Revision of Palaearctic mountain midges (Diptera

Iowa State University

From the SelectedWorks of Gregory W. Courtney

January, 1994

Revision of Palaearctic mountain midges(Diptera: Deuterophlebiidae), with phylogeneticand biogeographic analyses of world speciesGregory W. Courtney, Smithsonian Institution

Available at: https://works.bepress.com/gregory_courtney/38/

http://www.iastate.edu

https://works.bepress.com/gregory_courtney/

https://works.bepress.com/gregory_courtney/38/

Revision of Palaearctic mountain midges (Diptera: Deuterophlebiidae), with phylogenetic and biogeographic analyses of world species G R EG O R Y W . Washington, D.C., U.S.A.

COU RTN EY Department of Entomology, Smithsonian Institution,

I

Abstract. The mountain midges (Diptera: Deuterophlebiidae) of the Palaearctic Region are revised to include eight species. Four new species are described: D. brachyrhina sp.nov., D.oporina sp.nov. and D. blepharir sp.nov. from the Himalayas (Assam and Sikkim), and D. bicarinara sp.nov. from southern Korea. A lectotype is designated for D.mirnbilis Edwards, and a key to adult males of all Palaearctic species is provided.

Larval, pupal and adult characters were used to reconstruct the phylogenetic and biogeographic relationships of world species of Deuterophlebiidae. Based on features of the adult male, the Himalayan species D. brachyrhina and D.oporina are considered the most primitive deuterophlebiids. The Nearctic species D.inyoensis is proposed as the sister group of the remaining species. Relationships among the latter are based primarily on larval and pupal characters; however, lack of information about the immature stages of several Palaearctic species contributes to a poorly resolved phylogeny. Several alternative hypotheses are presented and discussed. All phylogenetic alternatives suggest that the Nearctic fauna originated from at least two invasions of North America.

Introduction

The nematocerous family Deuterophlebiidae, commonly known as mountain midges, is one of the most specialized and distinctive of dipterous insects. Their immature stages are aquatic and adapted for life in swiftly flowing, often torrential, streams. Among the structural adaptations to these habitats are eversible, crochet-tipped larval prolegs and markedly streamlined pupae. Adults demonstrate equally aberrant features, including vestigial mouthparts and a longevity of only a few hours (Courtney, 1991a). Because of their structural and ecological specializations, and the controversy regarding their phylogenetic placement, deuterophlebiids have been of interest both to diptetists and aquatic entomologists. Unfortunately, the paucity of morphological and taxonomic studies has greatly hindered our ability to reconstruct the phylogenetic relationships of these unusual flies.

Corrcspondcncc: Dr G. W. Courtncy, Dcpartment of Entomo- logy. NHB 169, Smithsonian Institution. Washington, D.C. 20560, U.S.A.

The Deuterophlebiidae contains a single genus, Deutero- phlebia Edwards, with fourteen species, six from western North America (D.coloradensis Pennak, D.sha.sta Wirth, D.nielsoni Kennedy, D. inyoensis Kennedy, D.personata Courtney and D. vernalis Courtney) and eight from eastern and central Asia (Dmirabilis Edwards, D.nipponica Kita- kami, D.fyo.setiensi.s Kitakami, D.sajunica Jedlicka & Halgos, and four new species described in this paper). This paper provides diagnoses, descriptions and data on the distributions and biology of all Palaearctic species. Character systems introduced largely by Courtney (199oa) are a basis for keys to adult males of Palaearctic Deuterophlebiu.

Previous research on deuterophlebiid systematics has treated mostly familial relationships, e.g. Hennig, 1973; Rohdendorf, 1974; McAipine et al., 1981: Wood & Borkent, 1989: Courtney, l W b , 1991b) or accounts of new species (e.g. Kitakami, 1938b; Pennak, 1945; Wirth, 1951; Kennedy, 1958, 1960; Courtney, 1Wa). This study bridges the gap between these different aspects of systematics, by investigating the phylogenetic relationships of all species of the Deuterophlebiidae and by providing a basis for discussing the biogeographic relationships of the world mountain-midge fauna.

2 Gregory W . Courtney

Taxonomic history

The Deutemphlebiidae were originally described from two adult males collected in the mountains of Kashmir, India (Edwards, 1922). Shortly thereafter, Pulikovsky (1924) de .dbed Deuterophlebia larvae, pupae and adult females (dissected from pupae) from the Altai Mountains, Semipalatinsk (U.S.S.R.). Pulikovsky lacked adult males to compare with Edwards' description, so referred to the Altai specimens simply as Deuterophlebia sp. Pulikovsky provided a detailed description of larvae and brief descriptions of two pupal morphotypes, one of which remained unnamed for almost 60 years.

Brodsky & Brodsky (1926) and Brodsky (1930) provided other Deuterophlebia records from central Asia, with the latter including a detailed account of larval structure. Brodsky (Im. cit.) described two female morphotypes (dissected from pupae) but referred to most material as Dmirabifis. Brodsky (1980) provided a synthcsis of the geology, hydrology and faunistics of the Tien Shan in one of the most detailed reviews of mountain streams in central Asia. Data on deuterophlebiids, though based largely o n earlier papers (especially Brodsky, 1930), appeared in several sections of the book. Recent investigations on central Asian mountain midges include a description of the new species D.sajanica (Jedlicka & Halgos, 1981) and general reviews of the distribution of Palaearctic species (Jedlicka, 1Y80; Jedlicka & Halgos, 1985).

Subsequent to Edwards (1Y92). several years elapsed before further information about Himalayan deuterophlebiids was published. Larvae of 'an apparently undescribed species' were found in meltwater streams of the northern Pir Panjal Range and the south slope of the Himalaya (Singh, 1961), and additional observations were recorded by Mani (1962, 19681, Dubey & Kaul (1971) and Kaul (1979, 1986). Deuterophlebiids were also discovered in the Hindu Kush, northeast Afghanistan (Tokunaga, 1966). In spite of these records, the Himalayan fauna has remained poorly known.

The 1926 discovery of deuterophlebiid larvae, pupae and adult females at Kurama, a torrential stream near Kyoto, Japan (Kitakami, 1929), prompted considerable research on Japanese mountain midges. Detailed observations of the Kurama population resulted in one of the most comprehensive ecological studies to date (Yie, 1933). Additional sites were recorded between 1926 and 1937, but the lack of male specimens delayed species description (Kitakami, 1938b). Kitakami (1938a) reported the collection of males (pupae) and other life stages of KyQshO, and mentioned that the KyQshQ specimens were 'different than those . . . from HonsyQ' (page 215, translated from Japanese): however, the Kyashii and Honsyii specimens were both included under D.nipponica in Kitakami's revisionary study (1938b). The latter paper also contained a description of D.tyosenensis from Korea. Several years later, Kitakami (1954) collected pupae and more than 140 adults, including a large series of adult males, from the Hakugan River, North Korea. Kitakami gave mensural data for adults of D.mirabilis, D.nipponica, Pulikovsky's

Deuterophlebia sp. and the Hakugm species, the latter of which was simply listed as Deuterophlrhicl sp.

Detailed reviews of the history of research on Nearctic deuterophlebiids arc provided elsewhere (Kennedy, 1958; Courtney, 1989, 1Y9Oa).

. Methods

Material. This study was based on examination of larvae, pupae andlor adults of all known species of Deuterophlebia. Representatives of Palaearctic species were borrowed from, or are depbsited with, the following institutions (acronyms used throughout the text): BMNH, The Natural History Museum, London, England; CNC, Biosystematics Research Centre, Ottawa, Canada; GWC, collection of author; PZW, collection o f P. Zwick, Limnologische Flup-station des Max-Plank-lnstituts fur Lirnnologie, Schlitz, Germany; USNM. National Museum of Natural History, Washington. U.S.A.; ZSM, Zoologische Staats- sammlung, Mlinchen. Germany.

Specimen prepuru'ioti. Specimens were prepared accord- ing to procedures outlined in Courtney (I9Wa, b). Evalu- ation of characters on air-dried, pinned adults proved difficult because of distortion and shrinkage, and rcquircd rehydration in dilute (approximatcly 10%) potassium hydroxide (KOH). Material trcatcd in KOH was dissected and permanently mountcd in cithcr Eupord or Canada balsam.

Term7 fur strncfiires. Terms for structures arc bascrl primarilyon the Manual of Nearctic Diptcra [Tcskey (IYXI) for larvae, McAlpine (1981) f o r adults]. with deviations from this system, or terms applicablc specifically to thc Deuterophlebiidae, from Courtncy ( 199Oa. b). Thc rcadcr should consult Courtney (IWOii) for a dctailed charac- terization of the family. Abbreviations for adult features include f l - f 4 , flagellomere 1-4, and rl-15. tarsomere 1-5. The system for larval chaetotaxy is based on Courtney (19YOb) and includes the form of sensilla prl and sbt (thoracic segments, Figs 23, 24), pm (abdominal segments I-VII, Figs 23, 24), ar, mt and ap (abdominal segment VIII, Fig. 25). Descriptive format. Complete descriptions of known life

stages are given for new species; otherwise, I refer to the original description. The latter is supplemented by certain taxonomic information not included in the original reference (e.g. pupal microsculpture). When applicable. sample sizes are indicated prior to each data set, with values presented as a range followed in parentheses by a mean and standard deviation. Vaiues used for adult antenna1 article ratios are relative to the shortest article, usually the pedicel, and represent the mean ratio of all measured individuals. Leg articles were measured along the dorsal margin; article percentages are mean values for article 1ength:total leg length. Abbreviations for immature life- stages are L, larvae; P, pupae. Taxa are arranged in a phyletic sequence (i.e. plesiomorphic species listed prior to apomorphic species), as determined by the cladistic analysis.

.

Key to adult males of Palaeatctic Deuteroph/ebia*

1 Antennal lciigth 7 mm or less,'length off4 approximately 90 times length of pcdiccl ............................... .2

Antenna1 length 8mm or mom. f 4 at least 120 times Iongcr than pedicel. ....................................... .3

2(1) Tibia and tarsus without capitate rnacrotrichia (Fig: 7):

-

psteromedial margin of dorsal plate with deep excision (Fig. 11); median clypeal bbe indistinct (Fig. 1); postgcna and oral region glabrous (Fig. 3); antennal j7-3 with approximately 5, 20 and 30 digitifom sensilla, respectively, on anteroventral margin (Fig. 5); hind rl thickened, its width greater than width of basal tibia (Fig. 6 ) . ............

Tibia and tarsus with capitate macrotrichia (as in Fig. 8); posterior margin of dorsal plate truncate (Fig. 12); median dypeal lobe distinct, truncate (Figs 2, 4); postgena and oral rcgion with microtrichia (Fig. 4); antennal f l -3 each with fewer than 10 digitiform sensilla on anteroventral margin: hind rl not thickened, its width subequal to width of t2-5 and lcss than width of basal tibia .......... D.oporina sp.n.

.................................. D. brachyrhina spa.

3(1) Postgcna with microtrichia, subgcnal suture indistinct (as in Fig. 4) ............................................ .4

Postgcna glabrous, subgenal suture usually distinct (as in Fig. 3) ............................................ .5

4(3) Compound eyes with microtrichia between facets. .........

- Compount cyes glabrous .................. D.mirabilis Edwards

5(3) Antcnnalfl bulbous distally, with about 25 digitiform scnsilla on antcrovcntral margin; mcdian clypcal lobe indistinct ....

Antcnnalfl not bulbous distally, with 10 or fcwcr digitiform scnsilla on antcrovcntral marpin; median clypcal lobc short. convex .......................... D.nipponica Kitakami

-

..................................... D. blepharis spa.

. . . . . . . . . D. birahata sp.n., D.sajanica Jcdlicka & Halgos -

Deuterophlebia brachyrhina spnov. (Figs 1,3,5-7, 9, 11.26-29)

Diagnosis. Male similar to D.oporina, but tibia and tarsus without capitate macrotrichia; median clypeal lobe indistinct; hind tl thickened, its width greater than that of tibia base; posterior margin of dorsal plate with deep median notch.

Larva. Unknown. Pupa. Unknown. Adult 8. Median clypeal lobe indistinct (Fig. l), with

about 25 setiform sensilla; lateral lobes poorly developed, densely set with microtrichia. Anterior tentorial pits approximately 6Opm posterior to clypeal incision. Com- pound eyes glabrous. Subgenal suture distinct, extending from lateral clypeal lobe to posterior tentorial pit; postgena and oral region glabrous, except for a few microtrichia and setifom sensilla near anterior clypeal margin (Fig. 3):

* Males unknown for D.ryosenensis; malcs of D.nipponica unavailable For examination. so its placement in the key is tentative and based primarily on Kitakami (1938b).

mental tooth acute. Antennal f1,p andf3 slightly bulbous distally, with a cluster of approximately 5 , 20 and 30 digitiform sensilla, respectively, on anteroventral margin (Fig. 5) . Coxa, trochanter and femur without macrotrichia, but with dense microtrichia. Fore coxa markedly short, subspherical. Fore, mid and hind tibia with simple macrotrichia, without capitate macrotrichia (Fig. 7). Hind tibia glabrous posteriorly, with a patch of elongate, apparently stiffened macrotrichia near tibio-tanal joint (Fig. 6). Hind tl thickened, its width greater than that of tibia1 base; tl glabrous posteriorly. All tarsi without capitate macrotrichia.

Terminalia (Figs 9, 11). Dorsal plate with deep pos- teromedial notch; posterior margin with setiform and chaetiform sensilla; posterolateral angle acute; lateral margins parallel-sided. Gonostylw length subequal to length of dorsal plate, greater than length of aedeagal sheath. Aedeagus internally smooth.

Measurements ( N = 6). Total body length 1.9-2.2mm (2.04 f 0.09). Antennal length 5.6-6.9mm (6.13 2 0.02); article length ratio 4 1:2.5:2:2:91. Wing length 4.0-4.6 mm (4.23 2 0.21). width 2.0-2.3 mm (2.12 2 0.13). Leg article percentages: foreleg: 4:424:39:16:4:4:3:3; midleg: 8:5:27:33:12:4:4:3:3; hindleg: 8:5:33:33:12:3:3:2:3.

Adult 9. Unknown. Type material. Holotype d: INDIA: Assam, Nyuk-

madong; NEFA, Kameng Fr. Div., 7UlOft [-27"26'N 92%8'E], 23.iv. 1961 (Schrnid) (CNC); specimen on slide in Euparal.

Paratypes. Same data as holotype, 196 [7 on slides, 12 pinned]; 18.iv.l961,7~3 [pinned]: INDIA: Assam, Kelong, NEFA, Kameng Fr. Div., 6OOOft, 26.ix.1961 (Schrnid). 1d [on slide]; deposited in CNC. USNM and GWC.

Derivation of specific epithet. From the Greek for 'short snout', in reference to the indistinct median clypeal lobe.

Distriburian. Known only from the Himalayas of northern Assam (Fig. 28).

Bionomics. Collection records suggest that D. brachyr- hinu is active during the spring, and is isolated temporally from other species (cf. D.oporina and Dmirabilis). How- ever, collection of a single specimen from Kelong on 26.ix.61 may refute this hypothesis and indicate that D.brachyrhina passes through more than one annual generation.

huterophlebia oporha sp.nov. [Figs 2,4, 10, 12, 26-29)

Diagnosis. Male similar to D. brachyrhina but tibia and tarsus with capitate macrotrichia; apex of median clypeal lobe truncate; mental tooth bilobed; posterior margin of dorsal plate truncate. Female distinguished from other species by two prominent setiform sensilla on the dorsal margin of the scape and by the markedly elongate basal cercat article.

Larva. Unknown. Pupa. Unknown. Adult d . Median clypeal lobe pronounced, subquadrate,

with truncate apex (Figs 2,4); clypeus medially with about


Rgs 1-8. Deuterophlebia adult males: 1, D.brachyrhina, cranium do& view; 2, D.opOha, cranium, dorsal view; 3 , D.brachyrhina, oral re@on, ventral View; 4, D.oponna, oral region, ventral view; 5, D.bmachyrhina, antenna1 flagellomeres 1-3, anterodorsal view: 6, D.brachyrhina, tibio-tarsal joint, posterior view; 7. D.brachyrhina, distal end of foretibia; 8, D.personata, distal end of foretibia. Abbreviations: clm, median clypeal lobe; pg, postgena. Scale bars=O.lmm (Figs 1, 2), 20pm (Figs 3-5, 7, 8), lOpm (Fig. 6).

Revision of Pulaeurcfic mountain midges 5

15 setiform sensilla; lateral lobes slightly developed, with sparse microtrichia. Anterior tentorial pits approximately 4Opm posterior to clypeal incision. Compound eyes glabrous. Subgenal suture indistinct; postgena and oral region with microtrichia; mental tooth bilobed (Fig. 4). Each of antenna1 fl, f2 and f3 distally with about six digitiform sensilla borne on anteroventral tubercle. Coxa, trochanter and femur without macrotrichia; fore and mid tibia distally with abundant capitate macrotrichia, especially on ventral margin. Hind tibia with capitate macrotrichia on ventral margin. All tarsi with capitate macrotrichia on ventral margin.

Terminalia (Figs 10, 12). Dorsal plate subquadrate; posterior margin truncate, with setiform sensilla. Gonostylus length subequal to length of both dorsal plate and aedeagal sheath. Aedeagus internally smooth.

Measurements (N= 7). Total body length 1.9-2.2mm (2.09 kO.11). Antennal length 5.5-6.0mm (5.74 C 0.25); article length ratio 2.5:1:2.5:1.5:1.5:87. Wing length 3.7-4.0mm (3.8520.11), width 1.6-1.9mm (1.772 0.10). Leg article percentages: foreleg: 7:423:36: 16:5:43:3; midleg: 8:4:26:3O: 17:5:4:3:3; hindleg: 8:4:31:35:9:4:4:3:3.

Adult 9. Similar to male. Median clypeal lobe pronounced, approximately 45 pm long, with about ten setiform sensilla; clypeus laterally without pronounced lobes or dense microtrichia. Anterior tentorial pits approximately 12 pm posterior to clypeal incision. Antenna: dorsal margin of scape with two prominent setifom sensilla; pedicel with about four setifom sensilla; fl-3 distally with two, three and four digitiform sensilla, respectively; f4 with five setiform sensilla and three digitifom sensilla.

Terminalia. Anterior bridge of genital fork posteriorly convex and extending less than half the distance to accessory gland opening. Cercus with length of basal article at least twice the length of distal article; articles distinguishable by constriction on all but medial margin.

Measurements (N = 2). Total body length 1.4-2.1 mm. Antennal length 0.33 mm; article length ratio 2.5:1:3.5: 1.5:1.5:1. Wing length 3.3-3.4mm, width 1.3-1.4mm. Leg article percentages: foreleg: 7:4:23:41:6:4:4:4:9; midleg: 8:4:23:39:6:4:44:8; hindleg: 9:4:25:384:44:4:10. Length of tllt5: foreleg: 0.65-0.70; midleg: 0.60-0.72; hindleg: 0.45.

Type material. Holotype d: INDIA: Assam, Phutmg, NEFA, Kameng Fr. Div., 7300ft [=27"14'N 92'14'E], 1-2.x.1%1 (.%hid) (CNC); specimen on slide in Euparal.

AMotype 0 : Same data as holotype (CNC); specimen on slide in Euparal.

Paratypes: Same data as holotype, 736 [12 on slides, 61 pinned] 39 [l on slide, 2 pinned]; INDIA: Sikkim, Ramtang, 5780ft, 13.x.1959 (Schmid), 9 6 [l on slide, 8 pinned]; Sikkim, Chunjom, 68OOft, 12.x.1959 (Schrnid), 5d [l on slide, 4 pinned]; deposited in CNC, USNM and GWC.

Derivation of specific epiher. From the Greek for 'autumnal', in reference to the Season when this species has been collected.

Distribution. Known only from the Himalayas of Sikkim and northern Assam (Fig. 28). The species presumably occurs in adjacent regions of Bhutan and Nepal.

Bionomics. Collection records suggest that D. oporina is a 'fall' species that is isolated temporally from D.brachyrhina but not D.mimbifis. The paratypes from Ramtang, Sikkim (13.x.59) were from samples that contained both species; it is unknown if these were captured from the same swarms.

Deuteropblebia blepbaris sp.nov. (Figs 26-29)

Diagnosis. Male and female similar to Dmirabilis but differentiated by the presence of microtrichia between the eye facets.

h r v a . Unknown. Pupa. Unknown. Adult 6 . Median clypeal lobe convex, approximately

4Qpm in length, with about 20 setiform sensilla; lateral lobes pronounced, densely set with microtrichia. Anterior tentorial pits approximately 7 0 p posterior to clypeal incision. Compound eyes with microtrichia between eye facets. Subgenal suture indistinct; postgena and oral region with microtrichia; mental tooth acute. Antennalfl distally with sma11 anteroventral tubercle bearing nine digitiform sensilla; each of f z and fJ distally with anteroventral tubercle' bearing about five digitifom sensilla. Coxa, trochanter and femur sparsely set with macrotrichia; fore and middle tibia with simple macrotrichia, distally with abundant capitate macrotrichia, especially on ventral margin. Hind tibia with simple macrotrichia on dorsal margin; capitate macrotrichia abundant on ventral margin. All tarsi with capitate macrotrichia on ventral margin.

Terminalia. Dorsal plate with shallowly emarginate posterior margin, medial notch, and several chaetiform and setifom sensilla; posterolateral angle blunt; lateral margins parallel-sided. Gonostylus length greater than length of dorsal plate or aedeagal sheath. Aedeagus internally smooth.

Measurements (N= 3). Total body length 2.0-2.9mm (2.37 2 0.48). Antennal length 12.4-12.7mm (12.51 2 0.20); article length ratio 3:1:3.5:1.5:1.5:175. Wing length 5.2-5.3mm (5.27kOO.O6), width 2.3-2.4mm (2.372 0.M). Leg article percentages: foreleg: 5:3:25:35:17:5:442; midleg: 8:426:29:185:4:3:3; hindleg: 7:433:36:7:443:2.

Adult Q (description based solely on a l l o w ) . Similar to male. Median clypeal lobe slightly convex, 10 pn long, with seven setiform sensilla; clypeus laterally without pronounced lobes or dense mimtrichia. Anterior tentorial pits approximately 30 pn posterior to clypeal incision. Antennal fl-4 narrow basally, expanded distally; each of jl-3 distally with slight anteroventral prominence bearing two, three, and three digitiform sensilla, respectively; f4 with four setiform sensilla and three digitifom sensilla.

Terminalia. Cercus with length of basal article greater than length of distal article; articles distinguishable by constriction of all but medial margin.

Measurements. Total body length 2.3 mm. Antennal length 0.50 mm; article length ratio 2.5:1:3: 1.5:2:3. Wing length 4.8 mm, width 2.0mm. Leg article percentages: foreleg: 7:3:23:44:7:3:3:3:9; midleg: 6:4:26:4O:6:3:3:3: 10;

6 Gregory W. Courmey

9-14. Dcutatophlebia: 9, D.brachyrhina, male tenninalia, do& view; 10, D.oporina, male tenninalia, dorsal View; 11, D.brachyrhim, dorsal plate of male; 12, D.oporina, dorsal plate of male; 13, D.mirabilis, dorsal plate of male; 14. D.bicarinrrta, pupal microsculpture. Abbreviations: Be, aedeagus: dp. dorsal plate; gs. gonostylus: p. gonocoxite. Scale bars=;Ulym (Fig. 9), 20pm (Figs 11, 13), lOpm (Figs 10, 12), 5pm (Fig. 14).

. , . . '

. . . -. ,---

r

Figs 15-17. Deurerophlebia female genitalia, internal: 15, D. vernalis, lateral view (semi-diagrammatic): 16, D.r*ernalis, dorsal view of genital fork; 17, D.iqyoensis, dorsal view of genital fork. Abbreviations: ag, accessory gland (base); gf, genital fork arms; gfb, anterior bridge of genital fork; gfp, posterior lobes genital fork: sVI11, stcrnite VIII; tVIII-tX, tcrgites VIII-X; asterisk, basal opening of accessory gland. Scale bar = 0.1 mm. (Adapted from Courtney. 1990a).

a * -. $. 3

a

a s.


hindleg: 8:4:28:39:4:3:3:3:8. Length of t11tS: foreleg: 0.71; midleg: 0.62; hindleg: 0.58.

Type material. Holotype 6 : INDIA: Sikkim, Yangsap [=2728'N SsOw'E], 9.x. 1959 (Schmid) (CNC); specimen on two slides in Canada balsam.

Alfotype 0 . Same data as holotype (CNC); specimen on slide in Canada balsam.

Puraqpes. Same data as holotype, 2 6 [pinned]; deposited in CNC and USNM.

Derivation of specific epithet. From the Greek for 'eyelash', in reference to the presence of microtrichia between the eye facets.

Distribution. Known only from Sikkim (Fig. 28). Bionomics. Deuterophlebia blepharis apparently co-

occurs with at least one species, D.mirabilis. The type series was from a sample that included seven adult (66, I ? ) D.mirabifis. I t is unknown if all were collected from the same habitat or swarm.

Deuterophlebia mlr8biliS Edwards (Figs 13, 26-29)

Deuterophlebia mirabilis Edwards, 1922: 382. Lectotype d (present designation), INDIA, Kashmir, Srinagar (BMNH) [examined]

Diagnosis. Male similar to D. blephuris and D.irtyoerisis; distinguished from the former by the absence of microtrichia between the eye facets, and from D.iriyoensis by the shape of the dorsal plate (in D.mirubifis, the posterior margin bears a median notch) and the presence of anteroventral tubercles on antenna1 fl-3 (absent in D.inyoensis). Female similar to D.blepharis and D.inyoensis: separable from the former by the absence of microtrichia between the eye facets, and from D.inyoensi.7 by the relatively short hind tl (length of tllt5 less than 0.6); distinguished from other species (e.g. D.shastu) by the presence of microtrichia on the postgend and oral region.

Larva. Unknown (not definitively associated with adult). Pupa. Unknown (not definitively associated with adult). Adulr d . Median clypeal lobe convex apically, of vari-

able length (30-60pm), bearing about 20 setiform sensilla; lateral lobes pronounced, densely set with microtrichia. Anterior tentorial pits approximately 60 pm posterior to clypeal incision. Compound eyes glabrous. Subgenal suture indistinct in most specimens; postgena and oral region with microtrichia; mental tooth acute. Antennal fl distally with small anteroventral tubercle bearing 4-8 digitifom sensilla; each of j2 and j3 distally with anteroventral tubercle bearing 5-8 digitifom sensilla. Coxa, trochanter and femur with sparse macrotrichia; fore and mid tibia sparsely set with simple macrotrichia, distally with abundant capitate macrotrichia, especially on ventral margin. Hind tibia sparsely set with simple macrotrichia on dorsal margin; capitate macrotrichia abundant on ventral margin. All tarsi ventrally with capitate macrotrichia.

Terminalia. Dorsal plate (Fig. 13) shallowly emarginate posteriorly, notched mediaiy , bearing chaetifom and setiforrn sensilla; posterolateral angle acute; lateral margins

parallcl-sided to slightly concave. Gonostylus length less than length of dorsal plate, greater than length of aedeagal sheath. Aedeagus internally smooth.

Measurements (N = 11). Total body length 2.1-3.1 rnm (2.66 20.28). Antennallength9.6-13.4mm (11.65 1.11); article length ratio 3:1:4:2:2:170. Wing length 4.2-6.0 mm (5.11 2 0.55), width 2.0-2.9 mm (2.39 k0.29). Leg article percentages: foreleg: 6:3:24:36: 17:5:4:3:2; midleg: 8:4:26: 30:17:5:4:3:2; hindleg: 8:4:31:36:8:4:4:3:2.

Adult 9 . Similar to male. Median clypeal lobe convex, approximately 4Oym long. with about ten setiform sensilla; clypeus larerally without pronounced lobes or dense microtrichia. Anterior. tentorial pits approximately 45 pm posterior to clypeal incision. Antennal j7-4 narrow basally, expanded distally; f2-3 distally with slight an- troventral prominence bearing 0-2. 3-5 and 4- 10 digitiform sensilla, respectively; f4 with four setiform sensilla and five digitifom sensilla.

Terminalia. Anterior bridge of genital fork extended posteriorly to more than half the distance to accessory gland opening. Cercus with length of basal article less than length of distal article; articles indisrinctly fused to each other, separated only by shallow. ventrolateral notch.

Measurements (N = 4). Total body length 2.4-2.6mm (2.47 z 0.02). Antenna1 length 0.50-O.54mm (0.52 2 0.02); article length ratio 2:1:3:1.5:2: 1.5. Wing length 5.2- 5.6mm (5.40 2 0.20), width 2.0-2.2 mm (2.12 t 0.12). Leg article percentages: foreleg: h:3:22:42:6:3:4:3: 11: midleg: 7:3:22:41:6:3:3:3: 1 1; hindleg: 8:3:26:40:4:3:3:3: 10. Length of tl/t5: foreleg: 0.58-0.63 (0.60 2 0.02); midleg: 0.56-0.67 (0.60 2 0.06): hindleg: 0.35-0.47 (0.40 -C 0.06).

Type muterial. Lertotype d: INDIA: Kashmir, Srinagar, 1 1 - 12,000 f t , 192 1 (Mitchell) (BMN H).

Lectorype designutiotr. The type series of D.mirabilis contained two adult males from Kashmir, both designated as cotypes by Edwards (1922). One specimen was mounted on a slide, while the other was stored in formalin. The slide-mounted specimen has apparently been lost (B. C. Townsend, The Natural History Museum, London, 1988, personal communication). I hereby designate as lectotype the remaining adult; this specimen evidently dried out at some time (R. W. Crosskey, The Natural History Museum, London, 1984, personal communication), was then rehydrated and placed in alcohol. The specimen has been removed from alcohol, dissected, and slide-mounted in Canada balsam [two slides]. Each slide carries the following labels: 'Kashmir, Srinagar, 11- 12,Ix)(!ft, 1921, coll. F.J. Mitchell, 414.' and 'LECTOTYPE: designated by G. W. Courtney 1993'.

Other material examined. CHINA: 'Uriimqui, Tienshan- Geb., 22000m, Antg. Oktober 1984, Auf Schnee am Bachrand' (Liibemu-Nestle) 4 6 39 (ZSM), 1d 1P (GWC): PAKISTAN: Chitral, Besti, 20.x. 1954 (Schmid) 2d (CNC), 2 8 (USNM); NWFP, Gittidas, 14.vii.1953 (Schmid) 26 (USNM); INDIA: Sikkim, Chachu, 29.vi. 1959 (Schmid), 4 6 (CNC); Chunjom, Sikkim, 68OOft, 12.x. 1959 (Schmid), 38 (CNC), lc3 (USNM); Sikkim, Copetang, 5780ft, 10.x.1959 (Schmid), 12d 10 (CNC): Sikkim.

6

10 Gregory W. Courtney

is also known from the Hindu Kush of n0rthea.t Afghanistan (Tokunaga, 1%), and may occur in parts of the Altai (Pulikovsky, 1924; Brodsky, 1930 [see below]).

Bionomics. Moni (1962) proposed that certain Himalayan populations have multiple generations, and collection records from Ramtang, Sikkim, suggest that D.mirabilis is bivoltine, with distinct spring and autumn generations. Such a pattern would be unusual for deuterophlebiids (Courtney, 1991a). Alternatively, two species may be present; however, adult characters currently provide no evidence of more than one species.

Remarks. Brodsky (1930) provided relatively detailed descriptions of larvae, pupae and pharate adults (dissected from pupae) from the Tien Shan, all of which were identified as D.mirabilis. Because I was unable to examine this material, and because Brodsky apparently collected two different species (see his discussion on pp. 313-314), I refer to this material as Deuterophlebia sp. Most of Brodsky’s specimens may indeed be D.mirabilis; however, until larvae, pupae and adult females are definitively associated with the Dmirabilis type series, or at least with D-mirabilis males from nearer the type locality, I remain non-committal about this material.

Deutetophlebia nipponica Kitakami (Figs 22, 26-29)

Deuterophlebia nipponica Kitakami, 1938b: 488. Syntypes (36, 574, ‘numerous’ pupae, larvae) JAPAN, Honshlj and KyQshD (depository unknown, syntypes apparently lost)

Diagnosis. Pupa similar to several other species ( e . g . D.coloradensis), but recognized by the presence of three pairs of raised, darkly sclerotized mesonotal bands, and one pair of similar bands on each of abdominal segments I-VII. Deuterophlebia bicarinura sp.nov. (see below) has similar abdominal sculpturing, but only on segments I and 11; these species are further separated on the basis of mesothoracic spines.

Larva. See Kitakami (1938b). Only one damaged larva was available for examination and it lacked most sensilla; therefore, I could not evaluate chaetotaxic characters.

Pupa (Fig. 22). See Kitakarni (1938b). Only one damaged female pupa was available for examination and it had been dissected: the specimen lacked its thorax, so I can provide no details about the gill or mesonotum. Because microsculptural features of the pupa were evaluated, a figure and partial description are provided. Abdominal segments I-VII each with transverse, raised, darkly sclerotized bands on either side of midline, and small, raised, darkly sclerotized dots farther laterad. Micro- sculpture: abdominal sutures with secondary ridges; tergites without microtubercles; microtrichia apparently absent.

Adult d . See Kitakami (1938b). Adulr 9 . See Kitakami (1938b). Postgenal vestiture

Material examined. JAPAN: Kurama River near Kyoto, absent.

23.x.1932 (S.Kirakarni?) [lL, lP] (USNM).

Distribution. Knowt, only from Japan (Honshii and KyDshQ) (Fig. 28).

Bionomics. Yie (1933) gave a detailed account of the bionomics of D.nipponica. Populations near Kyoto (if they still exist!) are apparently multivoltine and active through much of the year. Pupal collections consisting predominantly of females (Yie, 1933; Kitakami, 1938b) suggest that the species is parthenogenetic across much of its range.

Deutwophlebia tyosenensis Kltakami (Figs 20, 26-29)

Deuterophlebia tyosenensis Kitakami, 1938b: 501. Syntypes (7 pupae, 8 larvae) TyBsen (NORTH KOREA), Mt Ken@ and Kankydnanda (depository unknown, syntypes apparently lost).

Diagnosis. Pupa readily separated from most species by the presence of two spines on each side of the mesothorax; similar to D. sp. C from Nepal (see below), but recognized by differences in size and abdominal microsculpture.

Larva. See Kitakami (1938b). Pupa (Fig. 20). See Kitakami (1938b). Because only

one damaged female pupa was available for examination, I refer to Kitakami (1938b) for information about several characters. For reasons given under D.riipponica, a figure and partial description of the pupa are provided. Micro- sculpture: abdominal sutures without secondary ridges: tergites without microtubercles: microtrichia apparently absent .

Adult. Unknown. Material exurnitled. One pupa: ‘Kurama River, Kyoto,

JAPAN, 23.x.lY32’ (USNM); the specimen is presumably not from Japan, but from Korea (see Kitakami, I93Rb).

Distribution. Known from central and northern Korea (Kitakami, 1938b) (Fig. 28).

Bionomics. Unknown.

Deuferophlebia bicarlnata sp.nov. (Figs 14, 18, 23, 26-29)

Diagnosis. Larva similar to D.sajanica, but differing in dorsal crania! colour (dypeal lobe pale distally) and chaetotaxy (sensillum p m branched). Pupa similar to D.sajanica, but recognized by transverse, raised, darkly sclerotized bands on abdominal tergites I-11, and by abdominal microsculpture (i.e. microtubercles concentrated near sutures). Adult male separable from most species by indistinct median clypeal lobe; adults indistinguishable from D.sajanica.

Larva (Fig. 23). Dorsal colour: body pale to light brown. Dorsal cranium (instar IV) pale laterally; broad, dark medial band extending from prothorax to clypeus; clypeal lobe dark at base, pale distally. Distal antenna1 article pale, without digitiform microtrichia; pit-like sensillum on basal third of ventral branch. Chaetotaxy: prl. pm (Fig. 23), at and mt (as in Fig. 25b) branched setiforrn sensilla;

Reiiyioiz of Pulueurctic moccirrain midges 1 1

Figs 23-24. Deuterophlebiu, dorsal view of instar IV larval head, thorax, and abdominal segment I: 23, D.bicaririuta; 24, D . sp. B. Abbrcviations: pm. medial pseudopodal xnsilla; prl, prelaterotergal sensilla; sbt. subtergal sensilla. Scale bars = 0.5 mm.


sbt (meso- and metathorax) simple setiform scnsilla (Fig. 23); ap with two clusters of digitiform sensilla (as in Fig. 2%).

Measurements, instar IV (N= 10). Total length 3.0-3.8 mm (3.41 2 0.28), cranial width 0.54-0.62 mm (0.58 2 0.02): proleg with 7-12 crochet rows; antennal ratio 1:2: 1. P u p (Fig. 18). Male antennal sheath coiled around

periphery of ventral abdomen twice, terminating near head. Gill base thick, short, nearly twice as long as wide; gill with thick, posteriorly projected filament and three elongate filaments, two of the latter three (posterior-most pair) typically borne on a common base. Spine-bearing tubercle o n dorsolateral margin of mesonoturn, above and slightly posterior to gill base: spine finely tapered and elongate, its apex extending anteriorly to beyond outer margin of gill. Mesonotal bands transverse, raised, darkly sclerotized, between thoracic tubercle and ecdysial line; anterodorsal mesonoturn with 1-2 small, raised, darkly sclcrotized dots on either side of ecdysial linc. Metanotal bands similar to mesonotal bands, but less darkly sclerotized and shorter. Abdominal segments I and I1 each with transverse, raised, darkly sclerotized band on either side of midline. Abdominal segments I, 11, VI and V11 laterally with 7-9, 10-15, 8-10 and 8-12 thorn-like spines, respectively.

Microsculpture. Mesonotum moderately rugose; meso- and nietanotum densely set with mimtubercles. but without

associated microtrichia. Abdominal sutures without scc- ondary ridges: tergites with microtubercles mostly near sutures; microtrichia distributed generally over tergitc surface, not associated with microtubercles (Fig. 14).

Measurements. Male pupae (N = 4). Total length 2.2-2.6mm (2.39*0.15), width 1.5-1.9mm (1.6920.16). Female pupae ( N = 5 ) . Total length 2.0-2.4mm (2.25 -f 0.16), width 1.5-1.8mm (1.6520.13).

Adult d . Median clypeal lobe indistinct, slightly convex, with 15-20 setiform sensilla: lateral lobes pronounced, densely set with microtrichia. Anterior tentorial pits approximately 20 pn posterior to clypeal incision. Com- pound eyes glabrous. Subgenal suture distinct; postgena and oral region glabrous, except for a few microtrichia and setiform sensilla near anterior clypeal margin; mental tooth acute. Antenna1 fl slightly bulbous distally, with about 25 digitiform sensilla: each off2 and f3 distally with anteroventral tubercle bearing about 10 digitiform sensilla. Coxa, trochanter and femur with simple macrotrichia: fore and mid tibia sparsely set with simple macrotrichia, distally with abundant capitate macrotrichia, especially on ventral margin. Hind tibia dcnscly set with simple macrotrichia on dorsal margin; capitate macrotrichia abundant on ventral margin. sparsely distributed o n distal part of dorsal margin. All tarsi ventrally with capitate macrotrichia.

Teminalia. Dorsal platc concavc latcrally; posterior margin emarginatc. notched mcdially; with sctiform and

Fig. 25. Deurerophlebiu, instar IV larvae. dorsal view of abdominal segments VII and VIII: a. D.invoensis. b, D. vcmulis. Abbreviations: ap, apical sensilla; at, anterotergal sensilla; mt. mcdiotergal scnsilla. Scale bar = 0.25 mm. (Adaptcd from Courtney. 1Yx)a).

Revision of Pahearctic mountain midges 13

chaetiform sensilla; posterolateral angle acute. Gonostylus length greater than length of dorsal plate or aedeagal sheath. Aedeagus internally smooth.

Measurements (based on a single pharate adult dissected from pupa). Antennal length 9.8mm; article length ratio 2.5:1:3.5: 1.5:1.5:155.

Adult 9 . Similar to male. Median clypeal lobe broadly truncate, approximately 20 pm long, with 10- 15 setifom sensilla; clypeus laterally without pronounced lobes or dense microtrichia. Anterior tentorial pits approximately 30 pm posterior to clypeal incision. Antennal fI -4 narrow basally, expanded distally; each of f l - 3 distally with slight anteroventral prominence bearing, respectively, two, four and two digitiform sensilla; f 4 with four setiform sensilla and four digitifom sensilla.

Terminalia. Anterior bridge of genital fork posteriorly acute, extending more than half the distance to accessory gland opening. Cercus with length of basal article less than length of distal article; articles indistinctly fused, separated only by indistinct ventrolatenl notch.

Measurements (based on two phante adults dissected from pupae): Antennal length 0.32-0.34 mm; article length ratio 3: 1:2.5:1:1:1. Length of t l / t S : foreleg: 0.56-0.58; midleg: 0.52-0.60; hindleg: 0.31-0.40.

Type material. Holotype d (dissected from pupa): SOUTH KOREA: Samjong (620 m). Kayasan, Hapchon- gun, Kyongsangnamdo [=359)0" 128"00'E], 25-26.v. 1983 ( S . Uchida) (USNM); specimen on slide in Canada balsam.

AIIotype P [dissected from pupa]. Same data as holotype (USNM); specimen on slide in Canada balsam.

Paratype.y. Same data as holotype. 10 larvae [4 on slides, 6 in EtOH], 7 pupae [4 on slides, 3 in EtOH]; deposited in CNC, USNM, GWC.

Derivation of specific epithet. From the Latin for ridge or keel, in reference to the two pairs of transverse, raised, darkly sclerotized bands on abdominal tergites 1-11 of the pupa.

Distribution. Known only from South Korea (Fig. 28). Bionomics. Unknown.

Deuteropblebia sajanica Jedlicka & Halgos (Figs 19, 26-29)

Deuterophlebia sp. Pulikovsky, 1924: 45. Deuterophlebia sajanica Jedlicka & Halgos, 1981: 973.

Holotype d . MONGOLIA, Chbvsgbl ajmak, Ich- Choro-go1 (SNM) [not examined]

Diagnosis. Larva similar to D . bicarinata but differing in dorsal cranial colour (clypeal lobe dark) and chaetotaxy (sensilla p m setiform). Pupa similar to D.bicarinata, but differing in the lack of transverse, darkly sclerotized bands on abdominal tergites 1-11 and in abdominal microsculp ture (i.e. microtubercles distributed generally over tergite surface). Adult male separable from most species by indistinct median clypeal lobe: adults indistinguishable from D. bicarinata .

Larva. See Jedlicka & Halgos (1981). Dorsal colour:

body pale to light brown. Dorsal cranium (instar IV) pale laterally; broad, dark, medial band extending from prothorax to clypeus; clypeal lobe dark. Distal antennal article pale, without digitiform microtrichia; pit-like sensillum on basal third of ventral branch. Chaetotaxy: prl, pm (as in Fig. 23). at and mt (as in Fig. 25b) branched setiform sensilla; sbt (meso- and metathorax) simple setiform sensilla (as in Fig. 23); ap with two clusters of digitiform sensilla (as in Fig. 25a).

Measurements, instar IV ( N = 3 ) . Total length 3.0- 4.7 mm (3.61 2 O.Y7), cranial width; 0.62-0.64 mm (0.63 20.01); proleg with 7-11 crochet rows; antennal ratio 1:2:1.

Pupa (Fig. 19). See Jedlicka & Halgos (1981). Gill base thick, short, nearly twice as long as wide; gill with thick, posteriorly projected filament and three elongate filaments, two of the latter three (posterior-most pair) typically borne on a common base. Microsculpture: mesonotum moderately glabrous; meso- and metanotum densely set with microtubercles, but without basal microtrichia; abdominal sutures without secondary ridges; microtubercles and microtrichia distributed generally over tergite surface; microtrichia not associated with microtubercles.

Measurements. Male pupae (N = 5). Total length 2.5-3.3mm (2.97)70.31), width 1.8-2.3mm (2.1020.21). Female pupae (N = 3). Total length 2.7-2.8 mm (2.77 * 0.05), width 2.0mm.

Aduft d. See Jedlicka & Halgos (1981). Median clypeal lobe indistinct, slightly convex; lateral lobes pronounced, densely set with microtrichia. Anterior tentorial pits approximately 2 0 p posterior to clypeal incision. Subpnal suture distinct; postgena and oral region glabrous, except for a few microtrichia and setiform sensilla near anterior clypeal margin; mental tooth acute. Antennal f l slightly bulbous distally, with about 25 digitiform sensilla; each of j2 andf3 distally with anteroventrai tubercle bearing about 10 digitiform sensilla. Coxa, trochanter and femur with simple macrotrichia; fore and mid tibia sparsely set with simple macrotrichia, distally with abundant capitate rnacrotrichia, especially on ventral margin. Hind tibia densely set with simple macrotrichia on dorsal margin; capitate macrotrichia abundant on ventral margin, sparsely distributed on distal part of dorsal margin. All tarsi ventrally with capitate macrotrichia.

Terminalia. Gonostylus length greater than length of dorsal plate or aedeagal sheath. Aedeagus internally smooth.

Measurements [from Jedlicka & Halgos (1981) or personal observations of pharate adult dissected from a pupa (given in parentheses)]: Antennal length 7.5mm (11.7mm); article length ratio 2.5:1:3.5:1.5:1.5:120 (2.5:1:4.5:2:2: 185).

Adult 0 . See Jedlicka & Halgos (1981). Median clypeal lobe broadly truncate, approximately 20 pm long, with 10- 15 setiform sensilla; clypeus laterally without pronounced lobes or dense microtichia. Anterior tentorial pits approximatrly 30 pm posterior to clypeal incision.

Terminalia. Anterior bridge of genital fork posteriorly


acute, extended more than half the distance to accessory gland opening. Cercus with length of basal and distal article subequal; articles indistinctly fused to each other, separated only by shallow, ventrolateral notch.

Measurements [fro.m Jedlicka & Halgos (1981) or personal observations of two pharate adults dissected from pupae]. Antenna1 length 0.45 mm; article length ratio 3.5:1:2:1:1:1. Length of tl/t5: foreleg: 0.39-0.41; midleg: 0.38-0.44; hindleg: 0.26-0.30.

Type marerial. Holotype d [dissected from pupa]: MONGOLIA: ChiSvsgdl ajmak, Ich-Choro-go1 1650 m [=51"30'N 100"30'E], 11.viii.1976 (L. Jedlicka & J. Hufgos) (SNM) .

Material examined. MONGOLIA: Chovd ajrnak, 40 km NW Duut sum, Dund go1 nr Ulin Davaa, W.vii.80 (15. Jedlicka) [lL, 7P (46/39)] (GWC, PZW); RUSSIA: South Altai, Katen-Karagai, Buchtarma valley [-49"N 85"E] '23/VII-16' (B.Schwanwitrch) [2L, 1P( d)] (BMNH); Okhotsk Province, Ulja R Basin, Hetana R, 23.viii.85 (A. Rumitsin) [2L. 6P(2d/4Q), 281 (GWC); Primorye Province, Edinka R. 21.viii.86 ( A . Korionov) [5L, 3P(ld/ 2P)I (USNM).

Disrribution. Known from the Altai and Sayan Mountains of Russia and western Mongolia, through parts of far eastern Russia (Fig. 28). The distribution in Mongolia was summarized by Jedlicka & Halgos (1985).

Bionomia. Pulikovsky (1924) provided a brief account of the habitat and behaviour of the species; otherwise, bionomical data are unknown.

Unplaced species of Deuterophlebia

Deuterophkbia sp. A

Deuterophlebia mirabiiis Brodsky, 1930 289.

Remarks. As noted above (re Dmirabilis), material described by Brodsky (1930) as D.mirabilLF is here referred to Deuterophlibia sp., partly because immature stages have not been definitively associated with the D.mirubilis type series, or with adult males from near the type locality.

Deuferophlebia sp. 6 (Fig. 24)

Deuterophlebia sp. Singh, 1961: 109. Deuterophlebiidae Dubey & Kaul, 1971: 47.

Larva,(Fig. 24). Dorsal colour: body dark brown. Dorsal cranium (instar IV), including clypeus and mandible base, dark brown. Distal antennal article relatively dark, without digtiform microtrichia; pit-like sensillum near apex of ventral branch. Chaetotaxy: prl, pm, at and mt simple setiform sensilla; sbt (meso- and metathorax) 4anciform sensilla; up with two clusters of digitifom sensilla, without setifom sensilla.

Measurements. Instar I11 ( N = 3). Total length 2.8-

3.2 mm (3.05 2 0.23), cranial width 0.48mm, crochet rows 6-9, antennal ratio 1: 13:3. Instar IV ( N = 5) . Total length 2.8-4.8 mm (3.74 2 0.73), cranial width 0.70-0.74 mrn (0.72 c 0.02), crochet rows 10-14, antennal ratio 151.

Pupa. Unknown. Adult. Unknown. Material examined. INDIA: NW Himalaya, S slope of

Hampta Pass, headwaters of Alhni R, Sta. 11, 4000m [*32"15'N 77"15'E], 30.v.1970 (Kaul & Dubey) [6L] (GWC); Chandra Valley, 12,000ft [-3T15'N 77"15'E], 8.vi.1955, 10.vi.1955 (M. S. Mani) [2L] (GWC),

Distribution. Known only from the Pir Panjal Range and the Himalayas (Singh, 1961).

Bionornics. Singh (1961) and Dubey & Kaul (1971) provided a brief discussion of the larval habitat. Larvae were apparently collected in meltwater torrents in associ- ation with Simuliidae, Blephariceridae and other tor- renticolous insects.

Deuterophlebla sp. C (Fig. 21)

Larva. Unknown. Pupa (Fig. 21). Male antennal sheath coiled around

periphery of ventral abdomen twice, terminated near head. Gill base thick, short, nearly twice as long as wide; gill with thick, short, posteriorly projected filament and three elongate filaments, two of the latter three (posterior-most pair) typically borne on a common base. Mesothoracic tubercle on dorsolateral margin of rnesonotum, above and slightly posterior to gill base; each tubercle extending anterolaterally as 2 stout spines: spines broadly united basally, tapering to near outer margin of gill. Mesonotal bands transverse, raised, darkly sclerotized, between thoracic tubercle and ecdysial line; antetodorsal mesonoturn with 2-3 small, raised, darkly sclerotized dots on either side of ecdysial line. Metanotal bands similar to mesonotal hands, but less darkly sclerotized and shorter. Abdominal segments I, 11, V1 and VII laterally with 12, 16-20, 5 and 11 thorn-like spines, respectively.

Microsculpture: rnesonotum rugose; meso- and metano- turn irregularly set with ovoid microtubercles bearing apical microtrichia; abdominal sutures simple, without secondary ridges; tergites with microtubercles bearing apical rnicrotrichia; adjacent cuticle glabrous. Light microscopical examination of the thoracic and abdominal cuticle suggests a microsculptural arrangement similar to that of D.inyoensL, with microtrichia appearing as a corona around the microtubercle; although this suggests that microtrichia are in an apical position, confirmation may require SEM examination.

Measurements. Male pupa (N = 1). Total length 3.24 mm, width 2.40 mm.

Adult. Unknown. Material examined. NEPAL: Tributary to Dudh Kosi,

south of Tengboche, 29.x.1973 (K. D. & S. F. Kimball) [ IP] (USNM); slide-mounted in Canada balsam.

Distribution. Known only from Nepal. Remarks. The presence of two spines on each side of

Revision oj Palaearctic mountain midges 15

The presence of branched sht sensilla is one of several synapomorphies for these two species.

Character 3, sensilla prl: simple, setiform (0); branched, with two or these filaments (1).

Outgroup criteria again suggest that simple, setiform sensilla are plesiomorphic for the Deuterophlebiidae. Ontogenetic criteria (Nelson, 1978; Wheeler, 1990) indicate the same polarity, as early instar larvae of all species possess only simple sensilla. Among species for which larvae are known, the plesiomorphic state (Fig. 24) occurs in only D.inyoensis, shasta and D . sp. B. The apomorphic state (Fig. 23) is typical of D.nipponica, bicarinata, sajanica, personata, vernalis, nielsoni and coloradensis, and is used to help define a clade that also includes D.tyosenensis and shasta. This character state has not been determined definitively for D.tyosenenris; I assume that this species demonstrates the same state as other members of the clade. This hypothesis predicts that the apparently plesiomorphic condition in D.shasta is secondarily derived.

Character 4 , sensilla at and mt: simple, setiform (0); branched, with two or three filaments (1).

The distribution of character states is identical to that of character 3: the plesiomorphic state (Fig. 2%) occurs in D.inyoensis, shasfa and D. sp. B, whereas the apomorphic state (Fig. 25b) is typical of D.nipponica, bicarinata, sajanica. personata, vernalis, nielsoni and coloradensis.

Character 5 , sensilla group ap: two clusters of several digitiform sensilla (0): only two individual digitiform sensilla (1).

The plesiomorphic state (Fig. 2 5 ) occurs in D.inyoensis. nipponica, hicarinuta, sajanica. and D. sp. B. Unfortu- nately, the sensilla arrangement is unknown for most Palaearctic species, including D.tyosenemis. Under the accepted phylogeny (Fig. 27), the apomorphic state (Fig. 25b) supports monophyly of the Nearctic species D.shasta, personata. vernalis, nielsoni and coloradensis. Lack of data for most Palaearctic species requires that I only tentatively accept this character as a synapomorphy of the Nearctic faunna, exclusive of D.inyoensis. Homologous structures could not be identified in the Blephariceridae.

Character 6, instar I1 proleg: without crochets (0); with three rows of eversible crochets (1); with four rows of eversible crochets (2).

This proleg feature could potentially be among the most informative larval characters. The proleg structure of a later-instar deuterophlebiid is markedly different from that of a larval blepharicerid and represents one of three striking synapomorphies of the family. Except for the first instar, the deuterophlebiid proleg bears multiple rows of crochets. An increasing number of crochet rows presumably reflects greater structural specialization; thus, the presence of three crochet rows in larval instar I1 is considered more plesiomorphic than is four rows. This polarity decision supports the accepted phylogeny (Fig. 27) and possibly all alternative arrangements (Fig. 26). Three crochet rows are present in D.inyoensis and nipponica (Kitakami, 1938b), whereas four are typical of all Nearctic species exclusive of D. inyoensis. Unfortunately, second- instar larvae are unknown for most Palaearctic species.

the mesothorax suggests that this species may be a close relative of, or conspecific with, D.tyosenesis. However, the Nepal pupa and D.tyoseneniis are markedly different in microsculpture, size and distributions, suggesting they are indeed separate taxa.

Phylogenetic analysis

The monophyly of the Deuterophlebiidae has been established by Courtney (19!Nb, 1991b). Recent analyses of interfamilial relationships have provided compelling evidence for a sister-group relationship between the Deuterophlebudae and Blephariceridae (Wood & Borkent, 1989; Courtney, l W b , 1991b). Character polarities for species within the Deuterophlebiidae were therefore based on the assumption that blepharicerids are the nearest outgroup. Features of the larva, pupa and adult were used to evaluate the phylogenetic relationships of species of Deuterophlebia. Character numbers in the following discussion correspond to those in Table 1 and Fig. 27. Hypo- thesized plesiomorphic and apomorphic characters are listed sequentially, and character codes for the matrix and cladograms are given in parentheses. Any character that is polymorphic for a particular taxon is indicated by numbers separated by a slash mark (e.g. ‘O/ 1’). Phylogenies reconstructed initially by character-by-character (or ‘hand’) analysis were tested with the computer program Hennig86 (Farris, 1988). For the latter, I assumed equal character weights and used the implicit enumeration (ie*) option. In my discussion of characters and phylogenetic results, the taxon descriptors ‘basal’ and ‘apical’ are in reference to the accepted cladograrn (Fig. 27); i.e. they describe the relative deviation from the deuterophlebiid groundplan. as determined by the character analysis.

Larva

Character I , clypeal lobe: uniformly dark or with distinct pattern (0); uniformly pale (1).

Cranial colour patterns are relatively homoplastic in some deuterophlebiids (e.g. D.coIoradensis, see Courtney, 199oa); however, the colour of the clypeal lobe is rather conservative. The clypeal lobe is uniformly pale in only two species, D.nielsoni and coloradensis. Deuterophlebia bicarinata exhibits a similar arrangement, with the basal part of the clypeal lobe darkly pigmented and the apical part pale; I do not consider this condition as homologous. The presence of a pale clypeal lobe supports the monophyly of D.niei.soni and coloradensis.

Character 2, sensilla sbt: simple, setiform (0); branched, with two or three filaments (1).

Blepharicerid larvae demonstrate a variety of sensillar arrangements (Hogue, 1987), but only rarely are these structures branched. Outgroup criteria therefore suggest that the presence of simple, setiform sensilla is a groundplan feature of the Deuterophlebiidae. This arrangement is typical of all species except D.6icarinata and sajunica.

16 Gregory W . Courtiiey

Further data about this character in Palaearctic species will test this hypotheses.

Pupa

character 7, anterolateral margin of mesonotum: without spines (0); with one pair of anterolaterally projected spines (1); with two pairs of anterolaterally projected spines (2).

The plesiomorphic state (Fig. 22) occurs in D.inyoensis, nipponicu, vernalis, nielsoni, coloradensis and D . sp. A. The presence of mesonotal spines (Figs 18-21) could be used to define a clade comprised of D.shastu, personata, sujunicu, bicarinata and ryosenemis, with the last species showing the final stage in the transformation series (Fig. 20). This character state is also found in D . sp. C (Fig. Zl), a species presently known only from the pupa. If this pupal morphotype is conspecific with a species presently known only from adults (e.g. D.blephuris and rnirabilis), the presence of mesonotal spines must be interpreted as homoplastic or even plesiomorphic. I currently accept the former view, and suggest that the spined condition has arisen independently in D.shnsta + persotiuta and D.ryosenensi.s + (sajanica + bicaririutu). Homoplilsy is suggested further by polymorphism in D.colorudetuis - i.e. in certain populations, pupae have short, curved mesonotal spines (Courtney, 199Oa). Again, more data on Palaearctic species are needed to resolve these conflicting characters.

Character 8 , gill filaments: three, posterior filament absent (0); four, posterior filament short (I); four, posterior filament elongate (2).

Most Blephariceridae (subfamily Bkpharicerinae) possess four gill lamellae of more or less equal size and shape; however, their structure (Zwick, 1977) is markedly different from the gill filaments of deuterophlebiids. Furthermore, the gills of other blepharicerids (subfamily Edwardsininae) are even less similar (i.e. they lack 'filaments' altogether). Because these two groups of blepharicerids represent the basal dichotomy of the family, it is difficult to determine the groundplan condition for the Blephariceridae. Likewise, establishing the character polarity for the Deuterophlebiidae is problematic. For the cladistic analysis, I tested two alternative interpretations: (1) homology between the four filaments (lamellae) in deuterophlebiids and blepharicerids; and (2) non-homology of these structures. The latter interpretation required that I conduct a preliminary cladistic analysis of other characters, then, based on the resultant tree, select a functional outgroup from within the Deuterophlebiidae. This preliminary analysis suggested that the condition in D.inyoemis (i.e. three filaments) is plesiomorphic. This state is typical of most species known as pupae (Fig. 22, see also Figs 14-22 in Courtney, 1990a), whereas the presence of four filaments (Figs 18, 19) is unique to D.tyosenensiS, bicurinara, sajatiica. and D . sp. C. Even if homology between the four filaments (lamellae) of deuterophlebiids and blepharicerids is accepted, the tree

topology does not change (i.e. suggesting that the four- filamented condition is secondarily derived in deuterophlebiids). This provides further support for interpretation 2 (non-homology of structures).

Charucter 9. abdominal tergites: without sclerotized bands (0); with sclerotized bands on all or some segments

The pupae of D.nipponica (Fig. 22) and bicarinata (Fig. 18) possess transverse, raised, darkly sclerotized bands on the abdominal tergites; however, species differ in that bands are on segments I-VII in D.nipponica, but only segments 1 and 11 in D.hicarinata. Although a transformation series is suggested, other characters suggest that the apomorphic condition has arisen independently in these two species. The phylogenetic analysis supports the latter interpretation (Figs 26, 27).

(1).

Churucwr I f ) , postgena and oral region: with microtichia (0); glabrous (1).

The plesiomorphic state is typical of most species of Drrrrrrophlchiu and apparently of the Blephariccridae and other nematcxcrous flies. Thc prescncc of a glabrous postgena and oral region helps dctine ;I large clade consisting of nine spccies: D.nippoiiioa. tyosrrreti.si.s. sajunicu, bicuririuta, persoriutu. shastu. vertiuli.s, coiorudrtisis and tzie/.sorii. However, a glabrous postgcnil and oral rcgion also occurs rn D.bruchyrhiriu, prcsumably by convergence. Although the latter suggests the character could be homoplastic. 1 accept this feature as il moderately important indicator of the monophyly of the above nine spccios.

Character 11, male antennal length: short, length of f4 less than 90 times length of pedical (0); long, length of f4 more than I 10 times length of pedicel (1).

I consider this antennal feature to be one of the most informative characters in the analysis. The presence of an extremely elongate male antenna is a striking synapomorphy of the Deuterophlebiidae. Althouth the antennae of D.brachyrhiiru and oporina are elongate, they are substantially shorter than those of all other species of mountain midge. Comparison of the relative length of antennal articles demonstrates that most difference is related to the length of the last flagellomere cf4). The plesiomorphic state is considered an intermediate stage in the transformation from a more or less 'normal' nematocerous antenna (e.g. as in blepharicerids) to that of most members of the Deuterophlebiidae. The apomorphic state supports the monophyly of all Deuterophlebia except D . brachyrhina and oporina.

Character 12, male antennal f1: with cluster of 10 or fewer sensilla on anterior margin (0); with cluster of more than 10 sensilla on anterior margin ( I ) .

The plesiomorphic state is typical of the Blephariceridae and most Deurerophlebia. Only D . bicurinata and sajarzica demonstrate the apomorphic condition, which is one of several characters that support monophyly of the two species.

Hci.i.siutr or Puluearctic rnouriraiti rnidge.v 17

Character 13, male antennal fz-3: without raised tubercles on anterior margin (0); each with raised tubercle on anterior margin (1).

Raised tubercles on antennal fz-3 are absent in only three species, D.brachyrhina (Fig. S), oporina and inyoensis (see Fig. 52 in Courtney, IYWa). The apomorphic condition is typical of all other species (see Fig. 51 in Courtney, 199Oa). This character helps define a large clade comprised of eleven species.

Character 14, male tibia and tarsus: without capitate macrotrichia (0); with capitate macrotrichia along ventral margin (I).

In most mountain midges, the ventral margin of the tibia and tarsus bears numerous capitate macrotrichia (Fig. 8). I have argued (Courtney, 1990a, 1991a) that capitate macrotrichia are an adapation to the males' exclusively aerial life above torrential habitats. Male deuterophlebiids gcnerally fly at or near the stream surface, usually over turbulent reaches, which often leads to their being knocked into the water. Recovery is usually rapid, with males able to resume flight within a few seconds. This ability is presumably related to the mechanical and hydrofuge properties of elongate appendages, particularly those with cnpitate macrotrichia (i.c. legs). Similar structures, including long, filiform antennae, and legs with enlarged empodia or without tarsal claws, are seen in males of some surface-skating flies (e.g. Tokunaga, 1932). Among deuterophlebiids, capitate macrotrichia are lacking only in D.hruchyrhinu (Fig. 7). This condition is typical of all other Diptcra. as well as all other insects. The apomorphic state is both unique and innovative, and is considered a strong indicator of the monophyly of the Deuterophlebiidae, exclusive of D.hruchyrhinu. This character provides compelling evidence that D.hruchyrhina is the sister group of all other mountain midges.

Characrer 15, male gonostylus: short, its length subequal to length of dorsal plate (0); elongate, its length greater than length of dorsal plate (1).

Unlike the genitalia of most Diptera, those of deuterophlebiids furnish little phylogenetic information. Marked fusion of structures has obscured segmental boundaries and led to a rather simple arrangement. However, certain features of the male genitalia provide insights on relationships among plesiomorphic lineages. One pertains to the structure of the gonostylus. In D. brachyrhina, oporina and inyoensis, gonostyli are short and relatively unmodified (Figs 9, 10). with the ventromedial margin more or less even. In other species, gonostyli are elongate, distinctly tapered apically and sinuate ventromedially. The latter, presumedly apomorphic state supports the monophyly of the Deuterophlebiidae, exclusive of the above three species.

Character 16, bridge of genital fork (female): extended posteriorly less than half the distance to the accessory- gland opening (0); extended posteriorly more than half the distance to the accessory-gland opening (1).

The plesiomorphic state (Fig. 17) has been observed in D.oporitia and inyoensis, whereas the apomorphic condition (Fig. 16) is typical of most other species. The character

state has not been determined for D.hruchyrhinu, hlepharis and tyoset retais .

Discussion

Interspecific relationships

The present analysis evaluated sixteen characters and fourteen taxa (Table I; Fig. 27), including a generalized outgroup based primarily on the Blephariceridae. Autapo- morphies of individual species, which provide no information about relationships, were excluded from the analysis. Character analysis using the computer program Hennig86 (Farris, 1988) resulted in twenty-one equally parsimonious trees of length 24 and consistency and retention indices of 79 and 86, respectively. Reversing the polarity of character 8 (re the pupal gill) led to the same number of trees of slightly longer length (26) and lower consistency and retention indices. (73 and 81, respectively). By accepting the polytomy involving Dmzirabikv, blepharis and the remaining 'apical' clade. the number of equally parsimonious trees is reduced to seven (Fig. 26a-g). With one exception (see below), there was complete congruence between trees under either interpretation of character 8. Alternative hypotheses differ only with respect to relationships among the apical clade and indicate considerable stability of basal dichotomies. These basal dichotomies reflect features of the adult male, especially the following four characters: (14) tibia and tarsus with capitate macrotnchia (present in all species except D.hruchyrhina); (1 1) antenna elongate, length of f4 s 110 times length of pedicel (typical of all species except D. brachyrhina and oporina); (13) J?-3 with raised tubercle on anterior margin (typical of all species except D. hrachyrhitra. oporitaa and inyoemis); (15) gonostylus elongate, its length greater than length of dorsal plate (distribution as in character 13). Resolution of the more apical clade is problematic. I accept the tritomy involving D. blepharis and D.mirabilis largely because no characters permit further resolution of relationships. The most compelling synapomorphy for the remaining species is the glabrous postgena and oral region; however, this condition has apparently arisen independently in D. brachyrhirra. Relationships among the nine species above the tritomous node are poorly resolved, as indicated by the consensus tree (Fig. 26h).

In spite of the differences between alternative hypotheses (Fig. 26a-g), certain patterns are repeated. In five of the seven alternatives (Fig. 26a, b, e-g). D.nipponicu is the sister group of the remaining eight species. The character basis for this arrangement is unclear, but may reflect the absence of pupal mesothoracic spines, in combination with features of larval chaetotaxy. One tree has D.nipponica nested among the remaining eight species, as the sister group of an eastern Asian species group (Fig. 26c). This relationship is collapsed in another tree (Fig. 26d), which has D.nipponica as part of a tritomy involving the eastern Asian species group and a Nearctic species group. Another repeated pattern (Fig. 2612-g) suggests the monophyly of these Nearctic species. Support for this hypothesis


Table 1. Matrix of characters and alternate states used in the cladistic analysis of Deuterophlebia. IJ = plcsiomorphic; 1, 2 = apomorphic; ? =character state unknown.

Character ~~

Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Outgroup D. brachyrhina D.oporina D. blepharis D.rnirabi1i.s D.nipponicu D. tyosenensis D.sajanica D. bicarinara D. inyoensis D.shasta D . personata D. vernalis D. coloradensis D. nielsoni

Semaphoront

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? 0 0 0 0 1 0 0 ? ? ? ? ? ? ? ? ? O 1 0 1 1 ? ? ? ? ? ? ? ? ? 0 1 0 1 1 1 1 0 ? 1 1 0 1 0 0 1 1 1 0 I 1 1 1 0 ? ? ? ? ? 2 1 0 1 1 0 1 1 l ? 0 1 1 1 0 ? 1 2 0 1 1 1 1 1 1 1 0 1 1 1 0 ? 1 2 1 1 1 1 1 1 1 1 0 0 0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 2 1 0 0 1 1 0 1 1 1 1 0 0 1 1 1 2 1 0 0 1 1 0 1 1 1 1 0 0 1 1 1 2 0 0 0 1 1 0 I 1 1 1 1 0 1 1 I 2 011 0 0 1 1 0 1 1 1 1 1 0 1 1 1 2 0 0 0 1 1 0 1 1 1 1

l ?

L L L L L L P P P A A A A A A A

apparently comes from larval chaetotaxy (character 5: the presence of only two digitiform sensilla in sensilla group ap) and from the increased number of proleg crochets on instar 11 larvae (character 6). Unfortunately, larvae are unknown, or unassociated with adults, in four Palaearctic species, and character states have not been determined for some other Palaearctic species (due either to incomplete descriptions or lack of material for examination). So, although characters pertaining to larval chaetotaxy and proleg crochets could provide compelling insights into phylogenetic relationships, the current lack of comparative data makes definitive conclusions impossible. More data are clearly needed for all Palaearctic species. A final pattern seen in five of the seven alternative trees (Fig. 26b-f) suggests the monophyly of D.shasta and personata. In all but one of the five trees (Fig. 26b), D.shasta and personata are nested among other Nearctic species, presumably because of larval chaetotaxy; they then cluster with each other because of the shared presence of pupal mesothoracic spines. In Fig. 26(b) the node assignments of these characters are reversed; i.e. mesothoracic spines help place D.shasta and personata with the eastern Palaearctic group of D . tyosenensis, sajanica and bicarinata, then larval chaetotaxy clusters the former two species. As with larval characters, data on pupae are lacking for several Palaearctic species; pupae are unknown, or unassociated with adults, in D.brachyrhina, oporina, blepharis and mirabilis. When the polarity of character 8 (re the pupal gill) is reversed, six of the above trees (Fig. 26a-c, e-g) are retained. i n the tree that is not congruent, the difference pertains to the position of D.nipponica - instead of being part of an unresolved tritomy, this species is placed as the sister group of a Nearctic species group.

Based on the distribution of characters and on the

repetitious patterns shown in the seven alternative trees, 1 selected one (Fig. 26e) as the most acceptable phylogenetic hypothesis. This hypothesis was then usccl to plot the distribution of all characters (Fig. 27).

Biogeography

Although many question remain about the phylogenetic relationships among species of Dercterophlebio, I will attempt to place the current knowledge in a biogeographical framework. Amphi-Pacific distributions, as demonstrated by mountain midges. are typical of several aquatic and montane insect groups (e.g. the black fly Gymnopais Stone, caddisfly Uenoidae, beetle Amphizoidae and ice- cricket Grylloblattidae). Several genera of blepharicerids (Agathon Roder, Bibiocephala Osten Sacken, Philorus Kellogg) exhibit similar distributions. Unfortunately, the phylogenetic relationships both within and among these genera have not been determined, and the biogeographic significance of their distributions remains unclear.

The relationship between Nearctic and Palaearctic groups has been the subject of numerous biogeographical studies (e.g. Ross, 1956: Allen, 1983; Noonan, 1986, 1988), and their results have had important implications for dispersal routes (e.g. Beringian vesus trans-Atlantic) and timing. Phylogenetic analysis of species of Deuterophfebia suggests that the Nearctic fauna is not monophyletic and provides some indication of the direction and timing of faunal exchange. Superposition of the 'accepted' phylogeny (Fig. 27) and contemporary species distributions gives an area cladogram (Fig. 29) that suggests the Nearctic fauna originated from at least two invasions of North America: an early invasion by the ancestral D.inyoensis and a sub-

Revision of Palaearctic mountain midges 19

inyoensits ’ blepharir mirabilis

nipponica vernalis

coloradensis nielsoni shasta

l$f

a

’“E tyosenensis majmica

bicarinata

outgroup

inyoens is blepharis mirabilis

nipponica nieleoni coloradensis

personata

tyosononsis

vernalis

b

sa janica bicarinata

outgroup brachyrhina

oporina

nielsoni coloradensis personata shasta

C

sajanica bicarinata


inyoensis blopharis mirabil is

“4 tyomenenm is

vernalis

sa janica bicarinata

shasta personata coloradensis nielsoni

blephuim mirabilis

nipponica bicarinata ma j mica

shasta personata COl0rrdend.E nidsoni

e 21

vernalis


inyoends blophuis mirabilim

nipponica bicarinata

tyorenensis ma j anicr

prrmonata shasta vernalis coloradensis

nielsoni

f

outgroup

blapharis mirabilis

nipponica bicarinata majanica

tyosenenmis mhasta personata

9

vernalis coloradensis nielsoni

c nipponica - porsoruta - vernalis J nielsoni coloradensis

bicarinata sajmica r ’ tyosenensis

Fig. 26. Equally parsimonious trees from Hennig86 analysis of Deuterophlebia; Nelsen consensus tree at lower right.

sequent invasion by the ancestor of all other Nearctic species. The phylogenetic relationships and amphiPacific distribution of extant mountain midges support an hypothesis of Beringian dispersal for Deuterophlebia. The Beringian land bridge existed periodically until the Late Wisconsinan glaciation, and presumably could have provided ideal conditions for dispersal of cool-adapted taxa. It is doubtful that either Nearctic invasion occurred across the Pleistocene land bridge. Although our knowledge about rates of cladogenesis in deuterophlebiids is limited, a Pleistocene invasion would have left little time for the

apical clade of Nearctic species to differentiate into five species. A mid-late Tertiary transgression is more prob- able. This event would have been preceded by an earlier invasion by the ancestral D.inyoensis. Trans-Atlantic dispersal routes, available until c. 20-35 million years ago (Noonan, 1988), could have provided an alternate corridor, but such an hypothesis is highly improbable for deuterophlebiids. This hypothesis, which requires subsequent extinction of mountain midges in both Europe and eastern North America, is much less parsimonious than an exclusively Beringian alternative.

20 Gregory W . Courttwy

Character state coding

c] Plesiomorphic m + l Apomorphic unknown

I 2 3 4 5 6 7 8 9 10111213141516

brachyrhina

oporina

in yoensis

blepharis

mirabilis

nipponica

tyosenensis

sajanica

bicarinata

personata

shasta

vernalis

coloradensis

nielsoni

I I I I I I I I I I I I I I I I

I I I I I I I I I I I I I I I I

Fig. 27. Charac‘tcr state distribution and hypothesizcd phylogcnctic rclationships of Drrrteropldrbici. Numbcrs rcfcr to characters in the text and tables.

Within the Palaearctic and Nearctic regions, the deuterophlebiid fauna contains two relatively widespread species and several known from just a few localities. The Nearctic species D. inyoemis and coloradensis are widespread in western North America, but noticeably absent from northern and central British Columbia (Courtney, 19%). This suggests a vicariant pattern, with Beringian and ‘southern’ elements. Because disjunct populations show little evidence of morphological divergence, a late Wisconsinian vicariant event, presumably involving glaciation, is suggested. Alternatively and perhaps more likely, this distributional gap may reflect the inaccessibility of northern

and central British Columbia and concomitant lack of collecting. One of the more puzzling discoveries is the paucity of deuterophlebiid ,records from Alaska. Streams in the Brooks, Alaska and Chugach Ranges contain seemingly suitable habitats, yet neither deuterophlebiids nor blepharicerids are present (personal observation). The absence of blepharicerids is particulary enigmatic, since these flies are widespread and apparently much more vagile than are deuterophlebiids. The Palaearctic species D.rnirabilis and sajatzica are widespread in central and eastern Palaearctic. the former known from the Himalaya, T e n Shan and Altai mountains. and D.sujutiicu in the

Revision of Palaearctic mountain midges 2 1

ranges of central and northeastern Asia (Fig. 28). The latter species has apparently colonized some arctic and subarctic streams in northeastern Russia (Makarchenko, 1991, personal communication). At present, no Holarctic species are known).

Among other species, . D.personafa is also relatively widespread (Courtney, 1990a). As in D.inyoenris and coloradensis, the distribution of D.personata suggests a vicariant pattern, with most records from California, Oregon, Washington and southern British Columbia, but a disjunct population in the southern Yukon. Interestingly, northern populations (from Washington, British Columbia and the Yukon) consist exclusively of females, providing strong evidence of geographic parthenogenesis (Courtney, 199Oa). The only other mountain midge that may be parthenogenetic is D.nipponica, a species restricted to Japan. It is possible that parthenogenetic reproduction has

been a contributing factor to the colonization of recently glaciated habitats and islands. Colonization of new habitats may depend largely on downstream drift of immature stages or on flight by adults from adjacent watersheds. However, because of their short life (<2h) and limited flight abilities (Courtney, 1991a), adult deuterophlebiids appear to be extremely poor dispersers. Parthenogenetic reproduction may be a mechanism by which population.. first become established. The present distribution of man) species of Deuterophlebia may best be interpreted a: relictual; i.e. they were previously more widespread and the disjunct pattern seen today reflects extinction across much of their former range. Reasons for their demise are unclear, but may reflect a poor ability to adapt to long-term fluctuations in stream conditions. Some species (e.g. D.coloradenris) are, however, relativel) eurybiontic, inhabiting a wide range of [mostly 'lowland']

0 D. brachyrhina D. nipponica

A D. oporina D. bicarinata

0 D. blepharis & D.ujanicr 8 D. mitabills

I I I I I D. tyusenens

60' 75' 90' 105' 120' 150' 165'

50'

3'

0'

0'

0'

Rg. 28. Distribution of Palaearctic species of Deriterophfebiu.


I I I 0 nip

1 '.\ bl mi in --. .\\

Fig. 29. Area cladogram of world species of Deurerophlebia, based on phylogenctic hypothesis from Fig. 27. Abbreviations: bi, D.bicarinata; bl, D.blepharis; br, D.brachyrhiRa; co, D.coloradensis; in, D.inyoenric: mi, D.mirabilis; nie, D.nielsoni; nip, D.nipponica; op, D.oporim; pe, D.personata; sa, D.sajanica; sh, D.shastu: ty, D.&osenensis; ve, D.vernalis.

streams (Courtney, 1991a); this might explain the relative 'success' of this species (i.e. with respect to its distribution). On the other hand, D.inyoemis, a species restricted primarily to cold, mountain streams, is considered a stenobiont (Courtney, 1991a) yet its distribution dearly suggests that it, too, is 'successful.' Among Nearctic species, the inhabitants of streams falling between these two categories tend to show the most restricted distributions. Again, reasons for this pattern are unclear and warrant further study.

It is clear that many discoveries about the ecology, phylogeny and biogeography of mountain midges remain for future dipterists and aquatic biologists. Because of large gaps in our knowledge about Palaearctic deuterophlebiids, the phylogenetic and biogeographic hypotheses presented in this analysis should be considered tentative in many respects. Discovery of the immature stages of

all Palaearctic mountain midges, especially the Himalayan species D. brachyrhina and oporina, will test these hypotheses and, in turn, provide valuable insights into the origin of other montane and aquatic organisms.

Acknowledgments

I am grateful to the following individuals for providing material for this study: 3. M. Cumming (aC); L. Jedlicka, Comenius University, Bratislava, Slovakia; W. N. Mathis (USNM); W. Schacht (ZSM); S. Singh, St John's College, Agra, India; B. Townsend (BMNH); S. Uchida, Tokyo Meteropolitan University, Tokyo, Japan; P. Zwick (PZW). I thank W. N. Mathis and two anonymous reviewers for their valuable comments on the manuscript, and C. Morigaki, Oregon State University, for providing trans-

Rmisioti of Pulueurctic mountain rnidge.v 23

lations of Kitakami (1954). Yie (1933), and other papers on Japanese Deuterophlebia. Support for this study was provided by the Department of .Entomology, University of Alberta (through a Natural Sciences and Engineering Research Council of Canada grant to D. A. Craig) and the Department of Entomology, National Museum of Natural History, Smithsonian Institution.

References

Allen, R.T. (1983) Distributional patterns among arthropods of the north temperate deciduous forest biota. A n m k of the Missouri Botanical Garden, 70, 616-628.

Brodsky. K.A. (1930) Zur Kenntniss der Wirbellosenfauna der Bergstrome Mittelasiens. 11. Deuterophlebia mirabilis Edw. Zeitschriff f i r Morphologie und dkologie der Tiere, 18,

Brodsky, K.A. (1980) Mountain Torrent of the Tien Shun. A Faunisric-ecology Essay. Dr W. Junk Publishers, The Hague.

Brodsky, A.L. & Brodsky, K.A. (1926) Deuterophlebiu from the mountains of central Asia. Bulletin. University of Central Asia, 13, 23-26, [in Russian, with French summary].

Courtney, G.W. (1989) Morphology, systcmatics and ecology of mountain midges (Diptera: Dcuterophlcbiidae). Ph.D. thesis, Univcrsity of Albcrta, Edmonton.

Courtncy, G.W. (1Wa) Rcvision of Ncarctic mountain midges (Diptera: Dcutcrophlcbiidac). Journal of Naturul History, 24, 81- 118.

Courtncy. G.W. (lY9Ob) Cuticular morphology of larval mountain midgcs (Diptcta: Dcutcrophlcbiidac): implications for thc phylogcnctic rclationships of Ncmatoccra. Cunadiun Jourml of

Courtncy, G.W. (llY9la) Lifc history pattcrns of Ncarctic mountain rnidgcs (Diptcra: Dcutcrophlcbiidac). Journul of the North Americun Benfhologicul Society, 10, 177- 197.

Courtncy. G.W. (19Ylb) Phylogcnctic analysis of the Blepharicero- morpha, with special reference to mountain midges (Diptera: Deuterophlebiidae). Systematic Entomology. 16. 137- 172.

Dubey, O.P. & Kaul, B.K. (1971) Torrenticole insects of the Himalaya. IV. Some observations on the ecology and the characters - insect communities of the R. Alhni. Oriental Insects, 5,47-71.

Edwards, F.W. (1992) Deuterophlebia mirabilis, gen. et sp.n., a remarkable dipterous insect from Kashmir. Annals and Magazine of Natural History, 9, 379-387.

Fams, J.S. (1988) Hennig86, version 1.5. Computer program and documentation. Port Jefferson Station, New York.

Hennig, W. (19’3) Diptera (Zweifliigler). Handbuch der Zoologze, 4(2), 2/31 (Lfg. 20), 337pp.

Hogue, C.L. (1987) Blephariceridae. In: Griffiths, G.C.D. (editor). Flies of the Nearctic Region, 2 (4), 1- 172. E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart.

Jedlicka, L. (1980) Notes on the distribution of mountain midges (Dipten: Deuterophlebiidae) with special regard to Asian species. Acta Universitatrs Carolinae-Biologica, 1977, 31 1-315.

Jedlicka, L. & Halgos, J. (1Y81) Deuterophlebia sajunica sp.n., a new species of mountain midges from Mongolia (Diptera: Deuterophlebiidae). Biologia (Bratislava), 36, 973-981.

Jedlicka, L. & Halgos, J. (1985) Distribution of Deuferophfebia sajanico Jcdlicka et Halgos, 1981 in Mongolia. Acta Facultatis Rerum Naturalium Universitatis Comenianue. Zoologia, 28.

Kaul. B.K. (1979) Taxonomy of the families Blcpharocendae and

289-32 1.

Zoology, 611,556-578.

27-32.

Deuterophlcbiidac (Diptcra) in India. Abstracts, Workshop on Advances in Insect Taxonomy in India and the Orient, Manila (H.P.) [October 9-12, 1979), pg. 62.

Kaul, B.K. (1986) Ecology of some Diptcra (Nernatocera) at high altitudes on the Himalaya. Abstracts, First International Congress on Dipterology. Budapest, Hungary [August 17-24, 19861, pg. 122.

Kennedy, H.D. (1958) Biology and life history of a new species of mountain midge, Deuterophlebia nielroni, from eastern California. Transactions of the American Microscopical Society, 77,201-228.

Kennedy, H.D. (1960) Deuterophlebia inyoemis, a new species of mountain midge from the alpine zone of the Sierra Nevada Range, California (Diptera: Deuterophlebiidae). Transactions of the American Microscopical Society, 79, 191-210.

Kitakami, S. (1929) An unusual insect from Japan (Diptera: Deuterophlebiidae). Zoofogicd Magazine, 41, 379-380 [in Japanese].

Kitakami, S. (1938a) The Blepharoceridae and Deuterophlebiidae from KyOskyG, Yaku-sima Island, and Taiwan (Formosa). Kontyu, 12, 214-215 [in Japanese].

Kitakami, S. (1938b) The Deuterophlebiidae of Japan. Annof- utiones Zoologicae Japonenses, 17, 487-507.

Kitakami, S. (1954) Imagos of Deuterophlebia from North Korea. Konryu, 20,73-74 [in Japanese].

Mani, M.S. (1962) Introduction to High Altitude Entomology. Methuen & Co. Ltd, London.

Mani. M.S. (1968) Ecology and Biogeography of High Altitude Insects. Dr W. Junk Publishers, Thc Haguc.

McAlpinc, J.F. (1981) Morphology and tcrminology - adults. In: McAlpinc, J.F., Pcterson, B.V., Shcwcll, G.E., Tcskcy, H.J., Vockcroth, J.R. & Wood, D.M. (coordinators). iManual of Nearctic Diptera, Volume 1, Rcscarch Branch, Agricdturc Canada Monograph 27. pp. 9-63.

McAlpinc, J.F., Pctcrson. B.V., Shcwcll, G.E., Tcskcy, H.J., Vockeroth, J.R. & Wood, D.M. (1981) Introduction. In: McAlpine, J.F. et ul. (coordinators). Manual of Neurctic Diprera, Volume 1, pp. 1-7. Research Branch, Agriculture Canada Monograph 27.

Nelson, G. (1978) Ontogeny, phylogeny, paleontology, and the biogenetic law. Systematic Zoology. 27,324-345.

Noonan, G.R. (1986) Distribution of insects in the northern hemisphere: continental drift and epicontinental seas. Bulletin of the Entomological Society of America, 32, 80-84.

Noonan, G.R. (1988) Faunal relationships between eastern North America and Europe as shown by insects. In: Downes, J.A. & Kavanaugh, D.H. (editors). Origins of the North American Insect Fauna, pp. 39-53. Memoirs of the Entomological Society of Canada 14.

Pennak, R.W. (1954) Notes on mountain midges (Deutero- phlebiidae) with a description of the immature stages of a new species from Colorado. American Museum Novitates. 1276, 1 - 10.

Pulikovsky, N. (1924) Metamorphosis of Deuterophlebia sq. [sic] (Diptera: Deuterophlebiidae Edw.). Transactions of the Entomological Society of London, 1924,45-62.

Rohdendorf, B.B. (1974) The Historical Development of Diptera [translated from Russian]. Hocking, B.. Oldroyd, H. & Ball, G.E. (editors). University of Alberta Press, Edmonton, Alberta.

Ross. H.H. (1956) Evolution and Classification of the Mountoin Caddisfles. The University of Illinois Press, Urbana, Illinois.

Singh, S. (1961) Entomological survey of Himalaya. Part XXXII. A note on the larva of an apparently undescribed species of Deuterophlebia Edw. (Deuterophlebiidae: Diptera-Nematocera) from North-West (Punjab) Himalaya. Agra University Journal

24 Gregory W. Courtiiey

offteseurch, 10, 1OY-113. Tcskey, H.J. (1981) Morphology and terminology - Larvae. In.

McAlpinc, J.F., Pcterson, B.V., Shcwcll, G.E., Tcskcy, H.J., Vockeroth, J.R. & Wood, D.M. (coordinators) Munrrul of Neurcric Dipteru, Volumc 1, Rescarch Branch. Agriculture Canada Monograph 27. pp. 65-88.

Tokunaga, M. (1932) Morphological and biological studics of a new chironomid fly, Pontomyiu pac@cu from Japan. Memoirs of the College of dgriciilture Kyoto Imperial University, 19, 1-56.

Tokunoga, M. (1%) Somc ncmatoccrous Diptcra OF the northeast of Afghanistan. Results of the Kyoto University Scientific Expedition to the Karakorum und Hindukush, 1W5, 8,

Wheclcr, Q.D. (1W) Ontogeny and character phylogcny. 273 - 2%.

Clcrdirlics, 6 , 225-268.

Wirth, W.W. (l9j1) A ncw mountain midge from California (Diptcra: Dcuterophlcbiidac). Pun-PaciJic Enmmologur. 27. 49-57.

Wood, D.M. & Borkcnt, A. (1Y8Y) Phylogcny and classification of thc Ncrnatoccra. In: McAlpine, J.F. a Wood, D.M. (coordinators). rClaniw1 of Nenrcric Diptera. Vdurnc 3, pp. 1333- 1370. Rescarch Branch, Agdculturc Canada Monograph 32.

Yic, S . (1933) Obscmations on a Japanese Deuterophlebiu (Dip- tcra). Transactions of the Nuturai ,%tory Society of Formma. 23, 271-2915 [in Japancse, with English summary and platc captions].

Zwick, P. (1977) Australian Blephariccridae (Diptcra). Austruliun Journal of Zoolop. Supplemenrury Series. 46. I- 121.

Acccptcd 11 Novcmbcr 1Y93

Documents

Revision of Palaearctic mountain midges (Diptera