© Entomologica Fennica. 29. VI.1990

A striking new species of Megaselia (Diptera, Phoridae) from Sulawesi, with re-evaluation of related genera

R. H. L. Disney

Disney, R. H. L. 1990: A striking new species of Megaselia (Diptera, Phoridae) from Sulawesi, with re-evaluation of related genera. - Entomol. Fennica I :25-31. Megaselia torautensis sp. n. from Sulawesi is described. In the li ght of its peculiar features the genu s Megaselia is rev iewed and the following Afrotropi­ cal genera synonymised with it: Epimegaselia Beyer, Metaplastophora Beyer, Quasipseudacteon Beyer, and Tarsophoromyia Beyer. The species included in these genera are related to the rest of Megaselia by indicating where they will key out in the existing keys to Afrotropical Megaselia. R. H. L. Disney, Field Studies Council Research Fellow, University Depart­ ment of Zoology, Downing Street, Cambridge, CB2 3£1, England

Traditionally the classification of species (i.e. the 1983b), taxonomists can now truly claim to be grouping of species into species-groups, subgen­ engaged in science. Only in the solving of nomen­ era, genera, tribes, etc) has been based on a straight clature problems is their task more akin to that of forward typological/phenetic sorting. This, how­ a lawyer than a scientist, in that precedence (the ever, has tended to give rise to classifications ever principle of priority) and typification are th e key at ri sk of revision as the morphological gaps to aniving at a correct solution, rather than the between supra-specific taxa are bridged by newly critical evaluation of biological data. di scovered species. With the best estimates indi­ The research programme that seeks to group cating that only about 10% of the world's fauna species into a hierarchical scheme of monophyletic has been described and named by taxonomists, the taxa may be causing a certain amount of instabil­ prediction must be that the majority of supra­ ity in the short term, but in the longer term the specific taxa proposed by traditional taxonomy resulting classification will more readily accom­ must be at risk of revision. modate new species without the necessity for While the Darwinian revolution initiated the constant revisions of supra-specific taxa. While process of freeing taxonomy from the dead hand there is frequently controversy regarding the cor­ of authoritative opinion and arbitrarily imposed rectness of a claim to have identified a mono­ classifications, it has required the clarifications of phyletic group, such a claim constitutes a legiti­ the Hennigian research programme to complete mate scientific hypothesis. As with all science the process. Now, in principle, biological classifi­ such hypotheses cannot be proved. They can, cation is an experimental science, which is seek­ however, be disproved; or at least the proposed ing to discover the correct classification through a autapomorphy of the characters, on which the process of advancing testable hypotheses. Phylo­ hypotheses of monophyly are based, can be di s­ genetic cladistics is truly heuristic. Such taxon­ proved by demonstration of homoplasy or re­ omy can no longer be likened to biological philat­ versed polarity of the homologous transformation ely. In four, of their five major tasks (see Disney series. 26 Disney: A new species of Megaselia, Phoridae • ENTOMOL. FENNICA Vol. l

The giant genus Megaselia Rondani currently It is against the above background that a strik­ includes nearl y 1400 valid species. It was previ­ ing new species from Sulawesi is described. It is ously estimated, from the rati os of known to assigned to the genus Megaselia. By so doing the unknown species in collections sent to the author, validity of several genera is called into questi on. that the true total lies between 5000 and 20 000 The synonymising of some Afrotropical genera species (Disney l983a). Subsequent experi ence with Megaselia is proposed. Such action renders suggests the higher fi gure is more likely to be the latter genus even more un wieldy. However it nearer the truth. Confronted with such an unman­ is considered that the priority with the Megaseliini ageable mass of species there has been a strong is to construct user-friendly identification keys to temptation to lop off morphological segregates recognisable genera and their included species, as and to designate these as separate genera. With the a first step in making progress in the classification description of new species, however, the gaps of thi s mass of species. Segregates should onl y be between such genera and the parent genus Mega­ removed and pl aced in separate genera when a selia have tended to be bridged again . Thus both case for their monophyly has been proposed and, Endonepenthia Schmitz and Plastophora Brues at the same time, their pl acement in a subgenus have had to be abandoned, and their incl uded been shown to be inappropriate on scientific species returned toMeg aselia (Disney 1978, 198 1, grounds. 1986a). A further complication confronts one when considering the legitimacy of the genus Megase­ M egaselia torautensis sp. n. lia and related genera. The ground plan of Mega­ Figs. l-4 selia appears to be close to the ground plan of the Phoridae in many respects (Disney 1988). Princi­ Type materi al: Holotype #, Sulawesi-utara, Dumoga­ pal proposed apomorphic features are a mesopleu­ Bone Nati onal Park, Toraut Forest, Ill.l 985 (R. H. L. Disney) (in Cambri dge Uni versity Zoology Museum). ral furrow, which serves to characterise the Me­ Paratypes: I #, 2oo same data as holotype except # topininae, and the Dufour's crop mechanism in 30-3 1.1.1 985, and oo 23. VII-3. YIII.I 985 (A. H. Kirk­ females, which serves to characteri se the Megase­ Spriggs) ( I o in National Museum of Wales, Cardiff) liini (Disney 1989). Within the Megaseliini the Diagnosis: The combination of two longitudi­ characterisation of the genera is far from being in nal hair palisades on the mid-tibia, an unforked terms of autapomorhic features. Even in terms of vein 3, two bristles and two hairs on the scutellum, morphological gaps many of the proposed genera and fine hairs and a pair of strong bristl es on the are so poorly characterised that they cannot be mesopleuron, apart from distinctive male and separated from Megaselia when attempting to female abdominal terminalia, will serve to distin­ construct a dichotomous key to genera, except guish thi s species from all other described Meg­ when dealing with the fauna of a limited region. aselia species except M. bruesi Disney, which has Even in the latter case the description of new pale yellow halteres and only very short hairs on species has tended to erode the disti nctions pro­ abdominal tergites l-5. posed. Such a situation is only tolerable if one is confident that one is dealing with monophyletic Male: Frons about as wide as long, bro wn and genera. Where this is not yet the case then prag­ with 70--80 hairs. Lower supra-antenna! bristles a mati c criteria, such as designation of genera in little shorter, weaker, and closer together than terms of clear cut morphological di stinctions, can upper pair, which are situated about the same level be the only justification for maintaining parts of as the anti a! bris tles. The latter are clearly closer to Megaselia s. l. as separate genera. the supra-antennals than to the antero-laterals, An additional complicati on in the Phoridae is which are clearly higher on frons. In one specimen that more than 50% of the genera (out of a current there is a weaker supernumerary supra-antenna] total around 235) are only known in one sex. Even bristle above the left upper supra-antenna!. Pre­ when the female is known the past reli ance on ocellars a little further apart than upper supra­ pinned specimens means it is frequently not known antennals, but closer to each other than either is whether Dufour's crop mechanism is present or from a medi o-lateral bristle. The latter clearly absent. lower on frons. Third antenna] segment pale brown, ENTOMOL. FENNICA Vol. I • Disney: A new species of Megaselia, Phoridae 27

Figs. 3-4. Megaselia torautensis sp. n.- 3: anterior face of male mid tibia;- 4: dorsal view of segments 5 and 6 of female abdomen. - Scale bars = 0.1 mm .

Front legs yellowish grey. Middle and hind legs more brownish, with apical third of hind femur progressively browner. With 5-7 of the hairs below basal half of hind fe mur as long as or Figs . 1-2. Megaselia torautensis sp. n. - 1: male clearly longer than those of antero-ventral row in hypopygium viewed from left side ; - 2: female ovi ­ positor viewed from left side. - Scale bars = 0.1 mm . apical half. Hind tibia with 12-18 postero-dorsal hairs, with those in lower half more robust and spine-like. Mid-tibia with both a dorsal and an anterior longitudinal hair palisade (Fig. 3). Fore spherical to ovoid, with a dorsal, pre-apical, shor­ tarsus with a postero-dorsal hair palisade on all tish-haired, brown arista. Palps pale brownish five segments. The segments are long and slender, with 4--5 apical bristles, the longest of which is no especially the metatarsus (the length rati os being longer than greatest width of palp. Proboscis with 3.0 : 1.4: 1.0 : 0.9: 1). pale brown labrum, simple Iabella lacking short Wings 1. 8-1.9 mm long. Costal index 0.54-- pale spines below. 0.57. Costal ratios 1.01 - 1.18 : I, vein 3 being un­ Thorax brown. Notopleuron with three bristles. forked. Costal cilia 0.07- 0.08 mm long. Vein A pair of humeral, intra-alar, post-alar and poste­ Sc runs to R 1, but is somewhat pale and subcostal rior dorsa-central bristles on scutum. Scutellum cell is very narrow. A small hair at base of vein 3. with an anteri or pair of short, fine hairs and a All veins brownish, including vein 7. Axillary posterior pair of long bristles. Mesopleuron with ridge with 3-5 bristles. Membrane distinctly 10- 14 small hairs and a posterior pair of strong brownish grey tinged. Haltere largely pale greyish bristles, the lower of which is longer and stronger. brown. Abdomen with dark brown tergites and brownish venter. The latter with, mostly fine, Female: Frons and its chaetotaxy as in male. In hairs on segments 3-6 below, most obviously on one specimen there is a set of fo ur small, supernu­ segments 5 and 6, and a few on the fl anks above. merary, supra-antenna! bristles above the normal Tergite hairs short and fine, except for stronger supra-antennals. Antennae as in male. Palps a postero-laterals. Hypopygium as Fig. 1, being little more slender than in male, and with bristles brown with a dirty yellow anal tube. a little longer. Thorax as in male. 28 Disney: A new species of Megaselia, Phoridae • ENTOMOL. FENNICA Vol. I

Abdomen with colouring and hairing as in morphic features include the unforked vein 3, the male apart from development of bristles on venter modified ovipositor segments, and probably the at rear of segments 4-6. In particular segment 6 elongated anal tube of the male. with a posterior row of long bristles encircling the The sexual dimorphism in the number of fore segment (Figs 2 and 4). Ovipositor (Fig. 2) well tarsal segments with a postero-dorsal hair pali ­ developed. Dufour's crop mechanism weakly sade is unusual. The plesiomorphic state is proba­ sclerotised. bl y the complete seri es of the male. Legs as male, except fore tarsus with postero­ The supernumerary supra-antenna! bristles in dorsal hair palisades on fi rst four segments onl y. some specimens casts serious doubt on the taxo­ Wings 1. 8- 1. 9 mm long. Costal index 0.56- nomic weight given to this character at the generic 0.58. Costal ratios 0.94 : I. Costal ci lia 0.07 mm level. Indeed the taxonomic weight given to the long. Otherwise as male. number and inclination of the bristles at the front of the frons is undoubtedly misplaced. Mutants in A number of features of M. torautensis invite both, on one side at least, are frequent in Megase­ comment. lia. The cases reported above reinforce the grow- The presence of two hair palisades on the mid tibia (Fig. 3) is in contrast to the single dorsal hair palisade of most known M egaselia species. Double hair palisades on the hind tibia have been sug­ Table 1. The numbers of longitudinal hair palisades on the tibiae. gested as a ground-plan feature of the Phoridae (Disney 1988). In the undoubtedly monophyletic Hind Mid Front genus Woodiphora Schmitz there is a transforma­ tion seri es from two, to one, to no hair palisades on PLATYPEZIDAE both the mid and hind tibiae (Disney 1989). The Plesioclythia argyrogyna (De Meijere) 2 2 Platypezid P/esioc/ythia argyrogyna (De Meijere) not only has double hair palisades on both its hind SCIADOCERIDAE and mid tibiae, but also has a single palisade on its Sciadocera rufomaculata White 0 0 0 front tibia. The simplest interpretation, on the PHORIDAE principle of out-group comparison, is that the Aenigmatias lubbocki (Verrall) 4- 5 3 1 ground plan of the Phoridae included double hair Aenigmatistes foveolatus Schmitz 1 1 0 Cataclinusa pachycondylae (Brues) 2 2 0 palisades on its mid tibia. Table 1 summarises the Diplonevra abbreviata (von Roser) 3 2 0 recorded pattern s. The most parsimonious inter­ D. bifasciata (Walker) 3 2 0 pretation of the double hair palisade on the mid D. concinna (Meigen) 2 2 0 tibia of M. torautensis is that this is a plesiomorphic, D. florea (Fabricius) 2 2 0 ground-plan, feature that has been retained. Fur­ D. funebris (Meigen) 2 2 0 thermore the evidence suggests that the reduction D. glabra Schmitz 2 2 0 D. watsoni Disney 2 2 0 from 2 to 1 palisade on both the mid and hind tibiae Dohrniphora cornuta (Bigot) 1 0 has taken place independently in Woodiphora and Epicnemis flavidula (Brues) 1 0 Megaselia. Megaselia spp. (most species) 1 1 0 Another feature interpreted as being plesiomor­ M. bruesi Disney 1 2 0 phic, in M. torautensis, is the distinct 'coll ar' be­ M. torautensis sp. n. 1 2 0 Multinevra macropygidia Disney 5-7 3 1 tween the epandrium and the anal tube. According Myopiomyia harmani Disney 1 0 to the most parsimonious interpretation of the Palpiclavina tonkinensis Silvestri 0 0 homologies of the principal sclerites of the hy­ Plectanocnema nudipes (Becker) 4-5 2-3 0 popygium this collar is the fused tergite and ster­ Woodiphora bilineata Borg meier 2 2 0 nite of segment 10 (Di sney 1986b, 1988, 1990). W. biroi (Brues) 1 0 0 W. papuana Disney 2 Other features considered to be plesiomorphic are 0 0 W. parvula Schmitz 2 1 0 the presence of3 not 2 notopleural bristl es, vein Sc W. retroversa (Wood) 0 0 0 terminating in R 1, costal index exceeding 0.5, the W. salomonis Beyer 1 1 0 short costal cilia, and vein 7 being distinct. Apo- ENTOMOL. FENNICA Vol. I • Disney: A new species of Megaselia, Phoridae 29

ing evidence that these characters should be any sustainable justification. Elsewhere (Disney downgraded in terms of their taxonomic weight. 1989) it was concluded that a more satisfactory Certainly the validity of some genera separated basis for recognition of tribes within the Metopin­ off from M egaselia must be called into question. inae is required. Certainly the inclination of bristles Four such genera, from Africa, are re-assessed at the front of the frons is not adequate grounds for below. erecting a tribe. Information on variation in the placement and inclination of bristles in general would suggest that single gene mutations can Megaselia bisetigera (Beyer) comb. n. bring about striking differences. Likewise super­ numerary bristles are frequent, as evidenced by Tarsophoromyia bisetigera Beyer, 1965: 198. the supernumerary 'supra-antenna!' bristles re­ The genus Tarsophoromyia was erected for a ported in two specimens of M. torautensis above. species only known in the female sex. It has an These must be due to simple gene mutations. ovipositor modified in a manner associated with In view of the above considerations the con­ parasitoid habits, combined with swollen fore­ cept of the genus Epimegaselia cannot be sus­ tarsal segments. Both characters are widespread tained. I thus transfer E. cauda/is to the genus in the genus Megaselia, the only unusual feature Megaselia and synonymise Epimegaselia with being that swollen fore-tarsal segments are nor­ the latter. mally restricted to the male sex. If T. bisetigera is Megaselia cauda/is will run to couplet 5 on to be placed in a separate genus then it must be page 56 in Beyer's (1965) keys. inferred that there is a stronger case for transfer­ ring several hundred species from Megaselia to new genera. I conclude that Beyer failed to make Megaselia congrex (Beyer) comb. n. out a case for placing T. bisetigera in a new genus, Metaplastophora congrex Beyer, 1965: 188 even in typological terms. I herewith transfer it to Megaselia, and synonymise Tarsophoromyia with This species was placed in Metaplastophora the latter genus. along with the type species M. rotundicauda Beyer. Megaselia bisetigera runs, allowing for ec­ The validity of the genus is called into question centricities of construction that altemates cou­ under the latter (see below). plets based on the female or the male sex only, to If there were a case for placing this species in couplet 30 on page 54 of Beyer's ( 1965) keys. a separate genus to Megaselia then it would be difficult to justify placing it in the same genus as M. rotundicauda. Although Beyer ( 1965) stressed Megaselia cauda/is (Beyer) comb. n. that both have a compressed ovipositor he gave less taxonomic weight to the obvious differences Epimegaselia cauda/is Beyer, 1959a:74. in the fom1s of the ovipositors. For example cerci The genus Epimegaselia was distinguished are present in M. congrex but absent in M. rotun­ from M egaselia by the presence of an extra pair of dicauda. bristles on the front margin of the frons, or at least In the keys to Afrotropical Megaselia (Beyer by an extra pair of reclinate bristles. Beyer ( 1959a) 1965) this species will run to couplet 4 an page reported only a single pair of proclinate supra­ 48. antenna! bristles. The 'extra' pair of bristles coul d be a pair of supra-antennals that happen to be reclinate rather than porrect. Megaseliafurvicolor (Beyer) comb. n. The porrect, as opposed to proclinate, inclina­ Quasipseudacteon furvicolnr Beyer, 1959b:387. tion of the supra-antenna! bristles is the basis for the recognition of the tribe Beckerinini (Schmitz The genus Quasipseudacteon was distin­ 1956). In spite of this genera without supra-anten­ guished from Megaselia by a modified ovipositor, nals were subsequently added to the tribe, without a supemumerary pair of reclinate bristles near the 30 Disney: A new species of Megaselia , Phoridae • ENTOMOL. FENNICA Vol. 1

front margin of the frons and a lack of proclinate 1978, 1986a). Beyer's emphasis on the compres­ supra-antenna! bristles (Beyer 1959b). sion of the ovipositor tends to obscure the consid­ The evidenc~ suggests that the terminalia of erable difference in the structure of the ovipositor the female abdomen have been independently in these two species. For example there are no modified in relation to the independent evolution cerci in M. rotundicauda, but cerci are present in of parasitoid habits in Phoridae in many clades, M. congrex. and in the genus Megaselia in particular (e.g. Once again we are confronted with an attempt Disney 1978, 1986a). Thus "modified ovipositor" by Beyer to select part of the diverse radiation of is not a single character, and certainly not a syna­ ovipositor forms in Megaselia, on the basis of a pomorphic character. Until the different types of single ill-defined characteristic (in this case the modification have been elucidated, and homolo­ compression of the ovipositor), and to erect a new gous transformation series identified, these char­ genus. This might be justified if detailed studies acters cannot be used in classification. had been made of the adaptive radiation in the The highlighting of the frontal bristle charac­ morphological modifications in the Metopininae, ters by Beyer rests on the assumption that recti­ and monophyletic clades identified. Beyer, how­ nate or ponect bristles cannot be supra-antennals ever, proposed a number of new genera on the in the Metopinini, as opposed to the Beckerinini. basis of poor descriptions and uncritical evalu­ This, however, is a circular argument. If the incli­ ation of the structures used as a basis for erecting nation of these bristles is given taxonomic weight the proposed genera. He then adds to the confu­ at the species level only then the frontal chaeto­ sion, in the case of M etaplastophora, by declaring taxy of Quasipseudacteon is best interpreted as that the males cannot be distinguished from Meg­ being ofthe normal M egaselia type, except unlike aselia. I have no hesitation in synonym ising Meta­ most (but not all) Megaselia species there is only plastophora with Megaselia. one pair of supra-antennals (which happen to be In the keys to Afrotropical Megaselia (Beyer reclinate). As with Epimegaselia (see under M. 1965) this species will run to couplet 4 on page 48. cauda/is above) we must discount the supposed difference from Megaselia in the frontal chaeto­ taxy. Megaselia subulicauda Schmitz Until at least one unambiguous apomorphic character is demonstrated, on which to base a Megasela subulicauda Schmitz, 1929:39 generic distinction, the genus Quasipseudacteon Hemiplastophora subulicauda (Schmitz) Beyer, 1965: 186. cannot be sustained. I thus transfer Q. furvicolor New synonym. to Megaselia and synonymise Quasipseudacteon Beyer (1965) transferred Megaselia subuli­ with Megaselia. cauda to a new genus Hemiplastophora on the Megaselia furvicolor will run to couplet 3, grounds that it has a somewhat elongated tubular under "4 scutellaren", on page 48 of Beyer's ovipositor. It seems, from his choice of name, that ( 1965) keys. he saw this as intermediate between Megaselia and Plastophora. With the synonymising of Plas­ tophora with M egaselia (Disney 1978, 1986a) the Megaselia rotundicauda (Beyer) comb. n. genus Hemiplastophora cannot be sustained. I therefore return H. subulicauda to the genus Meg­ Metaplastophora rotundicauda Beyer, 1965:1 86. aselia, and synonymise Hemiplastophora with Beyer placed this species and M. congrex (see Megaselia. above) in a new genus Metaplastophora on the Megaselia subulicauda will run to couplet 20 basis of the ovipositors being somewhat com­ on page 50 of Beyer's ( 1965) keys. pressed. In his key to genera (Beyer 1965) he was Acknowledgements. This paper is based on specimens col­ unable to separate the male of this species from lected by the author and Dr. A. H. Kirk-Spriggs while we Megaselia and Plastophora. The latter genus has were partipants on Project Wallace, sponsored by the Royal since been synonymised with Megaselia (Disney Entomological Society of London and the Indonesian Insti- ENTOMOL. FENNICA Vol. 1 • Disney: A new species of Megaselia, Phoridae 31 tute of Sciences (Results of Project Wallace No. 79). My Britain. - Zeitschr. Ang . Zoo!. 70:225-234. participation in Project Wallace was made possible by 1983b: A synopsis of the taxonomist's tasks, with par­ grants from the Field Studies Council Study Leave Fund ticular allention to phylogenetic cladism. - Fld. Stud. (fi nanced by an anonymous trust), the British Ecological 5(5):841-865. Society, the Royal Entomological Society and the Percy 1986a: A new genus and three new species of Phoridae Sladen Memorial Fund (administered by the Linnean Soci­ (Diptera) parasitizing ants (Hymenoptera) in Sulawesi. ety). - J. Nat. Hi st. 20:777-787. 1986b: Two remarkable new species of scullle-fly (Diptera: Phoridae) th at parasiti ze tem1ites (Isoptera) in Sulawesi.- Syst. Entomol. II :4 13-422. References 1988: An interesting new species of Megaselia from Sul awesi, the ground plan of the Phoridae (Diptera) and Beyer, E. M. 1959a: Neue Phoridengattungen und -Arten phylogenetic implications for the Cyclorrhapha. - a us Angola (Phoridae, Diptera). -Camp. Diam. Angola Syst. Entomol. 13:433-44 1. 45 :55-76. 1989: A key to Australasian and Oriental Woodiphora 1959b: Diptera (Cyclorrhapha) Phoridae. - In: (Diptera: Phoridae), affinities of the genus and descrip­ Hanstrom, B., Brinck, P. & Rudebeck, G. (eds.), South ti ons of new species.-J. Nat. Hi st. 23: 11 37- 11 75. African Animal Life 6:376-389. 1990: Phylogenetic implications of some features of 1965: Phoridae (Diptera Brachycera).- Exploration Sulawesi scullle flies (Diptera: Phoridae). - ln: Knight, du Pare National Albert, Mi ssion G. F. De Witte W. J. & Holloway, J. D. (eds.), Insects and the rain ( 1933-1935) 99: 1-211 . forest of South East Asia- A special Project Wallace Disney, R. H. L. 1978: A new species of afrotropical symposium. R. Entomol. Soc. London (i n press). Megaselia (Diptera : Phoridae), with a re-evaluation of Schmitz, H. 1929: Zur Kenntnis eini ger von Dr. Jos. Be­ the genus Plastophora. - Zeitschr. Ang. Zoo!. quaert gesammelter afrikani scher Phoriden. - Rev. 65:313-319. Zoo!. Bot. Afr. 18:37-43. 198 1: A new species of Megaselia from Nepenthes in 1956: Phoridae. -In: Lindner, E. (ed.), Die Fliegen der Hong Kong, with re-evaluation of genus Endonepen­ palaearktischen Region 4 (33) (Lief. 187):369-4 16. thia (Diptera: Phoridae). -Oriental Insects 15:201-206. 1983a: A useful new character in the giant genus Meg­ aselia (Diptera: Phoridae), with two new species from Received 17.JI.I989