Testing phylogenetic hypotheses

Testing phylogenetic hypotheses: new species of Gymnophora Macquart (Diptera: Phoridae) from Japan and Nepal, with an updated world classification Can. J. Zool. Downloaded from www.nrcresearchpress.com by "Institute of Vertebrate Paleontology and Paleoanthropology,CAS" on 06/06/13 For personal use only.

BRIANV. BROWN Department of Entomology, University of Alberta, Edmonton, Alta., Canada T6G 2E3 Received November 14, 1988

BROWN, B. V. 1989. Testing phylogenetic hypotheses: new species of Gymnophora Macquart (Diptera: Phoridae) from Japan and Nepal, with an updated world classification. Can. J. Zool. 67: 2543-2556. Thirteen species of Gymnophora are recorded from Japan and Nepal: G. inexpectata Beyer (Japan and Nepal), G. fastigiorum Schmitz (Japan), G. integralis Schmitz (Japan), and 10 new to science: G. priora (Japan), G. enigma& (Nepal), G. nepalensis (Nepal), G. pro& (Japan), G. longissima (Japan), G. quadriseta (Nepal), G. platypalpis (Nepal), G. gotoi (Japan), G. pararcuata (Japan), and G. s e t u k (Nepal). Gymnophora priora is related to the two species of the Neotropical G. cymatoneura-group, G. nepalensis, G. prolata, and G. longissirnu are classified in the Holarctic G. nigripennis-subgroup, G. gotoi and G. pararcuata are classified in the Palearctic G. integralis-series, and G. setulata is classified with the Nearctic G. luteiventris-subseries. Other new species have more questionable positions in the preexisting classification and point out some probable errors. Keys to the species of Japanese and Nepalese Gymnophora are given, as well as a key to world species of the G. nigripennis-subgroup. A phylogenetic, partially sequenced classification of the genus is given, including changes prompted by the discovery of the new species.

BROWN, B. V. 1989. Testing phylogenetic hypotheses: new species of Gymnophora Macquart (Diptera: Phoridae) from Japan and Nepal, with an updated world classification. Can. J. Zool. 67 : 2543 -2556. Treize espkces de Gymnophora sont connues au Japon et au NCpal : G. inexpectata Beyer (Japon et NCpal), G. fastigiorum Schmitz (Japon), G. integralis Schmitz (Japon), et de 10 espkces inCdites : G. priora (Japon), G. enigmata (NCpal), G. nepalensis (NCpal), G. pro& (Japon), G. longissima (Japon), G. quudriseta (NCpal), G. platypalpis (Nepal), G. gotoi (Japon), G. pararcuata (Japon) et G. setulatQ (NCpal). Gymnophora priora est apparentke aux deux espkces du groupe nCotropical G. cymatoneura, G. nepalensis, G. prolata et G. longissirnu sont classifiks au sein du sous-groupe holarctique G. nigripennis, G. gotoi et G. pararcuata font partie de la sCrie palkarctique G. integralis et G. setulata appartient h la soussCrie nCarctique G. luteiventris. Les autres espkces nouvelles occupent des positions plus incertaines dans la classificationp k existante et mettent en Cvidence certaines erreurs probables. On trouvera ici des clCs des espkces japonaises et nkpalaises de Gymnophora ainsi qu'une clC h 1'Cchelle mondiale des espkces du sous-groupe G. nigripennis. Une phylogCnie partiellement ordonnCe ou sont intCgrCs les changements occasionnCs par l'addition des nouvelles espkces est proposCe pour le genre Gymnophora . [Traduit par la revue]

Introduction One benefit of phylogenetic analysis (Hennig 1966) is that previous hypotheses about phylogenetic relationships, expressed in the form of nested character sets, can be tested by new data. The ability to predict character states correctly in newly discovered taxa corroborates the hypothesis, while character conflicts point out possible errors. Initial classifications of the Holarctic (Brown 1987a) and Neotropical (Brown 1987b) species of Gymnophora have been published, and are the hypotheses to be tested in this paper. The new taxa that test these hypotheses are several previously undescribed species of Gymnophora from Japan and Nepal, sent to me by my colleague Dr. T. Got& Also, some character states of the male and female terminalia are reexamined to provide new data. Methods and materials

Terms and methods used are the same as those used by Brown (1987a, 1987b). As in previous works on this genus, I have refrained from describing females when the association with the corresponding male is questionable. Species of Gymnophora show little consistent variation, except in the male genitalia, and variations in such characters as costal length and halter color within and between species make association of males and females difficult. The best evidence that a male and female are conspecific is, of course, collecting them in copula or rearing them from a single female parent. Until such evidence is available, it is better not to describe the females. The formal classification is derived from a phylogenetic hypothesis

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of the relationships between species. As there are no formal categories for taxa at some levels, I have introduced my own terms, genus, subgenus, species group (e.g ., G. commotria-group) , species subgroup, species series, and species subseries. Relationships in the classification are also indicated by phylogenetic sequencing (Wiley 1981). Uncertain relationships are indicated by sedis mutabilis. Collections in which material is deposited are indicated as follows: BLK, Biological Laboratory, College of General Education, Kyushu University, Fukuoka 812, Japan (T. Saigusa); BVB, collection of the author; ELK, Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka 812, Japan (T. GotG). ZMK, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 150-164, 5300 Bonn 1, Federal Republic of Germany (H. Ulrich).

Taxonomy Genus Gymnophora Macquart, 1835 This genus was diagnosed and described previously (Brown 1987a). Diagnosis ~~~~~~l setae reduced, most individualsonly with postocellar and vertical setae; median furrow absent or rudimentary; anepisternum divided, setulose posterodorsally; legs without isolated setae or setal combs; wing veins R2+3 and k + s present; Dufour's mechanism Present in female; male terminalia withdrawn into segment 6, flexed 90" counterclockwise; female terminalia with internal sclerotized loop.

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Formal classification of the world species GENUS Gyrnnophora SUBGENUS Cerocratia (for list of species see Brown 1987b (Neotropical Region) SUBGENUS Gyrnnophora s. str. Gyrnnophora cornrnotria-group (for list of species see Brown 1987b (Neotropical Region) Gyrnnophora cymatoneura-' 'group" sedis rnutabilis Gyrnnophora priora n .sp . (Japan) Gyrnnophora spiracularis Borgmeier (Neotropical Region) Gyrnnophora cymatoneura Borgmeier (Neotropical Region) Sedis rnutabilis Gyrnnophora inexpectata Beyer sedis rnutabilis (Burma, Japan) Gyrnnophora quadriseta n. sp. sedis rnutabilis (Nepal) Gyrnnophora quartomollis-group Gyrnnophora nigripennis-subgroup sedis rnutabilis Gyrnnophora nepalensis n. sp. sedis rnutabilis (Nepal) Gyrnnophora fastigiorurn Schmitz sedis rnutabilis (Nearctic Region, Japan) Gyrnnophora quartornollis Schmitz sedis rnutabilis (widespread Palearctic Region) Gyrnnophora prolata-series Gyrnnophora nigripennis Schmitz (widespread Palearctic Region) Gyrnnophora prolata n .sp . (Japan) Gyrnnophora longissima n .sp . (Japan)

Sedis rnutabilis Gyrnnophora enigmata n. sp. sedis rnutabilis (Nepal) Gyrnnophora carina Brown sedis rnutabilis (Nearctic Region) Gyrnnophora arcuata-subgroup Gyrnnophora rnultipinnacula Brown (Japan) Gyrnnophora platypalpis n .sp . (Nepal) Gyrnnophora integralis-series Gyrnnophora gotoi n .sp . (Japan) Gyrnnophora pararcuata-subseries Gyrnnophora arcuata (Meigen) (widespread Palearctic Region) Gyrnnophora integralis Schmitz (widespread Palearctic Region) Gyrnnophora pararcuata n. sp. (Japan) Gyrnnophora subarcuata-series Gyrnnophora luteiventris-subseries Gyrnnophora setulata n. sp. (Nepal) Gyrnnophora luteiventris Schmitz (Nearctic Region) Gyrnnophora subarcuata Schmitz (Nearctic Region) Gyrnnophora healeyae-subseries (for list of species see Brown 1987a (Nearctic and Palearctic Regions) SUBGENUS Gyrnnophora Schmitz, 1929 Diagnosis Flagellomere 1 round, small; surstyli not enlarged, with 9 setae or less; aedeagus with curved anterior lobe (Brown 1987b).

Key to the species of Japanese Gymnophora 1.

Aedeagus reduced, without prominent dorsal projections in frontal view (see Brown 1987a, Figs. 11, 23, 24); basiphallus with prominent left lateral projection (Figs. 25 -28); epandrium with small posteroventral surstylar crest (Fig. 4); cercus short. Female with a single median ventral sclerite on segment 8 (Figs. 42, 43) .................................................................. .................................................................................. Gymnophora nigripennis-subgroup species (see key below) Aedeagus with dorsally projecting structures, basiphallus ring-shaped without long lateral projection; surstylar crest various; cercus long or short. Female with pair of ventral sclerites on segment 8 (Figs. 44 -46). ............................................... 2

2 (1). Male ................................................................................................................................................ 3 Female (females unknown for G. inexpectata, G. multipinnacula, and G. priora) ...................................................... 8 3 (2). Hypandrium with elongate, narrow process on left lobe (Fig. 41 ; Brown 1987a, Figs. 50, 5 1) ....................................... 4 Hypandrium without elongate process ......................................................................................................... 5 4 (3). Basiphallus in frontal view laterally compressed, thin (Brown 1987a, Fig. 13).................................. G. integralis Schmitz Basiphallus in frontal view rounded, thick (Fig. 22) .................................................................. G. pararcuata n.sp. 5 (3). Left side of epandrium with ventromedial process and greatly enlarged surstylar crest (Fig. 7). ....................... G. gotoi n.sp. Left side of epandrium without process; crest, when present, small ....................................................................... 6 6 (5). Left side of epandrium with small posteroventral crest (Brown 1987a, Fig. 16); inner left arm of aedeagus with several peaks (Brown 1987a, Fig. 22). ............................................................................................ G. multipinnacula Brown Left side of epandrium lacking crest (Figs. 1, 2); inner left arm without peaks .......................................................... 7 7 (6). Epandrium strongly inflated, with dorsomedial depression containing the short cercus; hypandrium without broad process on left lobe .......................................................................................................................... G. inexpectata Beyer Epandrium- hypandrium oval in frontal view, without dorsal depression housing cercus; cercus long; hypandrium with broad process on left lateral lobe ....................................................................................................... G. priora n.sp. 8 (2). Tergite 8 crescent-shaped, with few or no setae (Brown 1987a, Fig. 31); tergite 7 with setae concentrated anterolaterally (Brown 1987a, Fig. 41). ........................................................................................................... G. integralis Schmitz Tergite 8 larger, symmetrical (Figs. 36, 37); tergite 7 with scattered setae ............................................................... 9 9 (8). Tergite 8 rounded triangular (Fig. 36); venter of segment 8 medially with group of spinulae (Fig. 44). ............... G. gotoi n.sp. Tergite 8 elongate pentagonal (Fig. 37); venter of segment 8 without medial group of spinulae (Fig. 45). ............................. ................................................................................................................................. G. pararcuata n. sp.

BROWN

Key to the Nepalese Gymnophora Key includes males only; female known only in G. platypalpus.


1.

Aedeagus reduced, without prominent dorsal projections in frontal view (see Brown 1987a, Figs. 11,23,24); basiphallus with prominent left lateral projection (Figs. 25 -28); epandrium with small posteroventral, surstylar crest (Fig. 4); cercus short. Female with a single median ventral sclerite on segment 8 (Figs. 42, 43) .................................................................. .................................................................................. Gymnophora nigripennis-subgroup species (see key below) Aedeagus with dorsally projecting structures, basiphallus ring-shaped, without long lateral projection; surstylar crest various; cercus long or short. Female with pair of ventral sclerites on segment 8 (Figs. 44 -46) ................................................ 2

2 (1). Hypandrium with four setae ............................................................................................. G. q d r i s e t a n.sp. Hypandrium with two setae (Fig. 41) ........................................................................................................ 3 3 (2). Left side of epandrium with elongate ventromedial process (Fig. 9) .................................................. G. setulata n.sp. Left side of epandrium with process short, when present (Fig. 5) ........................................................................ 4 4 (3). Left side of epandrium with small, narrow, curved surstylar crest (Fig. 5) ......................................... G. enigmta n.sp. Left side of epandrium with broad surstylar crest (Fig. 6) .......................................................... G. platypalpis n.sp.

Key to the world species of the Gymnophora nigripennis-subgroup A separate key was deemed necessary to summarize the slight differences between the species of this most difficult group. 1.

Male .............................................................................................................................................. 2 Female (female G. nepalensis unknown) .................................................................................................... 7

2 (1). Lateral arm of aedeagus narrow (Fig. 25) ................................................................................................... 3 Lateral arm of aedeagus broad basally (Figs. 26 -28) ..................................................................................... 5 3 (2). Right surstylus enlarged (Brown 1987a, Fig. 47) ...................................... Gymnophora quartomollis Schmitz (Europe) Right surstylus small (Brown 1987a, Fig. 46) .............................................................................................. 4 4 (3). Lower arm of aedeagus, in ventral view, with medial tooth on left side (Brown 1987a, Fig. 52) ..................................... .............................................................................................. G. fastigiorum Schmitz (North America, Japan) Lower arm of aedeagus relatively parallel-sided (Fig. 25) ................................................ G. nepalensis n.sp. (Nepal) 5 (2). Lower arm of aedeagus, in ventral view, with medial tooth on left side (Fig. 28) ............... G. nigripennis Schmitz (Europe) Lower arm of aedeagus relatively parallel-sided (Figs. 26, 27) .......................................................................... 6 6 (5). Broad part of lateral arm longer than elongate part (Fig. 26) .................................................. G. prolata n.sp. (Japan) Broad part of lateral arm shorter than elongate part (Fig. 27) ............................................. G. longissim n.sp. (Japan) 7 (1). Abdominal tergite 3 absent; tergite 8 pentagonal (Brown 1987a, Fig. 34) ........................ G. nigripennis Schmitz (Europe) Abdominal tergite 3 present, reduced or of normal size; tergite 8 various .............................................................. 8 8 (7). Abdominal tergite 8 elongate -oval (Fig. 34); only tergite 4 absent or rudimental ........................ G. prolata n. sp. (Japan) Abdominal tergite 8 various, but not elongate-oval; tergite 3 usually reduced ........................................................ 9 9 (8). Abdominal tergite 7 thin, wedge-shaped (Brown,1987a, Fig. 40); tergite 3 reduced, tergite 4 absent, 5 normal .................... .............................................................................................................. G. quartomollis Schmitz (Europe) Abdominal tergite 7 broader, wedge-shaped (Figs. 29,30; Brown 1987a, Fig. 38); tergite 3 reduced, 4 absent, 5 reduced ...... 10 (9). Abdominal tergite 8 pentagonal (Fig. 35); tergite 7 rounded, triangular (Fig. 30) ..................... G. longissim n.sp. (Japan) Abdominal tergite 8 rounded, triangular (Brown 1987a, Fig. 29); tergite 7 hourglass-shaped (Brown 1987a, Fig. 38) ........... .............................................................................................. G. fastigiorum Schmitz (North America, Japan) -

Species descriptions

The between G. G. quadriseray the G. cymatoneura-group, and the rest of the species (the G. quartomollis-group) are unknown, as indicated by sedis mutabilis. inexpectata9

Gymnophora inexpectata Beyer, 1958 (Figs. 1, 10, 11)

Description (male only; female unknown) Body length 2.1 -2.5 mm. Mean frontal ratio 0.92. Mean costal length 0.42 wing length (no variation); mean costal sector ratio 7.6:2.8: 1, range 6.2 -9.0:2.0-3.5: 1. Costa not thickened. Legs and halter yellowish.

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-

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Male terminalia: Left side of epandrium inflated, with mediolateral setae and setulae, cercus in dorsomedial depression; surstylus broad, with sinuous margin (Fig. 1). Right side of epandrium with surstylus short, with several setae. Right arm of aedeagus with small posterior projection. Anterior lobe of distiphallus broad, with lateral arm, dorsal curved plate, and narrow- left sclerite (Figs. 10, 11). Lower arm short, thin. Lateral arm long, dorsal. HOLOTYPE (male): BURMA: Karnbaiti, 7000 ft (1 ft = 0.305 m), 11.V. 1934, Malaise trap (ZMK). PARATYPE (male): BURMA: Kambaiti, 2000 m, 4.VI. 1934, Malaise trap (ZMK). OTHER MATERIAL EXAMINED: Japan. 1 0 , Fukushima, Mt. Hiuchigatake, Hinoemata-mum, 1660 m, 23.VIII. 1980, T. Got6 (ELK).

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Gymnophora cymatoneura-group Diagnosis Terminalia markedly asymmetrical, more so than in other species. Epandrium with elongate subcercal process. Hypandrium with process on left lobe. Remarks This was formerly a phylogenetically defined species group, equal to other such taxa in rank. Because of the uncertainty of the relationships at the base of the subgenus Gymnophora s.str., however, it does not now strictly belong in this rank. It is still refered to as a group because it is a monophyletic taxon that will likely be found to belong at this level in the future, regardless of the eventual resolution of this basal area of the phylogeny.

Gymnophora priora n. sp. (Figs. 2, 12, 13) Description (male only; female unknown) Body length 2.0 -2.8 mm. Mean frontal ratio 0.94. Mean costal length 0.48 wing length, range 0.46 -0.50; mean costal Costa with sector ratio 6.4: 1.7:1, range 4.6-8.3:l.l-2.3:l. slight thickening before R,. Legs and costa yellowish. Male terminalia: Epandrium with long median process. Left side of epandrium with mediolateral setae; surstylus broad (Fig. 2). Right side of epandrium with surstylus moderately elongate, with several apical setae. Left lobe of hypandrium enlarged, deflecting right lobe and aedeagal guide counterclockwise, with short process. Right arm of aedeagus with anterior and posterior dorsal projections and left sclerite (Figs. 12, 13). Anterior lobe with small dorsal peaks. Lateral arm with basal expansion. Remarks This species is most easily recognized by the enlarged left hypandrial lobe. This is the first species of Gymnophora of the Holarctic Region to belong to a species group with Neotropical members. Derivation of speczjic epithet From the Latin for earlier or former, refeiring to the less derived structure of this species relative to that of the other members of the G. cymatoneura-group. H ~ L O T Y P E (male): JAPAN: Gifu, Hirayu, 12.X. 1982, 1300 m, T. Got6 (ELK) PARATYPE: JAPAN: 1 0 , Gunma, Ichinose-Sanpeitbge, 1420- 1760 m, Katashina-mura, 20.VIII. 1980, T. Got6 (ELK).

Gymnophora quadrisefa n. sp. (Figs. 3, 14, 15) Description (male only; female unknown) Body length 2.2-2.4 mm. Mean frontal ratio 0.88. Eye slightly reduced posteriorly. Mean costal length 0.51 wing length, range 0.51-0.52. Mean costal sector ratio 8.8:2.1:1, range 8.6 -9.0:2.0-2.2: 1. Costa with slight thickening before R,. Legs and coxae brown. Male terminalia: Epandrium without setulae; with deep posteromedial process. Left side of epandrium with surstylus slightly enlarged; with several posterolateral and one posterodorsal setae (Fig. 3). Left side of epandrium with several setae apically on surstylus. Each hypandrial lobe with large apical and small medial seta. Right arm of aedeagus extended dor-

sally into tall lobe (Figs. 14, 15). Outer left arm absent. Inner left arm peaked, with left lateral extension ending in small hook. Lateral arm short. Lower a r k broad. Remarks This species is easily recognized by the possession of four setae on the hypandrium, rather than the usual two. Derivation of specific epithet Named after the four setae on the hypandrium. HOLOTYPE (male): NEPAL: Dobang Kharka, 83"24'E, 28"36'N, 2400 m, 20.X.1971, Malaise trap, A. Nakanishi (BLK). PARATYPE:NEPAL: I 0 , Dobang Kharka, 83"24'E, 28"36'N, 2400 m, 26 -28.X. 1971, Malaise trap, A. Nakanishi (BLK) . Gymnophora quartomollis-group Diagnosis Epandrium with small to large surstylar crest on left side. Gymnophora nigripennis-subgroup Diagnosis Costa without abrupt thickening; male terminalia with cercus short; epandrium and hypandrium distinctive (Fig. 4); surstylar crest posteroventral; aedeagus reduced, basiphallus with left lateral process (Figs. 25-28); female with single sclerite with long setae ventrally on segment 8 (Figs. 42, 43).

Gymnophora enigmata n .sp . (Figs. 5, 16, 17) Description (male only; female unknown) Body length 2.2 mm. Frontal ratio 1.O. Costal length 0.42 wing length. Costal sector ratio 6.5: 1.3:1. Costa with abrupt thickening before R,. Legs and halter brown. Male terminalia: Epandrium with short posteromedial process. Left side of epandrium bare, with posterodorsal extension and several setae mediolaterally; surstylus broad, slightly extended, narrow epandrial process present posteroventrally (Fig. 5). Right side of epandrium shallow; surstylus elongate, setulose, with several setae. Right arm of aedeagus with dorsal projection and left lateral process separate (Figs. 16, 17). Anterior lobe tapered to thin apex laterally. Distiphallus broad. Lower arm bifurcate dorsally. Lateral arm broad with small anterior process. Derivation of specific epithet The name of this species reflects the enigmatic nature of its phylogenetic placement. HOLOTYPE (male): EAST NEPAL: Basantapur, 2300 m, 27"07'N, 87"24'E, 6-10.V.1972, Malaise trap (BLK). Gymnophora fastigiorum Schmitz This species, previously known only from North America (Brown 1987a), is the first to be found in both the Nearctic and Palearctic regions. MATERIAL EXAMINED: JAPAN:1 Q , Fukushima, Hinoematamum, Ozenuma, 1660 m, 21.VIII.1980, T. Got6 (ELK); 1 Q , Nagano, Azumi-mum, Shimashimadani, 1000- 1300 m, 14.X. 1982, T. Got6 (ELK); 11 0 , 15 Q , Yamanashi, Kitakoma-gun, Mount Kitadake, 2240 m, 27 .VIII. 1980, T. Got6 (ELK); 14 0 , 15 Q , Ashiyasu-mum, Kitazawat6ge, 2000 m, 22-24.VII.1981, T. Got8 (ELK).

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Gymnophora nepalensis n. sp. (Figs. 4, 24) Description (male only; female unknown) Body length 2.1 -2.3 mm. Mean frontal ratio 0.94. Mean costal length 0.45 wing length, range 0.42 -0.47; mean costal sector ratio 12.5:1.9: 1, range 8.3 - 14.6: 1.1-2.8: 1. Legs and halter brown. Male terminalia : Epandrium (Fig. 4), hypandrium, and aedeagus similar to those of other G. nigripennis-group species, except lateral arm of aedeagus not basally expanded or apically extended; lower arm without prominent lateral processes (Fig. 24).

Derivation of specific epithet This species is named after the country where is was collected. HOLOTYPE (male): NEPAL:Dobang Kharka, 2400 m, 83"24'E, 28"36'N, 25.IX. 1971, Malaise trap, A. Nakanishi (BLK) . PARATYPES: NEPAL:Dobang Kharka, 83"24'E, 28"36'N, 2400 m, 2 0,24.IX.1971,7 0,8.X.1971,6 0 , 10.X.1971, 3 0,19.X.1971,3 0 , 19-2O.X.1971,4 0,26- 28.X.1971, Malaise trap, A. Nakanishi (BLK, BVB); 2 0, Thudam, 27"45'N, 87"32'E, 21 -30.VI. 1972, 3500 m, Malaise trap (BLK) .

Remarks There were many female specimens that may have been conspecific with these males, but there was considerable variation among these females in shape and size of the tergites and sternites of the ovipositor. As these sclerites are normally the best taxonomic characters to use to differentiate between species, I could not determine whether one or several species were involved and I have left the females undescribed. Gymnophora prolata-series

Diagnosis Lateral arm or aedeagus broad basally.

Gymnophora prolata n. sp . (Figs. 26, 29, 34, 39, 42)

Description Body length 2.1 -2.3 mm. Mean frontal ratio 1.1. Female abdomen with tergite 4 absent or rudimentary; all other tergites present. Mean costal length 0.47 wing length, range 0.46 -0.49; mean costal sector ratio 8.3: 1.6:1, range 7.5 - 8.8: 1.0-2.0: 1. Legs and halter yellowish-brown. Male terminalia: Epandrium, hypandrium, and aedeagus similar to those of other G. nigripennis-group species (Figs. 4, 25-28; Brown 1987a, Figs. 5, 11, 17, 18,23,24,52,54,55), except lateral arm of aedeagus basally expanded, produced into long, thin distal point (Fig. 26). Female terminalia: Tergite 7 wedge-shaped (Fig. 29); sternite 7 thin, wedge-shaped. Tergite 8 elongate-oval (Fig. 34). Derivation of specific epithet From the Latin for extended, referring to the elongate lateral arm of the aedeagus. Remarks Most easily recognized by the elongate projection on the male lateral arm of the aedeagus. Like this species, the female of G. quartomollis lacks tergite 4, but G. quartomollis has tergite 3 reduced as well.

HOLOTYPE (male): JAPAN:Fukuoka, Mount Tachibana, 4.XII. 1976, T. Got8 (ELK). PARATYPES: 2 0 , 2 Q , same data as hblotype (BVB, ELK).

Gymnophora longissima n .sp. (Figs. 27, 30, 35, 43) Description Body length 1.7 -2.5 mm. Mean frontal ratio 0.94. Female abdomen with tergite 3 reduced, tergite 4 absent, tergite 5 reduced anteriorly. Mean costal length 0.49 wing length, range 0.48-0.51; mean costal sector ratio 7.5:1.9:1, range 6.8-9.5:1.3-3.0:l. Legs and halter yellowish. Male terminalia: Epandrium, hypandrium, and aedeagus similar to those of other G. nigripennis-group species (Figs. 4, 25-28; Brown 1987a, Figs. 5, 11, 17, 18,23,24,52,54,55), except lateral arm of aedeagus extremely elongate (Fig. 27). Female terminalia: Tergite (Fig. 30) and sternite 7 broad, wedge-shaped, with some variation in shape of tergite 7. Tergite 8 rounded triangular (Fig. 35). Derivation of specific epithet From the Latin for longest, referring to the extremely long lateral arm of the aedeagus. Remarks Tergite 7 of the female varies between specimens and it is possible that more than one species is represented. The association of the male and female specimens is based on their co-occurrence in trap samples. The best distinguishing characters are the elongate lateral arm of the male aedeagus and the distinctive sternite 7 of the female terminalia. HOLOTYPE (male): JAPAN: Tokushima, Mount Tsurugisan, Ichiu-mum, 1500 m, 16.X. 1980, T. Got8 (ELK). PARATYPES: JAPAN: 1 0, Gifu, Hirayu, 1300 m, 12.X.1982, T. Got8 (ELK); 6 o , 4 Q , Kumamoto, Mount Hakuchbzan, Izumi-mum, 1300 m, 4 -6.X. 1979, T. Got8 (BVB, ELK); 1 0 , 1 Q , Nagano, Mount Togakushi, Kamiminochi-gun, 1500 m, 17.X. 1982, T. Got8 (ELK); 1 o , 2 Q , Tokushima, Mount Tsurugisan, Ichiu-mum, 1500 m, 15- 17.X.1980, T. Got8 (ELK). Gymnophora spp. (G. nigripennis-subgroup) Four female specimens of an unknown species were collected in Hokkaido. I have left these undescribed until females of this species have been collected with male specimens. Also, three female specimens from Gunma Prefecture, Japan, and at least three "species" from Nepal have been left undescribed. The limits of species and the amount of variation between female specimens of the same species are unknown in this group. Unfortunately, this often makes it impossible to associate ,the females with the males, especially as more than one species can be found in a single locality. Gymnophora arcuczta-subgroup Diagnosis Outer left arm of aedeagus separate from right arm.

Gymnophora platypalpis n. sp. (Figs. 6, 18, 19, 33, 38, 46, 49) Description Body length 2.3 mm. Frontal ratio 0.96. Eyes reduced. Palpi enlarged, elongate. Costal length 0.55 wing length. Costal


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BROWN


sector ratio 10.8:2.8: 1. Costa slightly thickened along entire length, without abrupt swelling. Fork of Rs acute. Legs and halter brown. Male terminalia: Epandrium with moderately deep, elongate posteromedial process. Left side of epandrium dorsally setulose; surstylus broad, ventral, slightly developed; several setae present posterodorsally; flat posteroventral lobe present (Fig. 6). Right side of epandrium posteriorly setulose; surstylus elongate, narrow, with several setae. Left hypandrial lobe with short broad process. Basiphallus with small, ventral, left lateral projection. Right arm with thin, elongate dorsal extension with thin apical hook (Figs. 18- 19). Inner left arm with elongate peak, ending abruptly at articulation with outer left arm. Outer left arm bifurcate at both ends; anteriorly, dorsal furcation articulates with apex of inner left arm, vent& furcation with basiphallus process (Fig. 49). Distiphallus broad, membranous, with broad apical hook. Lateral arm elongate. Lower arm broad. Female terminalia: Tergite 7 wedge-shaped (Fig. 33); sternite 7 a thin strip. Tergite 8 rounded triangular; venter of segment 8 bilobed, with large apical sclerites (Fig. 46). Internal sclerotized ring thickened ventrally. Derivation of specific epithet The name reflects the enlarged palpi of this species.

Remarks The association of this male and female needs to be verified by further collections, as they are from different localities. Both have the distinctive reduced eyes and enlarged palpi. HOLOTYPE (male): NEPAL:Thudam, 3500 m, 27"45'N, 87"32'E, 21 - 30.VI. 1972, Malaise trap (BLK). PARATYPE: NEPAL: 1 Q , Dobang Kharka, 2400 m, 83"24'E, 2g036'N, 27.X. 1971, A. Nakanishi (BLK).

lobe unmodified. Basiphallus broad dorsally. Right arm lacking ventral knob; dorsal extension in three sections: right anterodorsal sclerite, left anterodorsal curved sclerite (homologue to outer left arm, as shown by connection to basiphallus), and large posterodorsal extension. Inner left arm present, ending in small lateral lobe (Figs. 20, 21). Lateral arm long, narrow. Lower arm with large flat plate attached. Female terminalia: Tergite 7 parallel-sided, anterior onefifth slightly widened; with trapezoidal anterior extension on intersegment 6-7 (Fig. 31). Sternite 7 thin, posteriorly pointed. Tergite 8 long, rounded, wedge-shaped (Fig. 36). Segment 8 ventrally with median patch of spinulae and sclerotized lateral lobes (sternite 8) (Fig. 44). Sclerotized ring not seen. Derivation of specific epithet Named after the Japanese phoridologist Dr. T. Got6, who collected and allowed me to study this material. HOLOTYPE (male): JAPAN: Tottori, Mount Daisen, 7.VIII. 1978, T. Got6 (ELK). PARATYPES: JAPAN:1 0 , 2 Q , Tottori, Mount Daisen, 7.VIII.1978, T. Got6 (ELK, BVB). Gymnophora pararcuata-subseries Diagnosis Hypandrial process elongate.

Diagnosis Left hypandrial lobe with process. Anterior lobe of distiphallus broad, lightly sclerotized. Female with bent internal sclerotized loop (in three of four species).

Gymnophora integralis Schmitz 1920 This widespread European species is recorded for the first time from Japan. MATERIAL EXAMINED: JAPAN:Fukuoka, Mount Hikosan, 1 Q , 4.XII.1968,2 0 , 1 Q , 23.1.1969,4 0 , 1 Q , 12.11.1969, 2 0 , 1 Q,3.IV.1969, 1 0 , 9 . I V . 1 9 6 9 , 3 Q , 12.1V.1969, 1 Q , 22.XII.1969, 2 0 , 27.111.1970, 1 Q , 8.1V.1970, 1 Q , 18.XI.1970, 1 0 , 12.1V.1971, 1 0 , 12.1V.1972, 1 ~ , 2 8 29.IV. 1972, 1 0 , 2 4 . V . 1972, 1 0 , 24.IV. 1979, K. Takeno, Malaise trap (BVB, ELK); Gifu, Hirayu, 1300 m, 1 0 , 12.X. 1982, T. Got6 (ELK); Tokushima, Mount Tsurugisan, Ichiumum, 1500 m, 2 0 , 3 Q , 15-16.X.1980, T. Got6 (ELK).

Gymnophora gotoi n. sp . (Figs. 7, 20, 21, 31, 36, 44)

Gymnophora pararcuata n. sp . (Figs. 8, 22, 32, 37, 40, 41, 45)

Description Body length 2.2 - 3.0 mm. Mean frontal ratio 1.1. Female with tergites 1 and 2 normal (short, wide darkly sclerotized), 3 vestigial (light sclerotization in the center of the segment), 4 absent, 5 with sclerite-shaped patch of thin membrane, 6 rounded square. Mean costal length 0.56 wing length, range 0.54-0.57. Mean costal sector ratio 8.3:2.3:1, range 7.010.5: 1.7 -3.3: 1. Costa slightly thickened proximal to R, . Legs and halter yellow. Male terminalia: Epandrium with small median process below cercus. Left side of epandrium bare; surstylus broad, elongate; small, narrow posteroventral process present; several setae present laterally (Fig. 7). Right side of epandrium bare; surstylus narrow, elongate, with several ventral setae. Left hypandrial lobe with small, broad process; right hypandrial

Description Body length 2.5 - 3.5 mm. Mean frontal ratio 0.95. Female with abdominal tergite 2 trapezoidal; tergite 3 absent or reduced, lightly sclerotized; tergites 4 and 5 absent; tergite 6 large, wedge-shaped. Female abdomen with pair of sclerotized dorsal glands between segments 5 and 6. Mean costal length 0.56 wing length, range 0.54 -0.58. Mean costal sector ratio 9.5:2.3: 1, range 5.4- 11.2:1.3-3.0: 1. Costa with slight thickening before R,. Fork of Rs acute. Legs yellowishbrown. Halter yellow. Male terminalia: Left side of epandrium with small posterodorsal lobe; surstylus broad, with several setae posterodorsally (Fig. 8). Right side of epandrium with surstylus small, with several apical setae. Left hypandrial lobe bare, with process (Fig. 4 1); small (sensory?) pits present posterolaterally. Right

Gymnophora integralis-series

-

-

-

FIGS.1-9. Epandria, left lateral view. Scale bar = 0.1 rnm. Fig. 1. Gymnophora inexpectata. Fig. 2 . G. p r i o r ~ .Fig. 3. G. quadriseta. Fig. 4. G. nepalensis. Fig. 5. G. enigmata. Fig. 6 . G. platypalpis. Fig. 7. G. gotoi. Fig. 8. G. pararcuata. Fig. 9. G. setulata. c , surstylar crest; v, ventral epandrial process.

Can. J. Zool. Downloaded from www.nrcresearchpress.com by "Institute of Vertebrate Paleontology and Paleoanthropology,CAS" on 06/06/13 For personal use only. 2550 CAN. J. ZOOL. VOL. 67,1989

BROWN


hypandrial lobe setulose. Basiphallus with dorsal and left lateral expansions (Fig. 22). Anterior lobe broad, curved backwards. Possible supra-aedeagal sclerite present on right side. Distiphallus with heavily sclerotized, curved base. Right arm heavily sclerotized. Outer left arm present. Lower arm broad. Lateral am narrow. Cercus long. Female terminalia: Tergite 7 wedge-shaped, anteriorly sinuous (Fig. 32). Tergite 8 triangular, with scattered setae (Fig. 37). Segment 8 ventrally with subequal, lateral lobes and with long sclerite (Fig. 45). Thickening of vaginal wall ringshaped, bent (Fig. 40). Derivation of specijc epithet The name reflects the similarity of this species to European G. arcuata (Meigen). HOLOTYPE (male): JAPAN:Fukuoka, Mount Hikosan, 20-22 .XI. 1972, K. Takeno leg (ELK) PARATYPES: JAPAN:Fukuoka, Mount Hikosan, 1 Q , 19.X.1968, 1 0 , 29.X.1968, 1 Q , 30.X.1968, 1 Q , 14.XI.1968, 1 Q, 13.XI.1969, l o , 12.XI.1970, l o , 1 Q , 2-4.XI.1971, 1 Q , 22.XI.1971, 1 Q , 20-22.XI.1972, 1 0 , 31.X. 1973, 1 0 , 13.XI. 1973, K. Takeno leg. Malaise trap (BVB, ELK); Kumamoto, 1 0 , Mount Hakuchozan, Izumimum, 1300 m, 6.X. 1979, T. Got8 (ELK). Gymnophora subarcuata-series

Diagnosis Epandrium with prominent left ventrolateral process. Left surstylar crest posterodorsal. Anterior lobe of aedeagus peaked behind basiphallus. Gymnophora luteiventris-subseries

Diagnosis Epandrium with setulae (modified to spinuli in two species) on left side.

Gymnophora setulata n .sp. (Figs. 9, 23, 24)

Description Body length 2.4 -2.6 mm. Mean frontal ratio 0.94. 'Mean costal length 0.46 wing length, range 0.45 -0.48. Mean costal sector ratio 5.6:1.1:1, range 2.7-8.6:0.8-1.4:l. Costa with slight thickening before R,. Legs and halter yellowish. Male terminalia: Epandrium with elongate median process. Left side of epandrium setulose mediolaterally; surstylus small, crest-shaped; epandrial process narrow, elongate; flattened, dark, posteroventral lobe present; several setae present posterodorsally (Fig. 9). Right side of epandrium setulose; surstylus slightly elongate, with few setae. Basiphallus with short left ventrolateral process. Outer left am broad; origin extends from basiphallus process to basiphallus vertex (Figs. 23, 24). Inner left arm with small right lateral peak; extended across left side of aedeagus. Supra-aedeagal sclerite elongate, broadly bifurcate at apex. Distiphallus apically broadened, not twisted. Lateral and lower arms broad.

255 1

Remarks This species clearly belongs with G. luteiventris and subarcuata, both of which are found only in the Nearctic Region. Derivation of specijc epithet Named after the setulae on the left side of the epandrium. HOLOTYPE (male) : NEPAL:Dobang Kharka, 83"24'E, 28"36'N, 2400 m, 26-28.X.1971, Malaise trap, A. Nakanishi (BLK). PARATYPE: 1 0 , same data as for holotype (BLK).

Phylogeny The previous phylogenetic hypotheses for this genus (Brown 1987a, 1987b) were inadequate. The following synapotypies, proposed previously, were refuted or had to be reinterpreted on the basis of the new evidence: in Brown (1987a), 1, 2, 3, 9 (Fig. 50a); in Brown (1987b), 19 (partially), 20 (Fig. 50b). The major groups of the genus Gymnophora are discussed below, and their synapotypies shown in the cladograrns (Figs. 51 -54). Postulated apotypic character states Note: Synapotypies are not given for groups not discussed in this paper, but are in previous publications (Brown 1987a, 1987b). 1. Curved anterior lobe present The Gymnophora cornmotria-group is not treated here (see Brown 1987b). 2. Epandrial setulae reduced to posterodorsal comer on left side According to the previous hypothesis of relationship (Fig. 50b), the Gymnophora cymatoneura-group belonged to a monophyletic group with the G. quartomollis-group. New research shows that the "swelling of the left side of the basiphallus" (Brown 1987b), linking the G. cyrnatoneuragroup to the G. quartomollis-group, may in fact be a primitive character shared by Anevrina Lioy , Crinophleba Borgmeier, and other phorid genera. Also, the former G. carina-group (Fig. 50b) was found to be paraphyletic, belonging within the G. quartomollis-group. Based on the existing information, I cannot resolve the relationships at the base of the subgenus Gymnophora, other than to note that the G. cornmotria-group is the sister-group to all of the rest of the species. An unresolved quadrichotomy is represented by the G. cymatoneura-group, G. quadriseta, G. inexpectata, and the remainder of the species. The species of the G. cyrnatoneura-group, G. inexpectata, and G. quadriseta exhibit many plesiotypic character states in their aedeagi, being similar to the species of the G. commotria group in form. This node of the tree is the area most in need of further resolution. 3. Hypandrium with left process (Brown 1987b, Fig. 105) 4. Genitalia markedly asymmetrical (Brown 1987b, Figs. 97- 100) 5. Subcercal epandrial process elongate (Brown 1987b, Figs. 97- 100)

FIGS. 10 -24. Aedeagi. Scale bars = 0.1 mrn (Figs. 10- 15, 18 -21, 23, and 24 to same scale; Figs. 16 and 17 to same scale). Fig. 10. Gymnophora inexpectata, frontal view. Fig. 1 1 . G. inexpectata, left lateral view. Fig. 12. G. pnora, frontal view. Fig. 13. G. pnora, left lateral view. Fig. 14. G. quudriseta, frontal view. Fig. 15. G. quudrista, left lateral view. Fig. 16. G. enigmata, frontal view. Fig. 17. G. enigmata, left lateral view. Fig. 18. G. platypalpis, frontal view. Fig. 19. G. platypalpis, left lateral view. Fig. 20. G. gotoi, frontal view. Fig. 2 1. G. gotoi left lateral view. Fig. 22. G. pararcuata, frontal view. Fig. 23. G. setulata, frontal view. Fig. 24. G. setulata, left lateral view. a , anterior lobe of distiphallus; 6 , basiphallus.


CAN. J. ZOOL. VOL. 67, 1989

FIGS.25 -41. Male and female terminalia. Scale bars = 0.1 rnm (Figs. 25 -40 to same scale). Figs. 25 -28. Aedeagi, ventral. Fig. 25. Gymnophora nepalensis; Fig. 26. G. prolata. Fig. 27. G. longissima. Fig. 28. G. nigripennis. Figs. 29 -33. Female tergite 7. Fig. 29. G. prolata. Fig. 30. G. longissima. Fig. 31. G. gotoi. Fig. 32. G. pararcuata. Fig. 33. G. platypalpis (showing details of segments 6 and 7). Figs. 34 -38. Female tergite 8. Fig. 34. G. prolata. Fig. 35. G. longissima. Fig. 36. G. gotoi. Fig. 37. G. pararcuata. Fig. 38. G. platypalpis. Figs. 39 and 40. Internal sclerotized ring of female. Fig. 39. G. prolata. Fig. 40. G. pararcuata. Fig. 41. G. pararcuata, hypandrium, ventral. hp, hypandrial process; lp, lateral process of basiphallus; t, lateral arm of aedeagus; w , lower arm of aedeagus; wt, tooth of lower arm; 6, 7, abdominal segments 6 and 7.


BROWN

FIGS.42 -49. Female terrninalia and male aedeagi. Scale bars = 0.1 rnrn (Figs. 42 -46 to same scale; Figs. 47 -49 to same scale). Figs. 42 -46. Venter of apex of female segment 8. Fig. 42. Gymnophoraprolata. Fig. 43. G. longissirnu. Fig. 44. G. gotoi. Fig. 45. G. pararcuata. Fig. 46. G. platypalpis. Figs. 47 -49. Aedeagi, dorsolateral (oblique) view. Fig. 47. G. carina. Fig. 48. G. multipinnacula. Fig. 49. G. platypalpis.f, fusion of curved plate to basiphallus; o, outer left arm; s, separation point of outer left arm and right arm.

These three characters unite G. priora with the G. cymatoneura-group species G. spiracularis and G. cymatoneura. The terminalia of G. inexpectata are also somewhat similar, but are so highly modified that it is difficult to tell if this is a synapotypy or a by-product of other structural differences. The structure of the aedeagus is uninformative on this matter, but G. inexpectata may be better placed in the G. cymatoneura-group as well. 6. Subcercal epandrial process apically broadened, spoonshaped (Brown 1987b, Figs. 97, 98) 7. Surstylar crest present (Figs. 4, 5, 7) The rest of the species, including the two members of the former G. carina-group (Fig. 50b), belong within the G. quartomollis-group. Gymnophora enigmata may belong in the G. nigripennis-subgroup, as the epandrium of this species and those of the subgroup are very similar, but there are no characters to indicate whether this similarity is derived or primitive. The aedeagus of G. enigmata is at a G. commotriagroup grade of development. The relationship of G. carina to the rest of the species is unknown, but its aedeagus is similar in organization to those of the G. commotria-group species. The species of the G. integralis-series and G. platypalpis lack a discrete, small surstylar crest. That of G. platypalpis is broader, but still recognizable, whereas that in G. integralis and gotoi is enlarged, and in G. arcuata and G. pararcuata it is apparently lost. It is possible that synapotypy 7 is misapplied here, and that the G. integralis-series and G. platypalpis are

not members of a group with the other G. quartomollis-group, species, but evidence from the aedeagus disputes this. 8. Female segment 8 ventrally, with a single sclerite (Figs. 42, 43) 9. Lateral arm of aedeagus broad (t in Figs. 26 -28) 10. Lateral arm of aedeagus with elongate projection (Figs. 26, 27) Within the G. nigripennis-subgroup, there are no characters apparent to link any of the other species with each other or with the newly named G. prolata-series. 11. Outer left arm of aedeagus attached to basiphallus Cf in Fig. 48) Gymnophora multpinnacula is the first species to show this distinctive character. The attachment is very narrow, and is part of a large curved plate on the outer left arm. The outer left arm itself has not yet separated from the dorsal extension of the right arm of the aedeagus. 12a. Outer left arm partially separate from right arm (s in Fig. 49) 12b. Outer left arm completely separate from right arm (Fig. 24) Species of the G. commotria-group have a lateral sclerite (see Brown 1987b), and G. canna has a sclerite that may be the precursor of the outer left arm (o in Fig. 47). The separation of the two "arms," however, is first shown in G. platypalpis (Fig. 49). This species is somewhat enigmatic, as I could not find a connection between the basiphallus and the

CAN. J . ZOOL. VOL. 67. 1989


Previous Hypotheses carina multipinnacula quartomollis nigripennis fastigiorum arcuata integralis subarcuata luteiventris healeyae marshalli talea

corino- group

t -I I Cerocratia

commotria-group

W . 1:

Y

19

21-22

carina-group (synapotypies in Brown 1987a)

&--cymatoneura-group quartomollis-group

FIG.50. Previous hypotheses of phylogenetic relationships. (a) Relationships of the Holarctic Region species of Gymnophora. (b) Relationships of Holarctic and Neotropical region species groups of Gymnophora.

outer left arm (synapotypy 10). I regard it as possible that the connection is either a thin membrane, or it was broken during the handling of the single male specimen. 13. Hypandrium with process on left lobe (hp in Fig. 4 1). The aedeagus of G. gotoi is similar to that of G. multipinnacula in that the attachment of ,theouter left arm to ,thebasiphallus is via a large curved plate. This is problematic, because the outer left arm of G. platypalpis is more darkly sclerotized and formed like that of the G. subarcuata-series and the rest of the

G. integralis-series rather than like that of G. gotoi. 14. Hypandrial process elongate 15. Anterior lobe expanded, tongue-shaped (a in Fig. 22) 16. Costal swelling of wing with small clear spot 17. Sclerotized internal ring of female bent (Fig. 40) The three species G. arcuata, G. pararcuata, and G. integralis form a clearly defined group, here designated as the G. pararcuata-subseries. Synapotypy 17 may be a synapotypy of the entire G. integralis-series, but I was unable to find a

BROWN

commotria - group


I

I ?

:

I

I

1 1

cyrnatoneura - group (see Fig. 52) inexpectata quadriseta nigripennis - subgroup (see Fig. 53)

1

multipinnacula platypalpis integralis - series (see Fig. 54)

luteiventris

subarcuata

19-22

I healeyae - subseries (see Brown 1987a)

FIG.51. Hypothesized relationships of species and species groups within the Gymnophora commotria, G. cymatoneura, and G. quartomollisgroups.

sclerotized ring in either of the female specimens of G. gotoi. 18. Paired glands in female abdomen sclerotized or darkcolored This character links G. integralis and G. pararcuata. The marked similarity in epandrial and aedeagal structure between G. pararcuata and G. arcuata is thus shown to be primitive, as were the similarities between G. subarcuata Schmitz and G. healeyae Disney (Brown 1987~). 19. Left side of epandrium with ventrolateral process (v in Fig. 9) The ventral process of the epandrium is found also in the

problematic G. gotoi, and perhaps also in G. arcuata (Brown 1987a, Fig. 3). This synapotypy may be more appropriately placed at the base of the group formed by the G. integralisseries plus the G. subarcuata-series (as shown in Fig. 51). 20. Surstylar crest postemdorsal (Fig. 9) 21. Anterior lobe (= inner left arm of Brown 1987a) peaked behind basiphallus 22. Supra-aedeagal sclerite (remnant of dorsal extension of right arm) present This group (the G. subarcuata-series) was discussed previously (Brown 1987~).

CAN. J. ZOOL. VOL. 67, 1989

gotoi

priora

+k

+-I

cymatoneura


3-5

6

I 7 arcuata

spiracularis

.

integralis

14-17

FIG. 52. Hypothesized relationships within the Gymnophora cyrnatoneura-group.

18

pararcuata subseries

pararcuata

FIG. 54. Hypothesized relationships within the Gymnophora integralis-series.

nepalensis

such as reproductive tracts, will provide further clues to the phylogeny of this genus.

fastigiorum quartomollis

7 nigripennis

prolata series

FIG. 53. Hypothesized relationships within the Gymnophora nigripennis-subgroup.

23. Left side of epandrium with lateral setulae (Fig. 9) The epandrial setulae, reduced in most species of northern hemisphere Gymnophora (synapotypy 2) have reappeared in G. setulata, G. luteiventris, and G. subarcuata, newly named the G. luteiventris-subseries. 24. Setulae of left side of epandrium flattened, spinulose (see Brown 1987a, Fig. 73)

Test of the phylogeny As the title of this paper indicates, the new species are to be used as a test of the previously reconstructed phylogeny. The new species fit into three categories: 1. Those that fit into the previously established groups, thus corroborating them, with few or no new character states needed. These species are G. nepalensis, G. prolata, G. longissimu (in the G. nigripennis-subgroup), G. pararcuata (G. integralisseries), and G. setulata (G. luteiventris-subseries). These groups are those most strongly supported by new data. 2. Those that formed recognizable sister-taxa to previously established groups with some modification, giving some support to previous groups. These are G. priora (G. cymutoneuragroup), G. gotoi (G. integralis-series), and G. platypalpis. Gymnophora platypalpis forced the reassessment of the characters of G. carina and G. multipinnacula. These groups are moderately supported. 3. Those that did not fit in any of the previously established groups. Gymnophora quadriseta and G. enigmuta belong to this group and point to continuing instability and uncertainty of the relationships at the base of the subgenus Gymnophora s.str. The obvious pattern is that the further from the base of the group one proceeds, the greater the stability of the cladogram. Also, most species from Japan fit into already established lineages (except G. inexpectata), whereas most of those from Nepal are more difficult to place. Hopefully, in the future, character states found in immature stages, female specimens (which are unknown for many taxa), and internal structures

Evolutionary aspects of characters used Unfortunately, few characters in this analysis were primary synapotypies, i. e., those found only in the groups in question. Many of the characters, such as hypandrial processes, occur in several groups of Gymnophora. Another interesting observation is that most of the characters used involving the epandrium -hypandrium were found on the left side. As this side is functionally ventral (adult male Gymnophora have their terminalia derotated 90°), it is likely to be the part of the terminalia that contacts the female during copulation. Thus, it would be the part of ,the terminalia most prone to develop sexually selected characters. Diversity of Gymnophora As a final note, it should be mentioned that since the start of my work on Gymnophora, the number of known species in this genus has grown from 23 to 53. There is no end in sight for new species, including several from Japan and Nepal that were represented only by female specimens. I believe that 65 -80 world species could exist, or approximately three to four times the number of previously described species. If these numbers are extrapolated to other phorid genera, many of which are probably even less completely known than Gymnophora, the number of living phorid flies is much larger than is currently recognized. R. H. L. Disney's statement (personal communication) that we know of less than one-quarter of the living species is probably close to the truth. Acknowledgments I would like to thank Dr. T. Got6 for sending me the specimens of Japanese and Nepalese Gymnophora, and Dr. G. E. Ball for reviewing this manuscript. I am also grateful to two anonymous reviewers who greatly improved the quality of this paper. This study was funded in part by Natural Sciences and Engineering Research Council of Canada (NSERC) grant A- 1399 to G. E. Ball and an NSERC postgraduate scholarship to the author. BEYER,E. M. 1958. Die ersten Phoriden von Burma. Soc. Sci. Fennica, Cornm. Biol. 18: 1-72. BROWN, B. V. 1987a. Revision of the Gymnophora (Diptera: Phoridae) of the Holarctic Region: classification, reconstructed phylogeny and geographic history. Syst. Entomol. 12: 271 -304. 1987b. Classification, reconstructed phylogeny and geographic history of the Neotropical phorid flies of the genus Gymnophora (Diptera: Phoridae). J. Nat. Hist. 21: 1477 - 1524. HENNIG,W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana. WILEY,E. 0. 1981. Phylogenetics: the theory and practice of phylogenetic systematics. John Wiley & Sons, New York.