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September 1997

The Paduniella (Trichoptera: ) of China, with a phylogeny of the World

Youwen J. Li Clemson University, Clemson, SC

John C. Morse Clemson University, Clemson, SC

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Li, Youwen J. and Morse, John C., "The Paduniella (Trichoptera: Psychomyiidae) of China, with a phylogeny of the World species" (1997). Insecta Mundi. 276. https://digitalcommons.unl.edu/insectamundi/276

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 281

The Paduniella (Trichoptera: Psychomyiidae) of China, with a phylogeny of the World species

Youwen J. Li and John C. Morse Department of Entomology, Clemson University, Clemson, SC 29634, USA

Abstract: The phylogenetic relationships of the species ofPadul~iellaare analyzed based on characters oflarvae, pupae, and adults (mainly male genitalia). The is monophyletic and most closely related to Psychotj~yia,and Metalype in the subfamily Psychomyiinae. Nine species groups are suggested. Eight species, including six new to science, are reported from the People's Republic of China for the first time.

Key words: Psychomyiinae, Psychottzyia,, Metalype, male genitalia, female genitalia, new species

Introduction relationships of the genus within the family Psy- The genus Padu7tiella was erected by Ulmer chomyiidae were analyzed by Li and Morse (in (1913) for the species Paduniella selnarai~gei~sis press). Paduniella is most closely related to Psy- Ulmer. Ulmer (1922) established a similar genus, cholnyia (s. 1.) and Metalype, but the relationships Psychomyiodes, based on Psychoinyiodes africai~a among these monophyletic genera remain unre- Ulmer from Cameroon. Lestage (1926) established solved. Malicky (1995) considered Metalype to be a the subfamily Paduniellinae to include Paduniella synonym of , but did not provide phylo- and Psychoinyiodes and two new closely related genetic evidence for his opinion. The latter two genera, Mesopadui~iellaand Propaduniella.. The genera are treated as outgroups in the present main character supporting Psychoinyiellodes as phylogenetic analysis. In this paper, we add more genus status is the discoidal cell of the fore wing characters to help clarify the relationships of this present. Actually, it is such an obscure vein that genus and Psycholnyia and Metalype. some researcher neglect it (Mosely 1936). However, No species were recorded from the People's this character is a plesiomorphy in this group and Republic of China before this research. This paper could not be served as evidence to support mono- reports eight species from China, including six phyletic group. Other three genera were also erect- species new to science. All type specimens are ed based on characters of wing venation that are deposited in the Department of Plant Protection, very hard to see, especially for some cross veins. Nanjing Agricultural University, the People's Re- Consequently, published descriptions vary in their public of China (NAU), and the Clemson University interpretations. For example, the main diagnostic Collection, Department of Entomology, character for the genus Propaduniella is the lack of Clemson University, South Carolina, USA (CUAC). anterior wing Fork V; for the type species P. ceyla7~- ica, anterior wing Fork V is absent in Ulmer's (1915) original illustration but present in Schmid's Paduniella morphology and groundplan (1958) later one. Such difficulties discouraged sub- sequent workers from recognizing any of the gen- Adult. Overall length (in following descrip- era other than Padui~iella. tions = distance from front of head to tips of folded For these reasons, forty species of Paduniella forewings) 2.8-5.8 mm. Forewings each 2.0-5.0 mm (sei~sulato), including the six new species in this long, yellow to yellow-brown. Forewings and hind- paper, have been included in the genus from the wings acute at apex (Fig. 1). Each forewing with Afrotropical(4 spp.), Oriental (34 spp.), West Pale- Forks 11,111, IV, and V; each hindwing with Forks arctic (1 sp.), East Palearctic (3 spp.), and Nearctic I1 and V. Each hindwing with acute projection on (1 sp.) Biogeographic Regions. costal margin at middle. Head with several warts In support of our higher classification of these on vertex: pair of large oval occipital warts, pair of species, this paper also analyzes the phylogeny of slender curved ocellar warts, single forked frontal world Padui~iella(s. I.) species based on available wart, and inconspicuous anterior warts (Fig. 2). information, mainly characters of male genitalia, Each maxillary palpus 6-segmented; each labial to determine any justification for recognizing Pro- palpus 4-segmented (Fig. 3). padui~iella,Mesopadui~iella, and Psychoinyiodes Female genitalia (Figs. 4-5). Abdominal seg- as distinct subgenera or genera. The phylogenetic ment VIII (VIII) synsclerotized, its ventral posteri- 282 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI

or margin broadly excised. Segment IX not evident. cata. [This is possibly the two lateral parameres Segment X (X) twice as tall as long, subconical, fused, but the homology of this character is very tapering from middle to posterior, with transverse difficult to determine. It does not exist in other row of long setae (trans. r. set.) near base of seg- genera of Psychomyiidae and , ment, internal ventral basomesal apodeme (apo- but may occur in (Ceratopsyche deme) forked and diverging; ventral meson cleft species) and (e.g., Polycelttro- from posterior end to anterior 115 of segment, pus colei Ross, 1941).] Inferior appendages each opening widest near anterior end of cleft; poster- with one compressed segment and with short mesal oventral margin projecting posteriad, closely ap- branch. pressed against ventral surface of segment IX (IX). (Mathis and Bowles, 1995; Figs. 32-35). Segment XI small, hairy, with pair of tiny, slender, Larva of P. ltearctica distinguished by two charac- one-segmented cerci (cercus). teristics from those of other psychomyiid genera for Male genitalia (Figs. 7-9). Male genitalia of which larvae are known: four well-developed teeth Padultiella species homogeneous, with distinctions on concave margin of each anal claw [similar to mainly involving tergum IX (t. IX), sternum IX (S. those of Psycholnyia species, but teeth lacking in IX), inferior appendages (inf. app.), superior ap- Tiltodes and species (Wiggins, 1996)l and pendages (sup. app.), and phallic apparatus [in- submental sclerites small and wider than long [like cluding paramere (para.), phallobase (phb.), and those of Tiltodes and Lype species, not longer than phallicata (phc.)]. Tergum IX smaller than tergum wide as in Psycholnyia species (Wiggins, 1996)l. VIII, usually triangular in dorsal and lateral views, Pupae (Mathis and Bowles, 1995). Pupa of P. with long, slender, sclerotized, lateral bands pro- ltearctica with 6-segmented maxillary and 4-seg- ceeding anteriad to juncture with corresponding mented labial palpi, mandibles whip-shaped, la- strips on sternum IX. Sternum IX shape nearly like brum with only 3 pairs of setae (rather than 5 pairs sternum VIII but broader in ventral view and with as in known pupae of other genera). long lateral bands directed anteriad. Lateral bands of tergum IX and sternum IX in each side, extend- Paduniella species of China ing anteriad and joining each other at small point Paduniella communis, new species in very acute angles. Superior appendages distinct (Figs. 4-5, 7-9) posteriorly and extending well beyond tergum X; Adult forewing length: 2.56-2.95 mm. Overall lateral bases of superior appendages extending length: 3.06-3.42 mm. Color in alcohol uniformly anteriad and indistinguishable from lateral bands pale yellow-brown, antennae annulate with brown. of tergum IX. Pair of narrow strips from joining Male genitalia (Figs. 7-9). Tergum IX broad, points of sternum IX and tergum IX directed an- round at posterior margin in dorsal view. Superior teroventrad and connecting these points to dorsal appendages each oval, acute at apex, almost twice side of base of phallic apparatus (= phallobase as long as tergum IX. No median process arising region, Fig. 7), fused sclerotized strips of phallic from sclerotized strips of segment IX. Inferior ap- shield and sclerotized strips of sternum IX in some pendages each with basal third broad, abruptly (Morse, 1975); similar strips also in narrowed to 113 basal thickness, then gradually genus except directed upward to base of Tiltodes, enlarged and rounded at apex; mesal branch aris- phallic apparatus (They probably developed in Ti- ing from near apex of mesal surface of basal part. and independently.). These strips nodes Padulziella Phallic apparatus vertical basally, then arched in sometimes not connected with phal- Paduniella caudoventrad, with deep anterior incision at verti- lobase. Most species with one or more median cal juncture of phallobase and phallicata and scle- processes (med. proc.) arising between anterior rotized strips of segment IX; phallobase very short, ends of sternal strips above phallobase and pro- about 118 as long as phallicata; dorsal paramere ceeding posteriad, these processes absent in sDme slender, arched dorsad then caudad, nearly as long (primitive) species. Phallic apparatus directed as phallicata, arising from dorsal concavity close to caudad, straight or slightly arched; phallobase phallobase, with short apical fork; phallicata com- broad, open in ventral view, short in some (primi- pressed, gradually clavate, and round at apex in tive) species, half as long as phallicata or as long as lateral view. phallicata in other (advanced) species; phallicata Female genitalia (Figures 4-5). Segment X more or less compressed, with slender dorsal spine- largest in basal third, with transverse ring of long like process arising between phallobase and phalli- INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 283

v Fused antenna1 trans, r. set. frontal

trans. r. set. I

Figs. 1-5. Adults of Padulriella species: la, forewing of Pa,dulriella furcata, n.sp.; lb, hindwing of same; 2, vertex of Padulriella uralerrsis Martynov, dorsal; 3, headof P~d~i~iellauraleirsis Martynov, lateral; 4, female genitaliaofPadulciella COI?LI~LUIL~S,n.sp., lateral; 5, same, ventral. 11,111, IV, and V = Forks 11,111, IV, and V; trans. r. set. = transverse row of setae. 284 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI

Treelength: 40 CI: 0.94 RI: 0.97 RC: 0.92

Fig. 6. Phylogenetic cladogram of species of Paduniella (s. l.), with Psychornyia and Metalype as outgroups. Underlined numbers = clade numbers; plain text numbers = homologues identifiedinTables 1and 2; apostrophe (')=parallelism or convergence. Nine species groups recognized: P. a~~da,rne~~e~tsisGroup (branch#46), P. uearctic Group (branch #47), P. 1nahil~dr.aGroup (branch #48), P. var~deli Group (branch#48), P. afr.icana Group (branch#51), P. ankya Group (branch#59), P. zc~.ale~~sisGroup (branch#60), P. ar~~urer~sisGroup (branch#61), P. subhalza,ra Group (branch#62). INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 285

Figs. 7-12.Male genitalia ofPadur~iella,species:7, Padu~~iellacor~~r~~~~r~is,n.sp., lateral; 8, same, dorsal; 9, same, right halfofsternum IX andphallic apparatus with paramere and right inferior appendage, ventral; 10,Padur~iella bifida, n.sp., lateral; 11,same, dorsal; 12, same, right halfofsternum IX and phallic apparatus with paralnere and right inferior appendage, ventral. inf. app. = inferior appendage, m. b. inf. app. = mesa1 branch of inferior appendage, para. = paramere, phb. =phallobase, phc. = phallicata, s. IX= sternum IX, sup. app. = superior appendage, t. IX = tergum IX. 286 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI setae along ridge between tapering and thick por- Etymology. colnmunis, Latin, meaning "com- tions. Sternum X split mesally, with pair of sharp mon," referring to the dominance of the species in internal apodemes arising from anterior margin. central eastern China. Segment XI round with pair of spine-like cerci. Diagnosis.The male of this new species is very Paduniella bifida, new species similar to that of Padur~iellai~earctica Flint, 1967 (Figs. 10-12) in the short phallobase, the absence of a median process above the phallic apparatus, and the undi- Male forewing length 2.3 -2.7 mm, overall length vided apex of each inferior appendage. However, 2.88-3.24 mm. Color in alcohol uniformly pale yel- their differences are obvious. In the new species, low-brown. tergum IX has a round posterior margin (P. nearc- Male genitalia. Tergum IX triangular in dor- tica has a rnesal point), the superior appendages sal view. No median process arising from sclero- are twice the length of tergum IX (P. nearctica has tized strips of segment IX. Each superior append- superior appendages slightly konger than tergum age straight, more than twice length of tergum IX. IX), the basal part of each inferior appendage is Inferior appendages each tapering to apex, with three times as wide as the apical part (less than apical incision as long as 118 length of inferior twice as wide in P. ~~earctica),the paramere tip is appendage, and short truncated rnesal branch aris- forked (unique in Paduniella), and the paramere is ing near base. Phallobase very thick in lateral view, arched (straight in P. nearctica). about half as long as phallicata; dorsal paramere Phylogeny. This new species is one of the arising at juncture of phallobase and phallicata, more primitive species of the genus, lacking the about 113 distance from base of phallic apparatus, median sternal strip process above the phallic ap- spine-like, mostly cylindrical, with apex widened, paratus. Its relationship with other species in the obliquely truncated, and depressed; basal 314 of genus remains uncertain. phallicata slender, apex expanded, round in lateral Type material. Holotype male: Song-cun, view. Ding-xi-he, 33 km E. of Jin-xian, An-hui Province, Diagnosis. The male genitalia of the species 120 m elevation, 8 June 1990, collected by Morse, are very similar to those of Padulziella vandeli Yang, and Sun (NAU). Allotype Female: same data Dkcamps, 1965 in the big phallobase; the incised as holotype (NAU). Paratypes: AN-HUI PROV- apex of each inferior appendage; the compressed, INCE: 4 Males 2 Females, Yang-jia-tan, Feng- round, expanded apex of the phallicata; and the yuan-shui, She-xian, 215 m elevation, 25 May 1992, obliquely cut apex of the paramere in lateral view. collected by Morse and Sun (CUAC); 11 Males, Yao- They differ in that the apical incision of each cun, Yong-feng-he, Lang-xi-xian, 23 May 1990, inferior appendage is much shallower, far less than collected by Morse, Yang, and Sun (NAU). JIANG- 114 of the length of the appendage; the inferior XI PROVINCE: 1Male, Lao-dong-qiao, Gui-xi-xian, appendages are acute apically, much narrower in 240 m elevation, 5 June 1990, collected by Morse the new species than in P. vandeli; and the forew- and Sun (NAU); 15 Males, Qin-hua-he, 57 km N. of ing length is about half that of P. vandeli's. Wu-yuan, Wu-yuan-xian, 250 m elevation, 25 May Phylogeny. The species is considered most 1990, collected by Morse, Yang, and Sun (NAU). closely related to P. valtdeli because of their uniquely HU-BE1 PROVINCE: 9 Males 1 Female, 50 km N. shared obliquely truncated paramere spine. W. of Yin-cheng, tributary of Da-fu-shui, Ji-shan- Distribution.The species is distributed in Si- xian, 90 m elevation, 17 July, 1990, collected by chuan and Jiang-xi Provinces, part of the Oriental Morse (CUAC); 9 Males, 47 km N. W. of Yin-cheng, Biogeographic Region of China. tributary of Da-fu-shui, Jin-shan-xian, 80 m eleva- Type materials. Holotype male, Si-mian-shan, tion, 17 July, 1990, collected by Yang and Wang Fei-long-he, Jiang-jin-xian, Si-ChuanProvince, 800 (NAU); 9 Males, Da-fu-shui, Tian-dian-Dam, Yi- m elevation, 7 July 1990, collected by Yang (NAU). cheng City, 40 m elevation, 16 July 1990, collected Paratypes: JIANG-XI PROVINCE: 2 Males, Xi-qi- by Morse and Yang (NAU). he, 10 km S. of Gui-xi, Gui-xi-xian, 30 m elevation, Distribution. The species is distributed in 4 June 1990, collected by Yang, Morse, and Sun Hu-bei, An-hui, and Jiang-xi Provinces, Oriental (NAU) . Biogeographic Region of China. Etymology. bifida, Latin, meaning "divided," referring to the incision at the apex of each inferior appendage. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 287

Paduniella bilobata, new species Yang, and Sun (NAU); YUN-NAN PROVINCE: 2 (Figs. 13-15) Males, Nan-wen-he-xiang, Nan-wen-he, Ma-li-po- xian, 600 m elevation, 12 July 1990, collected by Li Male forewing length 2.52-2.88 mm; overall and Ke (NAU). length 3.06-3.31 mm. Color in alcohol uniformly Etymology. bi-, Latin, meaning "two," and pale yellow-brown. lobatus, Latin, meaning "with a projection," refer- Male genitalia: Tergum IX triangular, with ring to the two median processes. concave sides in dorsal view. Superior appendages fused with tergum IX, each with length about twice Paduniella buddha, new species width in lateral view and about three times width (Figs. 16-18) in dorsal view, apex acute and pointed mesad in dorsal view and dorsad in lateral view. Inferior Male forewing length 3.65 mm; overall length appendages each tapering to blunt apex, mesal 4.38 mm. Color in alcohol uniformly pale yellow- branch 213 as long as appendage. Two median brown. processes arising between anterior tips of sclero- Male genitalia. Tergum IX subdorsally in- tized processes of segment IX almost equal length, cised, appearing three-lobed in dorsal view, with longer process reaching apex of phallicata, each middle lobe acute. Superior appendages each as process sharp and curved ventrad at apex. Phallo- long as wide basally, round at apex in dorsal view, base large, slightly shorter than phallicata; acute in lateral view, with minute ventrolateral paramere spine-like and arising from base of phal- spine basally. Inferior appendages nearly straight, licata, sharp at apex; phallicata compressed, slight- slightly tapering to blunt apex, with mesal branch ly and gradually broader toward apex in lateral appearing as small hairy wart about 314 distance view. from base. Two slender median processes between Diagnosis. The male of this species is similar anterior tips of sclerotized strips of segment IX, one to those of Paduniella wa~~gtalzraie7~sisMaliclry, process exceeding apex of phallicata, other process 1995, P. suwa7~namaliMalicky, 1993, and P. axake- much shorter, 113 as long as first process and 7~arnMalicky, 1995, in possessing two subequal positioned on left side of its base; phallobase thick median processes. However, the superior append- and nearly as long as phallicata, with ventral ages of the new species are broad, each with its surface excised close to base ofphallicata; paramere apicomesal corner acute and directed dorsomesad, spine about same length and basally nearly as thick not straight and round atthe apex as in those other as base of phallicata; phallicata mostly slender, species. apex depressed to spoon shape, curved dorsad. Distribution. The species is distributed in Diagnosis. The male genitalia of this species southeastern (An-hui and Jiang-xi Provinces) and are very similar to those of Paduniella rnaeklan- southwestern (Yun-nan Province) China, in the ge7~sisMaliclry, 1993 in that the median process has Oriental Biogeographic Region. a short basal lobe. However, the two species can be Phylogeny. The species is a member of the distinguished by the following characters: The Padu7~iellaafricana Group, as suggested by the median process has only one short lobe at its base two long median processes arising between the in this species (two in P. 1naekla1~ge7~sis)and the anterior tips of the segment IX sclerotized strips, small ventrolateral process of each superior ap- but its relationships with other species within the pendage is near the base of the superior appendage group are not resolved. in this species (near the apex in P. 1naekla7~gensis). Type materials. Holotype male: Song-cun, Distribution. This species is found only at the Ding-xi-he, 33 km E. of Jin-xian, An-hui Province, type locality in Si-chuan Province, southwestern 120 m elevation, 8 June 1990, collected by Morse, China, Oriental Biogeographic Region. Sun,andYang (NAU). Paratypes: JIANG-XI PROV- Phylogeny. The species is closely related to P. INCE: 4 Males, 61 km S E of Gui-xi, Lao-dong-qiao, 7naekla1~ge1~sisMalicky and Padul~iellafurcata, Xi-qi-he, Qui-xi-xian, 240 m elevation, 5 June 1990, sp. n., as suggested by the subdorsal incisions of collected by Morse and Sun (CUAC); 40 Males, 59 tergum IX, and the depressed and upturned apex of km S E of Gui-xi, Xi-qi-he, Gui-xi-xian, 210 m the phallicata uniquely shared by these species. elevation, 5 June 1990, collected by Yang (NAU); 88 Type materials. Holotype male, E-mei-he, 8 Males, Qi-hua-he, 57 km N of Wu-yuan, Wu-yuan, km W of Jing-shui, E-mei-shan, Si-chuan Province, 250 m elevation, 25 May 1990, collected by Morse, 288 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI

1040 m elevation, 1 July 1990, collected by Morse Paduniella uralensis bicornis Martynov and Yang (NAU). (Figs. 2-3, 22-24 ) Etymology. "buddha," the name of the founder Paduiziella r~raleizsisMartynov, 1914, pp. 5-10, 17, 19, of the religion of Buddhism. The type location of the 20 , 21, figs. 1- 5. Type 1ocality:Ural Mountains species, Emei Mountain, is the holy mountain of (Lakes Ilmen and Suratkul), Russia. Buddhism in China. Mesopadr~iziellauralei~sis (Martynov) - Lestage, 1926, p. 385. Paduniella furcata, new species Padu7ziella rsrale7zsis Martynov - Lepneva, 1928, p. 25. (Figs. 1, 19-21) Padz~lziellaura,lerzsis Martynov - Martynov, 1929, p. 30. Padz~niellar~rale~zsis Martynov - Martynov, 1934a, p. Male forewing length 2.60 mm, overall length 207, figs. 145-146. 3.17 mm. Color in alcohol uniformly pale yellow- Padr~niellaz~raleizsis bicorlzis Martynov - Martynov, brown. 1934a, p. 208, fig. 147. Male genitalia (Figs. 19-21). Tergum IX in- Padu~ziellazsralensis bicornis Martynov - Martynov, cised subdorsally, with middle lobe narrow and 1934b, p. 334. apically blunt. Superior appendages each tall, height Padz~niellar~ralel~sis Martynov - Martynov, 1948, p. slightly more than half of length, truncated, with 908, figs. 486c-e. tiny ventrolateral tooth near apex. Inferior ap- Padz~~~iellaurale7~sis Martynov - Tanida, 1993, p. 58. pendages each with large acute ventral tooth at Male forewing length 2.98 mm, overall length middle, apex truncated, mesal branch 213 length of 3.50 mm. Color in alcohol uniformly pale yellow- inferior appendage. Single median process arising brown, antennae annulate with brown. atjuncture of anterior apices of sclerotized strips of Male genitalia (Figs. 22-24). Tergum IX very segment IX and posterior end of phallobase and short, deeply notched mesally. Superior appendag- anterior end of phallicata; curved right at apical 11 es each slender, suddenly curved mesad subapical- 3, acute apically, and exceeding apex of phallicata. ly, slightly exceeding inferior appendages. Inferior Phallobase compressed; paramere spine arising appendages each thick basally, slightly more slen- from middle of phallicata, sharp, twisted, arched, der in middle, apically blunt, with mesal branch reaching apex of phallicata; phallicata slightly de- taller than base of appendage. Single median pro- pressed subapically, curved dorsad at apex. cess between anterior apices of sclerotized strips of Distribution. The species is known only from segment IX twisted, apex acute, extending slightly the type locality in Jiang-xi Province, in the Orien- beyond phallic apparatus. Phallobase short, about tal Biogeographic Region of China. 114 as long as phallicata; paramere spine arising at Diagnosis. The male genitalia of the species juncture of phallobase and phallicata, straight, are very similar to those of P. inaehla7~ge~~sisMal- very sharp at apex; phallicata straight, slender, icky and P. buddha, sp. n., in the short and rounded compressed and shaped like battle-ax at apex, with superior appendages each with the little ventrolat- acute apicodorsal and apicoventral points. era1 process, the subdorsally incised tergum IX, Distribution. The species is distributed in the and the spoon-like apex of the phallicata curved East Palearctic Biogeographic Region from the dorsad. However, the large tooth on the ventral Ural Mountains through the southern Ussuri River side of each inferior appendage is unique in the region of Russia to northeastern China. The nom- genus. inate subspecies occurs in the western end of this Phylogeny. The species is closed related to the range and the subspecies P. urale~~sisbicornis lineage composed of P. lnaekla1~ge7~sisand P. bud- Martynov, 1934 in the eastern end. dha, sp. n. Diagnosis. The male of this species is similar Type materials. Holotype male, Xi-qi-he, 10 to that of Paduniella ceylai~icaUlmer, 1915, in km S. of Gui-xi, Gui-xi-xian, Jiang-xi Province, 30 having the phallicata with an apicodorsal vertical m elevation, 4 June 1990, collected by Yang, Morse, extension at its apex and its apicoventral angle is and Sun (NAU). Paratypes: 5 Males, same data as acute. However, the apex of the phallicata in P. holotype (3 males, NAU; 2 males CUAC). uralensis extends ventrad, unlike that of P. ceylalt- Etymology. furca, Latin, meaning "a fork," ica. Also, the superior appendages of P. urale7tsis referring to the shape of the phallic apparatus in are curved mesad (straight, tapering to apices in P. lateral view. ceylanica) and tergum IX is concave mesally in P. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 289

Figs. 13-18. Male genitalia of PodUr~iellospecies: 13, Padul~iellabilobata, n.sp., lateral; 14, same, dorsal; 15, same, right half of sternum IX and phallic apparatus with paramere and right inferior appendage, ventral; 16, Padr~r~iellabuddha, n.sp., lateral; 17, same, dorsal; 18, same, right half of sternum IX and phallic apparatus with paramere and right inferior appendage, ventral. m. p. = median process. 290 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI

Figs. 19-24. Male genitalia of Padurriella species: 19, Padurtiella furcata, n.sp., lateral; 20, same, dorsal; 21, same, ventral; 22, Paduniella uralerrsisMartynov, left lateral; 23, same, superior appendages and tergum IX, dorsal; 24, same, right halfofsternumIX and phallic apparatus with paramere and right inferior appendage, ventral. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 291

/ \ *' med. proc. n

Figs. 25-31. Male genitalia of Padu~~iellaspecies: 25, Paclu.~~iella~para~~zure~~sis,n.sp., left lateral; 26, same, superior appendages and tergum IX, dorsal; 27, same, right half of sternum IX and phallic apparatus, with paramere and median process; 28, same, right inferior appendage, ventral; 29, Padu~aiellaa~r~u~~e~~sisMartynov, left lateral; 30, same, superior appendages and tergum IX, dorsal; 31, same, right half of sternum IX and phallic apparatus and right inferior appendage, ventral. 292 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI

Figs. 32-35. LarvaofPadur~iellar~earcticaFlint:32, right lateralview; 33, headand thorax, dorsalview; 34, head, ventralview; 35, right metathoracic leg, posterior view. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 293

urale~zsis(convex in P. ceyla~zica).The difference width in lateral view in P. a~nure~zsis),(2) the between Padulziella uralel~sisuralelzsis and P. superior appendages are about as long as broad uralensis bicorlzis is that the superior appendages basally in dorsal view (much longer in P. alnu7-en- of specimens from the Ural Mountains are more sis), (3) tergum IX is almost quadrate in dorsal view distinctively excised behind, with the lateral corner (triangular in P. anzurerzsis), and (4) the mesal also is somewhat produced in dorsal view. In our branch of each inferior appendage exceeds the specimens, the superior appendages are not pro- middle of the appendage (not reaching the middle of duced at their lateral corners, placing them in the the appendage in P. a~l~ure~zsis). subspecies P. urale~zsisbicornis. Distribution. The new species is distributed Phylogeny. Padu~~iellauralelzsis is a sister in Jiang-xi Province, southern China, Oriental species of P. ceyla~zica,as suggested by the dorsally Biogeographic Region. extended apex of the phallicata and its acute api- Phylogeny. The species is closely related to coventral angle. Padu~ziellaal~zure~zsis Martynov, as suggested by Materials examined. 2 males, 14 females, A- the truncated mesal branch of each inferior ap- shi River, Mao-er-shan Town, Shang-zhi-xian, Hei- pendage. long-jiang, 300 m elevation, 13 July 1993, collected Type materials. Holotype male, Xi-qi-he, 10 by Li and Sun (NAU). lim S of Gui-xi, Gui-xi-xian, Jiang-xi Province, 30 m elevation, 4 June 1990, collected by Yang, Morse, Paduniella paramurensis, sp. n. and Sun (NAU). Paratypes: 43 Males, same data as (Figs. 25-28) holotype (41 males, NAU; 2 males CUAC). Etymology. para,-, Greek, meaning "close, Male forewing length 2.81-2.95 mm, overall near," and Amur, area of Far East Russia adjacent length 3.31-3.53 mm. Color in alcohol uniformly to China. pale yellow-brown, antennae annulate with brown, eyes black. Paduniella amurensis Martynov Male genitalia. Tergum IX with posterior (Figs. 29-31) margin sinuous, almost straight, in dorsal view. Padt~r~iellaainzcrer~sis Martynov, 1934a, pp. 206, 208- Superior appendages triangular, about as long as 210, 334-335, figs. 148-149. tergum IX. Inferior appendages each with short Padur~iellaarnrcrertsis Martynov - Martynov, 193413, basoventral tooth, mesal branch extending beyond pp. 334-335. Padur~iellaar~zr~re~~sis Martynov - Martynov, 1935, pp. middle of appendage, appendage tapering to in- 151, 206. cised apex. Single median process arising between Padzei~iellaarnr~reizsis Martynov - Lepneva, 1953, p. anterior apices of sclerotized strips of segment IX 419. slender and sharp, twisted, exceeding apex of phal- Padz~r~iellaarnr~rer~sis Martynov - Tanida, 1993, p. 58. licata. Phallobase nearly as long as phallicata; Padt~rtiellaarnzcrerzsis Martynov - Vshivkova, 1995, p. paramere arising at their juncture, needle-lilie, as 56. long as phallicata; phallicata slender before en- larged, compressed apex almost as tall as phallo- Male forewing length 3.0 mm, overall length base, rounded in lateral view. 3.5 mm. Color in alcohol uniformly yellow-brown. Diagnosis. The male genitalia of this species Male genitalia. Tergum IX triangular, blunt are very similar to those of Padu~ziellaarnure~~sis apically. Superior appendages each broad at base, Martynov, 1934a, b in that superior appendages tapering to apex, more than twice as long as tergum are straight and tapering, the inferior appendages IX. Inferior appendages each withbasoventral tooth are incised apically and each has a truncated mesa1 inconspicuous, mesal branch obliquely truncated branch and a round ventromesal branch at middle well before middle of appendage, appendage taper- length; the single median process is twisted, long , ing to incised apex. Single median process arising slender and very sharp; the phallobase is nearly as between anterior apices of sclerotized strips of long as the phallicata; and the phallicata is expand- segment IX slender, sharp, exceeding apex of phal- ed and compressed laterally at its apex. The species licata. Phallobase longer than phallicata; paramere can be distinguished from P. arnurelzsis by four spine nearly straight, acute, reaching apex of phal- characters: (1) the phallicata is more slender and licata; phallicata compressed and round at apex in its length is more than ten times its width at the lateral view, width at middle 114 length. middle in lateral view (length about four times the 294 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI

Diagnosis. The species is similar to Paduniel- la para~nurensis,new species, resembling it and 7(5'). Phallicata about 1/12 as wide in middle as long differing from it as explained in its diagnosis above. (Fig. 25) ...... Paduniella paralnurensis, n.sp. Distribution. In addition to Si-chuan Prov- 7'. Phallicata about 113 as wide in middle as long (Fig. 29) ...... Padulziella arnurelzsis Martynov ince of China, Oriental Biogeographic Region, the species is distributed widely over the Amur region Phylogeny of world Paduniella species of Far East Russia and North Korea (East Palaearc- tic Biogeographic Region) and in India (Oriental To analyze the phylogeny of world Padu7tiella Biogeographic Region), although we are not con- species, 42 homologues were chosen from 40 of the vinced that the Indian specimens represent the 44 species of the genus (Tables 1 and 2). The same species. homologous condition for each of these characters Phylogeny. The species is closely related to was inferred by reference to the above groundplan. Padultiella paramureltsis, n.sp., for reasons cited The computer program PAUP (Swofford, 1993) above for that species. version 3.1, with its heuristic searching method, Materials examined. Si-Chuan Province: 1 was used to infer the phylogeny, employing the male, 17 km E of Ping-wu, tributary of Fu-jiang, following assumptions: all characters unordered; Ping-wu-xian, 1060 m elevation, 27 June 1990, base weight of each character I. collected by Yang and Li (NAU). 1 male 19 km E. of A single cladogram was inferred with tree- Ping-wu, tributary of Fu-jiang, Ping-wu-xian, 1090 length 40 (with base weight of each character = l), m elevation, 27 June 1990, collected by Morse CI 0.94, RI 0.97, RC 0.92 is depicted in Fig. 6. (CUAC) . Homologues 37-40 suggest that Paduniella is most closely related to the monophyletic lineage of Meta.lype and Psycholnyia. Metalype and Psycholny- Key to males of Paduniella species from ia share the four homologues 30-33. China Six homologues (1-6) strongly suggested that Padu~~iellais a monophyletic group. However, the 1. Median processes absent above phallic apparatus (Fig. 7) ...... 2 males, at least, are rather homogeneous within the 1' Median process or processes arising between anterior genus, such that discovery of informative infrage- ends of sclerotized strips of segment IX (Fig. neric homologues was difficult. Therefore, the cla- dogram of the species still has several unresolved polytomies. 2(1). Phallobase more than half as long as phallicata At the base of the cladogram for the genus, the (Fig. 7) ...... Padz~7ziellacomrnunis, n.sp. P. andarnane7~sis Group (branch #46, including 2'. Phallobase about 118 length of phallicata (Fig. 10) . Paduniella a7~da1na7~e~~sisMalicky, 1979, and Pa- ...... Paduniella bifida, n.sp. duniella salnpati Malicky, 1979) is monophyletic as 3(1'). Two median processes present (Fig. 13) ...... 4 indicated by homologues 7-10. The P. ~tearctica 3'. One median process present (Fig. 19) ...... 5 Group (branch #47, including P. nearctica Flint, Padu7~iellahatyaiensis Malicky, 1993 and Padu~t- 4(3). Median processes subequal in length (Fig. 13) .... iella ral~ogensisMalicky, 1993, is supported by ...... Paduniella bilobata, n.sp. homologues 11,12.The Paduniella lnahindra Group 4'. One median process much shorter than the other one (branch #48, including Paduttiella lnahindra (Fig. 16) ...... Paduniella buddha,, n.sp. Schmid, 1958 and Paduniellapa~tdyaSchmid, 1958) is suggested by homologue 13. The Paduniella 5(3'). Phallicata depressed and curved dorsad apically; vandeli Group (branch #49, including Paduniella inferior appendages each with apex entire (Fig. 19) ...... 6 bifida, sp. n. and Padu7tiella ualtdeli Decamps) is 5'. Phallicata compressed and expanded at apex; inferior supported by homologue 15. No homologue has appendages each with apex forked (Fig. 25) ... been found to associate these four Groups or Padu- ...... 7 niella colnlnunis sp. n. and Paduniella outtara Schmid, 1961 with other species in this clade. The 6(5). Tergum IX concave subdorsally (Fig. 20) ...... remaining species of the genus belong to a mono- ...... Paduniella furcata, n.sp. phyletic group (branch #50) suggested by homo- 6'. Tergum IX concave mesally (Fig. 23) ...... logue 16. This latter group is composed of nine ...... Paduniella uralertsis bicornis Martynov unresolved components: Paduitiella mahanawarta INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 295

Schmid, 1958, Paduiziella se~naraitgel~sisUlmer, species-group names attached to those that pres- Paduiziella methinee Chantaramongkol and Mal- ently seem to us most convincingly monophyletic. icky, 1986, Padui~iellatigridis Malicky, 1993, the Paduiziella subhakara Group (branch #62, sug- gested by homologue 27), Padui~iellaanzur-emis Group (branch #61, suggested by homologue 26), Checklist of Paduniella species Padui~iellauralensis Group (branch #60, suggest- ed by homologue 29), Paduniella aizlzya Group For each species in the following checklist, the (branch #59, suggested by homologue 25), and type country is noted, followed by an indication of Padu~~iellaafricana group composed of the rest of the major biogeographical region(s) from which the the species (branch #51, suggested by homologue species has been reported, where AT = Afrotropical, 17). EP = East Palearctic, NA = Nearctic, OL = Oriental, Homologue 12 appears independently twice and WP = West Palearctic. (branches #47 and 63) and homologue 13 twice Genus Paduniella Ulmer, 1913, p. 80; (branches #48 and 64). Discussion and taxonomic conclusions - type species: Padz~lriellase~nara~zge~rsis Ulmer (monoba- superspecific sic); synonym Mesopadz~lriella Lestage, 1926, p. 383, 384 (Martynov, 1934a, p. 206); type species: Padz~niella From the cladogram, nine monophyletic spe- z~ralelzsisMartynov (monobasic); cies groups are recognized. The type species of the synonym Propadz~lriellaLestage, 1926, p. 383, 384 (Mar- four genera mentioned in the beginning of this tynov, 1935, pp. 151, 206); type species: Paduniella paper are all in the monophyletic group, branch ceylanica Ulmer (monobasic); #50. The type species of Mesopaduiziella (Paduniel- synonym Psycholnyiodes Ulmer, 1922, p. 50 (AV Mar- la uralei~sis)(branch #30) and the type species of tynov, 1934a, p. 206); type species: Psychornyiodes Propadui~iella(Padui~iella ceylai~ica) (branch #31) africalra Ulmer (original designation). are sister species in the P. urale~~sisGroup (branch africalra (Ulmer, 1922, p. 52, figs. 3-7), Psycholnyiodes; Cameroon; AT. #60) ; of these two genera, Propaduiziella has page alnz~relrsisMartynov, 1934a, pp. 206, 208-210, 334-335, priority. The type species of Psycholnyiodes (P. figs. 148-149a-c; Russia (S Ussuri); EP. africaiza) is in the Paduiziella africaiza Group alrake~rar~zMalicky and Chantaramongkol, 1995, p. 23; (branch #51). The relationships of the type species Thailand; OL. of Paduiziella (P. seinaraizgel~sis)with these two alzdarnalzelzsis Malicky, 1979, p. 98, figs.; Andaman lineages and with other species in the monophyletic Islands; OL. group suggested by homologue 16 remain unre- al~gustaBanks, 1939, pp. 143-144, pl. 1 figs. 3, 5, 6; solved. Philippines; OL. Our purpose in inferring the relationships of aizlzya Mosely, 1939, pp. 29-30, figs. 85-90; Uganda; AT. bifida, new species; People's Republic of China (Sichuan the Padulziella species has been to establish work- Province); OL. ing hypotheses for subsequent biological work. We bilobata, new species; People's Republic of China (Anhui prefer to reflect our conclusions in the higher clas- Province); OL. sification of the group. If the included 44 species are borireelzsis Banks, 1931, p. 426; Malaysia (Sabah); OL. treated as a subfamily, then Padulziella, Psychol~zy- bz~ddha,new species; People's Republic of China (Si- iodes, and Propaduiziella would likely be consid- chuan Province); OL. ered different genera and many more generic names capelrsis Barnard, 1940, pp. 655-656, figs. 18a-e;Repub- would be needed and Mesopaduiziella as synonym lic of South Africa; AT. of Propadui~iellashould not be used. If the 44 ceylal~ic~Ulmer, 1915, pp. 42-43, 73, figs. 4-5; Sri Lanka; OL. species are treated as a genus, these genus-group colnrnzsl~is,new species; People's Republic of China names could be afforded subgeneric rank. Howev- (Anhui Province); OL. er, there are still many unresolved branches in our delzdrobiaMalicky and Chantaramongkol, 1993,p. 1159, cladogram. Furthermore, there are some character figs; Thailand; OL. parallelisms and convergences that weaken our fiZa,rnelrtosa Jacquemart and Statzner, 1981, p. 10, figs.; arguments. Therefore, to minimize future changes Zalre; AT. of species combinations, we prefer to retain all 44 fissa Martynov, 1935, pp. 150-151, 206, figs. 51a-c; species in a genus Paduiziella, with only informal India; OL. 296 Volume 11, Nos. 3-4, September - December, 1997, INSECTA MUNDI furcata, new species; People's Republic of China (Jiangxi Acknowledgements Province); OL. lzatyaieizsis Malicky and Chantaramongkol, 1993, p. We are very grateful to Dr. Michael Mathis at 1157, figs.; Thailand; OL. the University of Central Arkansas for providing lzoehleri Malicky, 1995, p. 24; Indonesia (Bali); OL. larvae and pupae of Padulziella xearctica. We are maehla7zge~zsisMalicky and Chantaramongkol, 1993, p. 1158, figs.; Thailand; OL. also thankful to Professor Yang Lianfang and other naga ad ha Schmid, 1961, p. 197, pl. 16 figs. 5-6;Pakistan; colleagues at Nanjing Agricultural University, Peo- OL. ple's Republic of China, noted in the text for helping inahal~awalzaSchmid, 1958, pp. 8, 9, 10, 11, 15, 16 18, collect the specimens. This research was supported 19, 20, 24, 28, 30, 36, 104-105, pl. 18 figs. 17-20;Sri by US National Science Foundation Grant No. Lanka; OL. DEB-9318074. This is Technical Contribution No. ~nahiizdraSchmid, 1958, pp. 18, 20, 24, 30, 36, 103, pl. 4288 of the South Carolina Agricultural Experi- 18 figs. 12-14; Sri Lanlia; OL. ment Station, Clernson University. inartyizovi Kumanslii, 1992, Korean People's Democrat- ic Republic; EP. References ina~~ryaSchmid, 1961, pp. 197-198, pl. 16 figs. 7-8; Pakistan; OL. Banks, N. 1931. Neuropteroid from North inethilzee Chantaramongkol and Malicky, 1986, p. 526, Borneo, particularly from Mt. Kinabalu. Journal of figs.; Sri Lanka; OL. the Federal Malaya Museums, Trichoptera 16: 420- nearctica Flint, 1967, pp. 310-311, figs. 1-4; United 26. States of America; NA. Banks, N. 1939. Neuropteroid insects from the Philip- outtara Schmid, 1961, pp. 196-197, pl. 16 figs. 3-4; pines. The Philippine Journal of Science 69 (2): 133- Pakistan; OL. 145, pl. 1. paizdya Schmid, 1958, pp. 5, 6, 10, 13, 19, 25, 26, 28, 36, Bowles, D. E., and M.L. Mathis. 1988. Description of 102-103, pl. 18 figs. 9-11; Sri Lanka; OL. the female of Paduiziella ~zearctica (Trichoptera: parainzsreizsis, new species; People's Republic of China Psychomyiidae). Entomological News 99 (1): 7-9. (Jiangxi Province); OL. Barnard, IC. H. 1940. Additional records and descrip- ra7zo~zgeizsisMalicky and Chantaramongkol, 1993, p. tion of new species of South African alderflies (Nleg- 1157, figs.; Thailand; OL. aloptera), mayflies (Ephemeroptera), sainpati Malicky and Chantaramongkol, 1993, p. 1156, (Trichoptera), stoneflies (Perlidae) and dragonflies figs.; Thailand; OL. (Odonata). Annals of the South African Museum 32 saizghalnittra Schmid, 1958, pp. 7, 11, 12, 13, 16, 18, 26, (6): 609-661. 28, 36, 104, pl. 18 figs. 15-16; Sri Lanka; OL. Chantaramongkol, P., and H. Malicky. 1986. Bes- selnaraizgeizsis Ulmer, 1913, pp. 81-82, figs. 2-3; Indone- chreibung von neuen Kocherfliegen (Trichoptera, sia (Java); OL. Insecta) aus Sri Lanka. Annalen Naturhistorischen siveci Chantaramongkol and Malicky, 1986, p. 526, Museums in Wien, Ser B, 88-89: 511-34, figs. figs.; Sri Lanka; OL. DBcamps, H. 1965. U11 Trichoptkres du genre Padulziel- subhakara Schmid, 1958, pp. 9, 10, 13, 16, 19, 20, 22, 25, la en Europe occidentale. Annales de Limnologie 28, 30,32, 36, 103-104,pl. 18 figs. 21; Sri Lanka; OL. l(2): 239-243. suwai~1zai?zaliMalicky and Chantaramongkol, 1993, p. Flint, 0.S., Jr. 1967. The first record of the Paduniel- 1157, figs.; Thailand; OL. lini in the New World. Proceedings of the Entomo- thitinza Chantaramongkol and Malicky, 1986, p. 526, logical Society of Washington 69: 310-311. figs.; Sri Lanka; OL. Jacquemart, S., and B. Statzner. 1981. Trichoptkres tigridis Malicky and Chantaramongkol, 1993, p. 1159, nouveaux du Zaire. Bulletin de 1'Institut royal de figs.; Thailand; OL. Sciences Naturelle de Belgique, Entomologie, Brux- uralelzsis Martynov, 1914, pp. 5-10, 17, 19, 20, 21, figs. elles 53 (21): 1-25, pls. 15. 1-5; Russia (S Ural); EP and WP; Kimmins, D. E. 1962. New african caddisflies (order subspecies Padulziella zsralellsis bicorizis Martynov, Trichoptera). Bulletin of the British Museum (Nat- 1934a, p. 208, fig. 147; Russia (S Ussuri). ural History), Entomology 12 (2): 83-121. valzdeli Dkcamps, 1965, p. 239, figs.; France; WP. Kumanski, K. 1992. Studies on Trichoptera of Korea vattagal7zaizi Schmid, 1958, pp. 5, 15, 36, 102, pl. 18 figs. (North) 111. Superfamily . Insecta 6-8; Sri Lanka; OL. Koreana 9: 52-77. vilzra~nasilzhaSchmid, 1958, pp. 4, 18, 20, 21,29, 30, 36, Lepneva, S. G. 1928. Lichinki rucheinikov Olonetskogo 105-106, pl. 19 figs. 1-6; Sri Lanka; OL. kraya karvae of Caddis Flies (Trichoptera) of Olo- ~~~zgtalzraieizsisMalicky and Chantaramongkol, 1993, nets Region]. Trudy Olonetskoi Nauchnoi Ekspert- pp. 1158, figs.; Thailand; OL. izy 6 (5): 1-125. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 297

Lepneva, S. G. 1953. Rucheiniki (Trichoptera). Zhivot- idae). Contributions of the American Entomological nyi Mir SSSR ( World of the USSR) vol. 4, Institute ll(2): 1-97. Lesnaya zona (forest region), pages 404-423. Mosk- NIosely, M. E. 1936. New African Trichoptera-1. Annals va-Leningrad, Izd. AN SSSR. and Magazine of Natural History, Ser. 10, vol. 12: Lestage, J. A. 1926. Notes trichopt6rologiques (9'"'' 429-451. Note), etude du groupe Psychomyidien et catalogue Mosely, M. E. 1939. British Museum (Natural History), systitmatique des genres et espitces decrits depuis Ruwenzori expedition 1934-5, Trichoptera 3(1): 1- 1907 (in Genera Insectorum). Bulletin et Annales de 40, 3 pls. la Societe Entomlogique de Belgique 65: 363-386. Ross, H. H. 1941. Descriptions and records of North Malicky, H. 1979 ("1978). Neue Kocherfliegen (Tri- American Trichoptera. Transactions of the Ameri- choptera) von den Andamanen-Inseln. Zeitschrift can Entomological Society 67: 35-126, 13 pls. der Arbeitsgemeinschaft Osterr., Entomologen 30 Schmid, F. 1958. Trichoptitres de Ceylon. Archiv fiir (314): 97-109. Hydrobiologie 54(1/2): 1-173. Malicky, H. 1993. Neue Trichopteren aus Thailand, teil Schmid, F. 1961. Trichopteres du Pakistan, 4me partie. 2: , , Polycentropo- Tijdschrift voor Entomologie 104(9): 187-230. didae, , Psychomyiidae, Xiphocentron- Swofford, D. L. 1993. PAUP, Phylogenetic analysis idae, Helicopsychidae, (Arbeiten iiber using parsimony, version 3.1. Laboratory of Molec- thailandische Kocherfliegen Nr. 12) (Fortsezung). ular Systematics, Smithsonian Institution, Wash- Linzer Biologische Beitrage 25 (2): 1137-1187. ington, District of Columbia. Malicky, H. 1995. Weitere neue I

Table 1. Characters used to infer a phylogeny of species 22. Paramere spine arising from base of phallicata = 0; in the genus Paduniella (s. 1.). 0 = plesiomorphy, 1 paramere spine arising from middle of phallicata = homologue. (with some species having it even closer to apex) = 1. 1. Each maxillary palpus 5-segmented = 0; each maxil- 23. Tergum IX not divided subdorsally = 0; tergum IX lary palpus 6-segmented = 1. divided subdorsallv = 1. 2. Each labial palpus 3-segmented = 1; each labial 24. Superior appendages without lateral teeth = 0; palpus 4-segmented = 1. superior appendages each with lateral tooth = 1. 3. Each hindwing with Fork I11 present = 0; each 25. Inferior appendages without conspicuous ventral hindwing with Fork I11 absent = 1. projections = 0; inferior appendages each with con- 4. Inferior appendages 2-segmented, without mesal spicuous ventral projection = 1. branch = 0; inferior appendages compressed, 1- 26. Mesa1 branch of each inferior appendage round = 0; segmented, each with small mesal branch = 1. mesal branch of each inferior appendage truncate = 5. No narrow transverse strips connecting dorsolateral 1. edges of sternum IX = 0; narrow transverse strips 27. Phallobase separated from phallicata = 0; phallo- connecting dorsolateral edges of sternum IX, ven- base fused with phallicata = 1. trally approaching phallobase = 1. 28. Sternal strip median process with less than three 6.Pupa with 5 pairs of labral setae = 0; pupa with 3 pairs long lobes = 0; sternal strip median process with of labral setae = 1. three long lobes = 1. 7.Paramere spine present = 0; paramere spine absent = 29. Phallicata apex with ventral angle obtuse = 0; 1. phallicata apex with ventral angle acute and curved 8. Superior appendages short, broad, and apically blunt dorsad = 1. = 0; superior appendages long, slender, and acute = 30. Hindwing m-cu crossvein present = 0; hindwing m- 1. cu crossvein absent = 1. 9. Superior appendages without long lateral basal pro- 31. Phallus straight at apex = 0; phallus "J-shaped" at cess = 0; superior appendages with long lateral apex = 1. basal process = 1. 32. Ocellar warts separated = 1; ocellar warts contigu- 10. Phallicata short, thick, and apically blunt = 0; ous = 1. phallicata long, slender, and acute = 1. 33. Male tergum X triangular = 0; male tergum X deeply 11. Superior appendages round at apex = 0; superior divided into two lateral parts = 1. appendages truncate at apex = 1. 34. Superior appendages without mesal apical spines = 12. Phallobase straight ventrally = 0; phallobase con- 0; superior appendages each with mesal apical cave ventrally = 1. spine = 1. 13. Paramere spine of phallicata dorsal = 0; paramere 35. Basal half of each inferior appendage not expanded spine of phallicata lateral = 1. mesally, straight, and with anterior surface sclero- 14. Apex of phallus compressed = 0; apex of phallus tized = 0; basal half of each inferior appendage more depressed = 1. or less expanded mesally and twisted slightly ante- 15. Apex of paramere spine not truncate at apex = 0; riorly at mesal edge, with anterior surface membra- apex ofparamere spine obliquely truncate at apex = nous = 1. 1. 36. Phallobase complete, without deep lateral excision = 16. Sternal strips without long median process = 0; 0; phallobase extended dorsally and ventrally, with sternal strips with long median process = 1. deep lateral excision = 1. 17. Sternal strip median process with one long lobe = 0; 37. Hindwing without prominent costal point = 0; hind- sternal strip median process divided, with one short wing with prominent costal point = 1. lobe and one long lobe = 1. 38. Larva anal claw without teeth = 0; larva anal claw 18. Sternal strip median process with one long lobe = 0; with teeth = 1. sternal strip median process with two long lobes = 39. Female abdominal segment X about same length as 1. other abdominal segments = 0; Female segment X 19. Inferior appendages without teeth at middle of elongated = 1. dorsal edges = 0; inferior appendages each with 40. Male phallic apodeme short = 0; male phallic apo- small tooth at middle of dorsal edge = 1. deme well developed = 1. 20. Inferior appendages without lateral teeth on mesal 41. Female with transverse row of setae on tergum X = branch = 0; inferior appendages each with small 0; female without transverse row of setae on tergum lateral tooth on mesal branch = 1. X = 1. 21. Sternal strip processes without short dents basally 42. Male tergum X lateral parts separated with superior = 0; sternal strip processes each with several short appendages = 0; male tergum X lateral parts fused dents basally = 1. with superior appendages for 112 length = 1. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 299

Table 2. Character states of Metalype, Psychornyia, and species of Pad~~~~rella.Plesiomorphy = 0; homologue = 1. 1 11 11 1 11 112 22222 22223 33333 33333 44 12345 67890 12345 67890 12345 67890 12345 67890 I2 Metalype 00000?000000000000000000000001 11111 II? 11 00 Psychomyin 00000 00000 00000 00000 00000 00001 1 I I00 01 11 1 11 andamanensis lI111 1011100000000000000000000000000111100 sampati Ill11 10111 000000000000000000000000001111 00 communis 11111 11000000000000000000000000000001111 00 nearctica 11111 11000 110000000000000 00000 0000001111 00 hatyaiensis 11111 11000 110000000000000000000000001111 00 ranongensis 11111 11000 110000000000000000000000001111 00 mahindra Ill11 11000001000000000000000000000001111 00 pandya IIllI 1100000100000000000000000000000111100 outtara 11111 11000000000000000000000000000001111 00 bifida 11 11 1 11000 00001 00000 00000 00000 00000 01 11 1 00 vandeli 11111 1100000001 0000000000000000000001 111 00 suwannamali 111111100000000111110000000000000000111100 anakenam 11 111 11000 00000 Ill 11 00000 00000 00000 011 11 00 wangtakraiensis 11111 1100000000 1111000000000000000001111 00 thitima 11111 1100000000 Ill00 1000000000000000l1ll00 martynovi Ill11 1100000000 Ill00 1000000000000000l11100 africana I1111 1100000000 1110000000001000000001111 00 filamentosa 11111 110000'0000 1ll000000000100000000ll1100 bilobata 11111 1100000000 1110000000000000000001111 00 magadha 11 11 1 1100000000 1100001000000000000001111 00 dendrobia I1 111 11000 00000 11000 01000 00000 00000 011 11 00 maurya lllll 1100000000 1100000000000000000001ll100 maeklangensis 11111 1100000010 1100000110000000000001111 00 buddha 11111 1100000010 110000011000000000000111l00 furcata Illll 1l00000010 11000001100000000000011l100 ankya Illll 1100000000 1000000001 000000000001111 00 koehleri 11111 1100000000 1000000001 000000000001111 00 uralensis 11111 1100000000 100000000000010000000l1l100 ceylanica 11111 1100000000 1000000000000100000001111 00 tigridis 1111 1 1100000000 10000 00000000000000001111 00 amurensis 11 11 1 1100000000 10000 00000 100000000001111 00 paramurensis 11 I I1 11000 00000 10000 00000 10000 00000 01 11 1 00 methinee 11 111 11000 00000 10000 00000 00000 00000 01 11 100 semarangensis 11111 1100000000100000000000000000000111100 mahanawana 1 I1 11 11000 00000 10000 00000 00000 00000 01 11 1 00 vattagamani 11 11 1 11000 00000 10000 00000 0 1000 00000 01 11 1 00 vikramasinha I I1 11 11000 00000 10000 00000 01 000 00000 01 11 1 00 sanghamittra 111111100001000100000000001000000000111100 siveci 11111 1100001lOO 1000000000010000000001111 00 subhakara 11111 1100001100 1000000000010000000001111 00