, thereafter parallel-sided or slightly tapering to segment 10 which is narrower and yellower. Viewed laterally (Figs. 7-81, from apex of segment 3, abdomen tapers regularly through segment 8, not depressed, segments 9 and 10 narrower. Lateroventral carinae well developed on segments 5 to 9, weak and shorter on 4. Just above the carina on each side of these segments is a pale yellow spot extending from the base of the segment to half its length. Seen from above, these appear as narrow, half-length yellow edges.

Caudal appendages sparsely covered with long brown hairs, yellow at base, either yellow or brown beyond. In dorsal view, they are straight-sided on outer margins and convex on inner margins, widest at a point about mid-length, and terminate in a sharp point.

The subgenital plate (Fig. 9) terminates in two rounded lobes which are separated by a semi-circular notch, distal ends of lobes barely surpass hind margin of segment 8. Near these lobes, on the sternum of segment 9, spaced apart by a distance equal to width of the notch, are two small elongate styli which are bent rearward. Between the styli is a dense growth of white hairs. The shape of the lobes and notch may become distorted during the drying process.

VARIATION

Males are slightly smaller than females. Of 14 males and two females described above, measurements in millimeters are as follows: total length: d 48.0-52.0, average 50.11; 9

Fig. 9. Neurocordulia xanthosoma (Williamson), ventral aspect of abdominal segments 8-10 of the female, showing the vulva lamina before the specimen was dried.


51.0-52.5, average 51.75; length of abdomen: d 34.5-38.0, average 36.4; 9 37.0-39.0, average 38.0; anal appendages (cerci): 2.5; length of hind wing: d 35.0-37.0, average 35.7; ? 40.0.

Often I have wondered about the great variation in the extent of color in the wings. I am now convinced that this is entirely governed by age, because all of the specimens taken late in their season have the wings completely suffused with color, while early in the season most of them do not. The color seems to accumulate very slowly throughout their life.

The number of crossveins in the midbasal space of front and hind wings is not of sufficient constancy to be reliable for use in a key. Of 41 males, one male had no crossveins in the midbasal space, four had one in one wing, two had one in two wings, four had one in three wings, and 30 had one in all four wings. Of 10 females, one had no crossveins in the midbasal space, eight had one in all four wings, and one had two in each left wing, and one in each right wing.

The number of cells in the triangles seems to be more stable with three in the forewings of all but one wing of one female with five, and two in the hind wings of all but one wing of one male with three. Needham & Westfall (1955:354) record the number as three-four in the front wings and three in the hind wings.

ACKNOWLEDGMENTS

I am most grateful to Mrs. Leonora K. Gloyd, Museum of Zoology, University of Michigan, for much encouragement and for the many hours she so willingly dedicated to editing and arranging this paper in proper order. I also wish to thank Professor Lewis Berner, Department of Zoology, University of Florida, for identifying the two species of mayflies.

LITERATURE CITED

Bick, G. H. 1951. Notes on Oklahoma dragonflies. Jour. Tenn. Acad. Sci. 26(3):178-180. Bick, G. H., and Juanda C. Bick. 1957. The Odonata of Oklahoma. The Southwestern

Naturalist 2(1):1-18. Bud, R. D. 1932. Dragonflies of Oklahoma. Biol. SUN. Univ. Okla. 4(2):50-57. Bud, R. D. 1932. Platycordulia xanthosoma Williamson (Odonata: Corduliinae). Entomol.

News 43(9):234-235. Byers, C. F. 1937. A review of the dragonflies of the genera Neurocordulia and

Platycordulia. Univ. Mich. Mus. Zool. Misc. Publ. 36, 36 p. Harwell, J . E. 1951. Notes on the Odonata of northeastern Texas. Texas Jour. Sci.

3(2):204-207. Needham, J. G., and M. J. Westfall, Jr. 1955. A manual of the dragonflies of North

America (Anisoptera), including the Greater Antilles and the provinces of the Mexican border. Univ. Calif. Press, Berkeley & Los Angeles, 8 vols. xii + 615 p.

Ortenburger, A. I. 1927. Odonata collected in southeastern Oklahoma during the summer of 1925. Univ. Okla. Acad. Sci., Bull. 6:291-221.

Williams, C. E., and S. W. Dunkle. The larva of the dragonfly Narrocordulia xanthosoma (Odonata: Corduliidae). In Manuscript.

Williamson, E. B. 1908. A new dragonfly (Odonata) belonging to the Cordulinae, and a revision of the classification of the subfamily. Entomol. News 19:428434.


THE SUBCOLONIZATION AND BUILDUP OF TETRASTICHUS JULIS, (HYMENOPTERA: EULOPHIDAE) A LARVAL PARASlTOlD OF 'THE

CEREAL LEAF BEETLE, (COLEOPTERA: CHRYSOMELIDAE) IN THE LOWER PENINSULA OF MICHIGAN1

P. A. Logan, F. W. Stehr, and R. J. Sauer2

ABSTRACT

Following initial establishment of the parasitoid, Tetrastichus julis (Walker), at a carefully managed field nursery, releases of parasitized Oulema melanopus larvae were made by Michigan county agents at preselected sites throughout the lower peninsula during 1970-74. A follow-up recovery program during 1971-75 revealed continued dispersion and population increase for T. julis. An independent census verified the increasing rates of parasitism.

INTRODUCTION

The cereal leaf beetle, Oulema melanopus (Linnaeus) of Eurasian origin, is a univoltine, economically important pest on small grains. It was found in southwest Michigan in the 1950's, but was not identified until 1962 (Castro, Ruppel, and Gomulinski, 1965). A biological control program involving releases of larval and egg parasitoids was begun by the United States Department of Agriculture in 1963 (Dysart, Maltby, andBrunson, 1973).

The larval parasitoid, Tetrastichus julis (Walker), was established at the Michigan State University W. K. Kellogg Farm by 1969 (Stehr, 1970).This parasitoid is bivoltine, with a facultative diapause. Some of the offspring of the f i s t generation emerge in mid-season as a second generation, and the rest enter diapause. Diapausing f i s t generation parasitoids and all of the second generation winter in the soil in the pupal cells of the host. There are usually four to six parasitoids per host larva.

METHODS

Tetrastichus julis was the subject of an extensive subcolonization program throughout the lower peninsula in 1971-74. County agents from the Michigan Cooperative Extension Service collected parasitized larvae from the Kellogg Farm area and released them at preselected "nursery sites" within their counties. Sites chosen were those which were unlikely to be disturbed before mid-season the following year (e.g. unplowed oat stubble or oats underseeded with alfalfa), in order to maximize survival.

A follow-up Extension recovery program in 1972-75 revealed widespread establishment and dispersal (Fig. 1). In 1972-73, county agents were asked to collect and rear larvae in units patterned after one developed by Gruber, Rivet, and Pietro (1972). They then returned pupal cells to Michigan State University for analysis of the rate of parasitism. In 1974-75, agents were provided sweepnets and preservatives with which to collect three samples over a two week period near the time of peak cereal leaf beetle larval density. These samples were returned to Michigan State University for dissection and analysis. Results and an interpretation of findings were sent to each agent.

l ~ i c h i ~ a n Agricultural Experiment Station Article No. 7518. This work was sup- ported in part by cooperative agreements 12-14-100-10, 905 (33) and 12-13-100-11, 208 (33& E nt. Res. Div., Agr. Res. Sew., USDA.

Respectively, graduate research assistant, associate professor, and associate professor and extension project leader, Department of Entomology, Michigan State University, East Lansing, Michigan.


Fig. 1. Michigan townships in which T. julis has been released and/or recovered, 1966-75.

Table 1. Summary of county agent recovery program 1974-1975.

Counties/townships represented

Number of samples,

Total larvae, x l i

Larvae dissected, 2: di

Dissected larvae that were parasitized, 2 pi

Weighted percent parasitism, 1:

Table 2. Parasitism rates in samples from MSU Department of Entomology cereal leaf beetle state census, 1973-1975. C w -l

1973 1974 1975 0\

No. of Mean Mean % Date D D 1 Fields CLB's3 Parasitism2

Allegan County-Dorr Township 6/12 755 12 81 18 6/20 935 13 85 2 6/26 1050 13 24 37

Barry County-Thornapple Township

Emmet County-Resort Township

Jackson County-Pulaski Township

No. of Mean Mean % No. of Mean Mean% Date D-Dl Fields CLB's3 Parasitism2

-. Date D-Dl Fields CLB's3 Parasitism2

Lake County-Chase/Piora Townships 6/12 631 11 95 3 6/16 606 14 96 17

z 6/22 820 11 108 0 6/27 746 17 166 11 712 987 10 2 3 20 715 911 19 8 8 8 Oscoda County-West Comins Township g 6/14 598 5 310 17 6/16 594 3 19 26 6/19 674 4 860 38 6/25 701 7 11 6 1 6/27 824 5 33 36 717 937 7 11 57

Shiawassee County-New Haven Township 6/13 731 30 363 17 6/12 690 29 506 44 6 / 4 4 575 30 40 58 6/22 926 29 167 2 6/17 770 23** 389 38 6/12 690 30 20 52 6/28 1044 6' 7 67 6/25 886 29 708 31

713 1060 29 50 61

l~egrec-days, base 4 8 ' ~ , accumulated from 1 April, at nearest NOAA weather station. 2 2 (parasitism rate for field i ) /2 i, given there were any larvae found. 3 ~ e a n per 100 sweeps. 'Samples lost in lab accident. 4

4 "Samples lost in lab accident; figure known to underestimate mean CLB's/100 sweeps.


RESULTS

Table 1 summarizes the Extension Service sweepnet survey for 1974-75. Agents from 33 counties out of 68 in the lower peninsula sent in pupal cells in 1972. Eighteen of these showed julis establishment. In 1973, 17 counties returned pupal cells, all samples showing parasitism by julis. The apparent rate of parasitism varies considerably depending on the time of sampling, due to the bivoltine nature of julis. The overall parasitism rate, adjusted for sample size, from 1974 may be compared to the 1975 rate if one assumes samples were made near the same physiological times in both years. We attempted to justify this assumption by roughly bracketing the time of the peak larval density by sampling a week before peak, during peak, and a week after peak. Thus, northern county samples were taken starting about two weeks later than southern counties. Starting dates were set at approximately 600 accumulated degree-days (base 48°F) (Gage 1974).

During 1973-75, larvae collected from several selected Michigan townships as part of the annual Michigan State UniversiD Department of Entomology cereal leaf beetle census were also analyzed for parasitism rate. This cereal leaf beetle census was repeated three times during mid-season at seven to ten day intervals. Up to 30 oat fields per township were sampled, using 100 sweeps with a 15-inch net as a standard sample (Ruesink and Haynes, 1973). Table 2 lists results from the Department of Entomology cereal leaf beetle census from 1973-75. The average parasitism rate per field in these areas generally increased over the three year period.

DISCUSSION

The data obtained through Extension Service and Department of Entomology activities indicate a rising rate of parasitism by julis. The increase in 1975 over 1974 could be due to a widely observed concurrent decline in beetle density caused by unknown factors such as weather or insecticide use. However, the net result is still a higher proportion of the cereal leaf beetle larval population being parasitized.

The parasitism rates in Table 2 are generally hlgher than those in TabIe 1. We do not know the reason for this, but the townships selected by the Department of Entomology for their survey were chosen because they contained above average acreages of oats. This abundance of oats from one year to the next provides a greater chance for both julis and cereal leaf beetles to find their respective hosts in proximity to overwintering sites, and may be a contributing factor to the higher rates of parasitism observed.

Parasitism by julis is becoming substantial only four years after its first subcoloniza- tions. This parasitoid's capacity for survival and increase in typical farm systems, despite damaging operations such as plowing and diking which destroy many overwintering parasitoids, is a positive factor for cereal leaf beetle management progarns

LITERATURE CITED

Castro, T. R., R. F. Ruppel, and M. S. Gomuhski. 1965. Natural history of the cereal leaf beetle in Michigan. Mich. St. Univ. Agric. Ex%. Sta Quart. Bull. 47:623-53.

Dysart, R. J., H. L. Maltby, and M. H. Brunson. 1973. Larval parasites of Oulerna melanopus in Europe and their colonization in the United States. Entomophaga. 18: 133-67.

Gage, S. H. 1974. Ecological investigations on the cereal leaf beetle, Oulma melanopus (L.), and the principal larval parasite, Tetrastichus julis (Walker). Ph.D. Thesis, Mich. St. Univ., East Lansing, Michigan.

Gruber, R., E. Rivet and C. Pietro. 1972. A technique for obtaining cells of a soil-pupating insect, Oulema melanopus. J. Econ. EntomoI. 65:904-6.

Ruesink, W. G. and D. L. Haynes. 1973. Sweepnet sampling for the cereal leaf beetle, Oulema melanopus. Env. Entomol. 2: 161-172.

Stehr, F. W. 1970. Establishment in the United States of Tetrastichus julis, a larval parasite of the cereal leaf beetle. J. Econ. Entomol. 63:1968-9.


DESCRIPTION OF THE LARVA OF DIOEDUS PUNCTATUS LeCON'rE (COLEOPTERA: TENEBRIONIDAE)

Daniel K. Young1

ABSTRACT

Larvae of Dioedus punctatus LeConte have been collected from logs, probably Ulmus americana, in the red-rotten stage of decay and determined from reared adults. They are described and illustrated.

INTRODUCTION

Larvae of Dioedus punctatux LeConte were collected in Ingham County, Michigan, on 8 September, 1974. They were taken along with numerous adults from a dead log, probably UImus americana, in the red-rotten stage of decay.

Several larvae were placed in a covered plastic petri dish with small amounts of the moist decaying wood and maintained at room temperatures. Of these, three pupated and thus a positive association was established through determination of the ensuing adults.

DESCRIPTION OF LARVA

Larvae of punctams possess the general alleculid-tenebrionid conspectus, differing in details of the head and ninth tergite as described below.

Mature larvae attain lengths of 6-7 mm and widths of 0.5-0.75 mm, with the body cylindrical and subparallel to slightly broader posteriorly. The head and body are creamy white to light yellowish-brown in color, brown to piceous in areas of heavy sclerotization such as the tips of the mandibles and the urogomphi; vestiture is sparse with a few moderately elongate setae associated with the head and ninth tergite, the remainder of the body possesses but a few short setae.

Head prognathous, exserted from prothorax, as wide as or slightly wider than prothorax. Epicranial suture and ocelli lacking; rim of occipital foramen but slightly reinforced, not more deeply pigmented than remainder of head capsule. Antennae prominent, 3-segmented, with second segment bearing an ovate, disk-like sensorium which occupies '/z of its inner surface; f i s t antennal segment slightly longer than second, thud segment 0.4-0.5 times as long as second and bearing a few short setae subapically and one elongate apical seta. Mouthparts retracted, with stout asymmetrical mandibles, the right one bearing a well developed transversely serrulate molar area, the left with a prominent molar area which is produced into a blunt molar tooth; apices of mandibles tridentate with middle tooth slightly longer than outer teeth; left mandible usually closing over the right. Clypeus distinct; labrum movable, symmetrical, slightly longer than wide and bearing several elongate, anteriorly directed setae. Maxillae movable, composed of cardo, stipes, undivided pad-like maxillary articulating area, maxillary mala, and a 3-segmented palpus. Mala bearing numerous stout, spine-like setae along inner and apical margins. Labium with ligula absent; palpi 2-segmented. Mentum slightly longer than wide, broadest distally; submentum trapezoidal, widest proximally; gula elongate and narrow, its length 5 times its width. Ventral epicranial ridges and hypostomal rods absent. Epipharynx similar in form and structure to that of Phthora canalicollis Lewis (Hayashi, 1966; pl. 14, K); hypopharyngeal sclerome heavily sclerotized and transversely rectangular.

Legs short, 0.4-0.5 times as long as width of thorax, each bearing a short terminal claw; legs narrowly separated, with intercoxal distances of about one coxal diameter. Thoracic spiracle ovate, located near anterior margin of laterotergite (mesothoracic preepipleuron sensu St. George, 1924).

lDepartment of Entomology, Michigan State University, East Lansing, Michigan 48824.


Figs. 1 & 2. Larva of Dioedus punctatus LeConte, abdominal segments 8 & 9: Fig. dorsal aspect; Fig. 2, lateral aspect.


Abdomen cylindrical, moderately sclerotized, tergites 1-8 subequal in length and of similar shape. Ninth tergite (Figs. 1, 2) nearIy twice as long as eighth, bearing two heavily sclerotized, darkly pigmented, fixed urogomphi and terminating as excavated concavity caudodorsally; the caudoventrally directed recurved urogomphi arise dorsally, anterad of excavated area (Fig. 1). Ninth sternite reduced, visible laterally (Fig. 2) or ventrally, arising from distal margin of eighth sternite. Tenth segment reduced, visible as lightly sclerotized area surrounding anal opening immediately behind ninth sternite, frequently obscured due to intimate association of ninth sternite with ventral aspect of ninth tergite. Circular spiracles of similar size located laterally on laterotergites 1-8.

DISCUSSION

Dioedus punctatus, from the eastern states, and Phthom americana Horn, a West Coast species, comprise the North American representatives of the tribe Phrenapatini. Larvae of P. americana have recently been associated with adults by Lawrence and Doyen (Lawrence, in litt.), and will be described in a future paper dealing with tenebrionid larvae (Doyen, in litt.).

The only other source of information known to the author relative to phrenapatine larvae is that provided by Hayashi (1966). Larvae of Phthom canalicollis Lewis and Tagalus sp., both from Japan, are described and figured. The descriptions agree rather closely with that of D. punctatus not only in general conspectus, but also in such salient features as the apparent lack of an epicranial suture, lack of ocelli ("sometimes visible" in Tagalus), and in the unique and bizarre form and structure of the ninth tergite.

LITERATURE CITED

Hayashi, N. 1966. A contribution to the knowledge of the larvae of Tenebrionidae occurring in Japan (Coleoptera: Cucujoidea). Ins. Matsumurana, Suppl. 1: 141.

St. George, R. A. 1924. Studies on the larvae of North American beetles of the subfamily Tenebrioninae with a description of the larva and pupa of Merinus laevis (Olivier). Proc. United States Nat. Mus. 65: 1-22.

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