Australian Entomologist, 2016, 43 (1): 17-30 17 A REVIEW AND KEYS TO GENERA AND SOME SPECIES OF THE FRUIT FLY TRIBES PHYTALMIINI, PHASCINI AND EPACROCERINI (DIPTERA: TEPHRITIDAE: PHYTALMIINAE) DAVID L. HANCOCK 8/3 McPherson Close, Edge Hill, Cairns, Qld 4870 Abstract The tribes Phytalmiini, Phascini and Epacrocerini are considered to form a related grouping that includes 18 primarily New Guinea genera: Phytalmiini with Diplochorda Osten Sacken, Ortaloptera Edwards, Phytalmia Gerstaecker and Sessilina McAlpine & Schneider; Phascini with Diarrhegmoides Malloch, Epinettyra Permkam & Hancock, Gressittidium Hardy, Othniocera Hardy, Paraphasca Hardy, Phasca Hering, Stigmatomyia Hardy and Xenosophira Hardy; and Epacrocerini with Epacrocerus Hardy, Proepacrocerus Hardy, Sophiropsis Hardy, Tanaodema Hardy, Tanymetopus Hardy and Udamolobium Hardy. Three genera occur in Australia. A key to genera and keys to the species of Ortaloptera and Diplochorda are included. The type locality of Diplochorda myrmex Osten Sacken, 1881 is considered to be in Papua New Guinea and 8D. trineata9 de Meijere, 1913 is regarded as a nomen nudum, with the species correctly named as D. trilineata de Meijere, 1915. Known host plants are newly fallen logs. Introduction Although sometimes treated in the past as a separate family or subfamily (e.g. Malloch 1939), there is general agreement among recent authors (e.g. Korneyev 1999, Hancock and Drew 2003) that the Phytalmia group represents a tribe (Phytalmiini) closely related to the Acanthonevrini and contains the genera Diplochorda Osten Sacken, Phytalmia Gerstaecker, Sessilina McAlpine & Schneider and Ortaloptera Edwards, which all have mid tibiae with 1 long, 1 medium, 1 short and several very short and thin apical black spines. The Phytalmiini appear to belong in a grouping within subfamily Phytalmiinae that also includes the tribes Phascini and Epacrocerini (sensu Korneyev 1999), which also have 1 long and several shorter midtibial spines. With the possible exception of Paraphasca biangulata (de Meijere) (see Hancock 201 1a), this group is restricted to the mainland of New Guinea and surrounding islands such as Salawati, Waigou, Biak, Japen and New Britain, plus northeastern Queensland, Australia, where three species occur. Host records are available for three species in the Phytalmia complex: Phytalmia alcicornis (Saunders) and P. mouldsi McAlpine & Schneider were reared from beneath the bark of recently fallen Dysoxylum gaudichaudianum (Meliaceae) trees (Hardy 1986, Dodson and Daniels 1988, Dodson 1989, 1999, Permkam and Hancock 1995); Phytalmia cervicornis Gerstaecker was attracted to D. gaudichaudianum (Dodson 1999) and reared from Xanthophyllum sp. (Xanthophyllaceae) (Dodson and Daniels 1988). The biology of P. mouldsi and other Phytalmia species has been studied in detail by Moulds (1977) and Dodson (1989, 1999), while an additional species was described by Schneider (1993) and their phylogenetic relationships were discussed by Schutze et al. (2007). 18 Australian Entomologist, 2016, 43 (1) No host or biological information is available for either Phascini or Epacrocerini but, as with Phytalmiini, they appear to be rainforest dwellers, with several species of Ortaloptera, Phytalmia, Othniocera Hardy, Paraphasca Hardy, Phasca Hering, Stigmatomyia Hardy, Epacrocerus Hardy and Sophiropsis Hardy collected in logging areas in Papua New Guinea (Hardy 1982, 1986, 1988, Schneider 1993). The following generic key is based on the works of Hardy (1982, 1986) and McAlpine and Schneider (1978). Terminology follows White et al. (1999). Key to genera of Phytalmiini, Phascini and Epacrocerini 1 Wing with R-M crossvein placed above basal third of cell dm, well before midline; wing pattern in apical half with a large oval brown area separated from a narrow costal band by an elongate, C-shaped hyaline band [2 species, keyed below] ............::sscccceeeeeeeeeeeeeeees Ortaloptera Edwards, 1915 4 Wing with R-M crossvein placed at or beyond middle of cell dm; wing pattern not as above 5.. raaccecenssy zssaanaa ceccoaeaa te aaamuiers) J iesemmannsedeunmnananerneceeesd> ? 2 2 Wing distinctly elongate; pterostigma elongate, if not much longer than cell c then very narrow and meeting costa at a sharply acute angle; cell bcu apically acute but at most weakly produced; wing pattern not dimidiate; scutellum with only 1 pair of strong, apical setae; head often with genal processes in males but not distinctly elongate; metathoracic postcoxal bridge strongly sclerotised (Tribe Phytalmiini) .......00... 3 4 Wing usually not distinctly elongate and pterostigma usually not narrow and elongate, if so then pattern dimidiate, brown anteriorly and reticulate posteriorly; cell bcu often distinctly produced at apex; scutellum usually with 2-3 pairs of strong marginal setae, if only 1 then head distinctly elongate and wing pattern dimidiate; metathoracic postcoxal bridge MEDIDOS eeina eee tata areias bihderehibhtitel cearhieSesisnsdesbbberecckbhltitetes 5 3 Wing with veins Ri, R23 and R4+5 closely approximate for much of their length and vein M strongly downcurved before R-M crossvein; pterostigma much longer than cell c; males with costa strongly arched in its outer half and with or without short, broad genal processes [9 species, keyed below; Nesadrama Perkins, 1939 is a synonym (McAlpine and Schneider 1978)] .............ceeeeeeeeeeeeeeeeeeees Diplochorda Osten Sacken, 1881 4 Wing with veins R;, Rz, and Ry,5 not closely approximate for much of their length and vein M not downcurved before R-M crossvein; pterostigma not much longer than cell c; genal processes variable .......... 4 4 Abdomen not petiolate, tergite 1+2 not constricted basally, anterior notopleural seta well developed, as long as posterior one; pterostigma about as long as cell c; males with or without short genal processes [3 species, keyed by McAlpine and Schneider (1978)] EATE EEE E STEET, Sessilina McAlpine & Schneider, 1978 Australian Entomologist, 2016, 43 (1) 19 4 Abdomen distinctly petiolate, tergite 1+2 constricted basally; anterior notopleural seta much weaker than posterior one or absent; pterostigma much shorter than cell c; males with long, antler-like genal processes [7 species, six keyed by McAlpine and Schneider (1978) plus one (P. robertsi Schneider) added by Schneider (1993); Elaphomyia Saunders, 1861 and Archiphytalmia Edwards, 1936 are synonyms (McAlpine and Schneider 1978); male genal processes illustrated by Dodson (1999) and Schutze et al. (2007)] ssesssisssooctttitresseesesssaana Phytalmia Gerstaecker, 1860 5 Wing cell bcu apically truncate or weakly acute but not produced into a distinct apical lobe; usually 2 pairs of scutellar setae, rarely 1 or 3; 1 pair each of long frontal and orbital setae, sometimes a weak upper orbital seta also present and sometimes all absent (Tribe Epacrocerini) .................04. 6 4 Wing cell bcu produced into a distinct apical lobe; usually 3 pairs of scutellar setae, rarely 2; usually 2-3 pairs of frontal setae, rarely 1; usually 2 pairs of orbital setae, if 1 then short and placed well behind middle of frons (Tribe Phascini) ............cccccccccccccccecccseesssscsecccccssseuseeeseeeseceeseteseeeaeees 11 6 Second antennal segment not lobate; face gently concave; wing pattern with brown apical area broadly separated from brown transverse band from costa over R-M crossvein into cell dm ...sssssssssooneesssesesssesssssrsssserree 7 4 Second antennal segment lobate on inner margin and extending beyond base of arista; face vertical or convex; wing pattern not as above ........... 8 7 Two pairs of scutellar setae; cell bcu apically blunt [1 species, illustrated by Hardy (1988)] cee ceeeeesseeestnneeeeeeees Proepacrocerus Hardy, 1988 4 Three pairs of scutellar setae; cell bcu apically acute [2 species, illustrated by Hardy C193 6). .cc Reslaataaatssanaen dadenatmeentets Sophiropsis Hardy, 1986 8 Scutellum with only 1 pair of distinct, apical setae; head narrow and elongate in lateral view; wing narrow and elongate, the pattern dimidiate, brown anteriorly and largely reticulate posteriorly [1 species, illustrated by Hardy (1987)] «eoneiacdececassadddenaaaereeseeaeoeeseaennees Tanaodema Hardy, 1987 4 Scutellum with 2 pairs of distinct setae; head broad in lateral view, often globose; wing relatively narrow but not distinctly elongate, the pattern not AS ADOVE: TETESI EEEE E AET anal rpc ta TEE EEIE 9 9 Head higher than long; face almost vertical; upper occiput narrow; thorax normal in shape, not elongate and slender [4 species, keyed by Hardy OESE2A N ES Epacrocerus Hardy, 1982 4 Head longer than high; face strongly receding; occiput inflated; thorax SION SAL and Slendër sssrini iiine naes | aaide iaeiae 10 10 Wing hyaline except a tinge of brown near apex; pterostigma less than half length of cell c; 1 long and 1 short costal spines above apex of vein Sc [1 species, illustrated by Hardy (1982)] ...... Tanymetopus Hardy, 1982 20 Australian Entomologist, 2016, 43 (1) 4 Wing with a complex pattern of brown and hyaline markings; pterostigma large, longer than cell c; 2 short costal spines above apex of vein Sc [1 species, illustrated by Hardy (1982)] 0.0.0.0... Udamolobium Hardy, 1982 11 Wing with R-M crossvein placed below middle or basal half of pterostigma, well before its apex ....... cee ccssseeceeeeeeeeeeeeeeeeeneeeeeeeetaeeeeees 12 4 Wing with R-M crossvein placed below or well beyond apex of PEL ORISAIIA seieren reann at re retete rran n errr E rreren tern ces 14 12 Wing with a distinct costal spine at apex of vein Sc; veins R3 and Rais strongly arcuate; intrapostalar setae distinct [1 species, illustrated by Hardy (1986)] .eeeseessseesseesessssssssrrereesssssssssssse Stigmatomyia Hardy, 1986 4 Wing without a distinct costal spine at apex of vein Sc; veins Rz, and R4 not strongly arcuate; intrapostalar setae absent ............ccsseeeeeeeeeeeees 13 13 Wing with a hyaline transverse band from costa to just below vein M in cell dm and cell dm mostly brown; 3 pairs of scutellar setae [1 species, illustrated by Hardy (1986)] -eeen Gressittidium Hardy, 1986 4 Wing without a hyaline transverse band from costa to just below vein M and cell dm mostly subhyaline to pale fulvous; 2 pairs of scutellar setae [1 species, illustrated by Permkam and Hancock (1995) and Hancock (2011b: 9, not  as stated)] ......... Epinettyra Permkam & Hancock, 1995 14 Wing with a broad hyaline transverse band from costa in cell r1 at apex of pterostigma across base of cell dm to posterior wing margin [2 species, keyed by Hancock (201 1a)] ......eeeeeeeeeeeeeeeeeeeees Paraphasca Hardy, 1986 4 Wing without a complete hyaline transverse band from cell r1 to posterior margin, the hyaline indentation triangular and not crossing vein M ..... 15 15 Two pairs of scutellar setae [2 species, illustrated by Hardy (1980)] ETE AA TA EIE EEA E A ETE Xenosophira Hardy, 1980 4 Three pairs of scutellar setae 0... eee eeceeeeeenneceeecesnseeeeeeeessneeeeeeeeaeees 16 16 Third antennal segment large and broad, with antennae at least half height of eye and usually as long as face; if antennae only 2/3 length of face then only 1 pair of frontal setae; middle pair of scutellar setae weak [3 species, keyed by Hardy (1986)] o.... eee eeseeeeeseeeeeeeees Othniocera Hardy, 1986 4 Third antennal segment not large and broad, with antennae less than half height of eye and distinctly shorter than face; 2-3 pairs of frontal setae; middle pair of scutellar setae distinct 2.0... eee eeeeeesneeeeeeneeeeeeeeeeeeeeneeees 17 17 Intrapostalar setae absent; arista with short hairs on dorsal surface only [1 species, illustrated by Hardy (1986)] ....... Diarrhegmoides Malloch, 1939 4 Intrapostalar setae present; arista long-plumose on both dorsal and ventral surfaces [6 species, keyed by Hardy (1986)] ............ Phasca Hering, 1953 Australian Entomologist, 2016, 43 (1) 21 Figs 1-3. Ortaloptera callistomyia females from Upper Manki logging area near Bulolo, Papua New Guinea: (1) lateral view; (2) dorsal view; (3) mid tibial spines. Photos by Barbara Baehr (Queensland Museum). 22 Australian Entomologist, 2016, 43 (1) Key to Ortaloptera species This genus is known only from mainland New Guinea and the following material was examined: O. cleitamina Edwards 4 holotype 2°, Dutch New Guinea, Mimika River, vii.1910, A.F.R. Wollaston, 1911-229 (in Natural History Museum, London); O. callistomyia Hering 4 1 £, 2 299, Papua New Guinea: Upper Manki logging area, near Bulolo, 50009, 15.xii.1972 (Q), 29 .xii.1972 (Q), 9.iii.1973 (4), F.R. Wylie and P. Shanahan, sticky trap (in Australian Museum, Sydney). 1 Thorax and abdomen uniformly black without pale longitudinal vittae; legs blackish brown [Indonesia (Mimika River, Papua Province) | eA RATE PAA maa AERO TREE TT tfO t ETRE na TE O. cleitamina Edwards, 1915 4 Thorax and abdomen reddish brown to black with distinct pale or yellow markings; legs mostly yellowish with a tinge of brown [Indonesia (Papua Province) and northern Papua New Guinea; illustrated by Hardy (1988) and in Figs TH]. eesasssonsverssssnndanpnudereeecetinesyes O. callistomyia Hering, 1941 Key to Diplochorda species This key is derived largely from Malloch (1939), with additional information provided by Osten Sacken (1881), Perkins (1939), Hardy (1974) and Permkam and Hancock (1995). Diplochorda is a mainland New Guinea genus, with one species extending as far west as Salawati Island and one as far south as Cape York Peninsula, NE Australia. In males the costa is strongly arched in its outer half and the genae in at least three species are expanded into short, blunt processes. The type species is Dacus turgidus Walker, 1865 (a synonym of Dacus concisus Walker, 1861). 1 Wing with apex of costal band extending broadly across R-M crossvein and apex of cell dm and into cell m almost to vein Cu,; hyaline discal area hatchet-shaped, subquadrate in centre of wing, extending along its length anteriorly across vein Ry,5 to vein R»,3; male unknown [southern Papua New Guinea (Western and Central Provinces: 1 9 examined, Central Province, 5 km NW Brown River bridge, forest, 29.xii.1985, J.W. Ismay, in Natural History Museum, London); Nesadrama petiolata Hardy, 1974 (described from Mindanao, Philippines but presumably mislabelled) is a synonym (McAlpine and Schneider 1978); illustrated as 8N. petiolata9 by Hardy (1974)] ertiecreherrtreneseaneusennenvereiukneld D. myrmex Osten Sacken, 1881 [The type female of D. myrmex was collected by L.M. D9Albertis at Katau (= Binaturi River, 09°08'29"S 142°57'10"E, ca 30 km west of Daru in Western Province, Papua New Guinea) during his 1876-77 expedition to the Fly River, not in Indonesian Papua as indicated by Norrbom et al. (1999). D9 Albertis (1880) made no mention of Katau during his travels in Indonesia9s West Papua Province in 1872-73 but recorded it (as 8Kataw9 ) during his 1876-77 expedition. Note that 8D. myrmex9 of Malloch (1939) is a misidentification of D. trilineata de Meijere.] Australian Entomologist, 2016, 43 (1) 23 4 Wing with apex of costal band not extending broadly into cell m; hyaline discal area elongate and not extending anteriorly across vein R445 to vein R543 Pererr eee ere eer arer eee rece rere EERE ETTI rere Tere rer ee Tee reyerer ee eT eee re eeS erer Tee Tee eee Tere ece e eee eee 2 2 Thorax with scutum anterior to suture largely or entirely black, without distinct longitudinal yellow vittae; male with broad genal processes ...... 3 4 Thorax with scutum anterior to suture yellow with 3 longitudinal black vittae; male unknown or without broad genal processes 1.0... ceeeeeeeeeee 5 3 Wing with costal band crossing R-M crossvein and filling most of cell r4,5 [northern Papua New Guinea; wing and head illustrated by Malloch CFDS 95] T a aaae AA AE aE aa ANa EAN D. aneura Malloch, 1939 4 Wing with costal band not crossing R-M crossvein and filling none or Gnily-part, faintly otee Tys araaarN aNR nes 4 4 Scutum posterior to suture largely yellow, with a medial longitudinal black vitta; scutellum yellow [Papua New Guinea and Australia (Iron Range, northern Qld); illustrated by Permkam and Hancock (1995)] beau E EE EE D. australis Permkam & Hancock, 1995 4 Scutum posterior to suture almost entirely black; scutellum black [eastern Indonesia (West Papua Province); illustrated by Saunders (1861: male GTi) ITEE E E T D. brevicornis (Saunders, 1861) 5 Legs yellow with faint vestiges of a brownish ring on femora; abdomen mostly yellow, with yellow band on tergite 1+2 not isolated and with a pair of large black lateral patches from tergites 2 to 4; face yellow; male unknown [eastern Indonesia (Arfak Mts, West Papua Province)] NECETE ELIO DIAE AT A LOEN ec dake D. ophion Osten Sacken, 1881 4 Legs with mid and hind femora broadly brown to black medially; abdomen mostly brown to black, often paler posteriorly and with yellow band on tergite 1+2 isolated; face with or without dark spots or band; male with genal processes represented by low ridges ...........ccseceeeeeeeeeeeees 6 6 Wing cell cu, hyaline; face yellow; abdomen with yellow band on tergite 1+2 large, separated from hind margin by less than half its own length [Papua New Guinea (Central and Eastern Highlands Provinces above 200m] ensAe aseae RA D. unistriata Malloch, 1939 4 Wing cell cu; brown at least anteriorly; face with a pair of black spots or band near epistome; abdomen with yellow band on tergite 1+2 small, separated from hind margin by about its own length or more ................. 7 7 Wing with costal band crossing R-M crossvein and filling all or most of cell r445 [eastern Indonesia (northern Papua Province) and northern Papua New Guinea; 8D. myrmex9 of Malloch (1939) is a misidentification; wings illustrated by Malloch (1939)] wee D. trilineata de Meijere, 1915 24 Australian Entomologist, 2016, 43 (1) [The name 8Diplochorda trineata9 appeared in a list of taxa reported at a meeting by de Meijere (1913) two years before its formal description (de Meijere 1915) as D. trilineata. De Meijere9s original (1913) report stated: 8Diplochorda trineata, eene nieuwe soort, waarbij de 63 de eigenaardige kopaanhangels missen, welke eenige der overige soorten kenmerken.9 [Diplochorda trineata, a new species, where the @ lack the peculiar genal processes, which feature in some of the other species.] However, use of the word 8eenige9 [some] indicates that this absence also applies to other species [e.g. D. concisa (Walker)] and the above does not constitute a description or definition under the strictly defined terms of the 8Code9 (ICZN 1999) but merely records 8a new species9 that lacks a character also lacking in other known species. Furthermore, de Meijere (1915) did not mention 8trineata9 or refer to his previous note when describing D. trilineata, clearly neither accepting nor having intended it as a valid name. Hence, the name D. trineata is regarded here as a nomen nudum, having appeared without a valid description, definition or indication, with D. trilineata de Meijere, 1915, thus regarded as the valid name.] Wing with costal band not crossing R-M crossvein and filling none or only part, faintly, of cell r445 ..eesessesssssesesssseesessssreeessssreeessssereessssereessssrrresss 8 Wing with costal band not crossing vein R445 into cell r4,5 or at most very faintly at its apex [northern Papua New Guinea; wing illustrated by Malloch (1939)] 2 ccsazalistssemmesrsit dtetaaeauaneisess D. minor Malloch, 1939 Wing with costal band crossing vein R445 into apical half to two-thirds of cell r4,5 in its anterior half and extending weakly as a narrow patch almost to apex of vein M [eastern Indonesia (West Papua Province) and southern Papua New Guinea; Dacus turgidus Walker, 1865 and Nesadrama longistigma Perkins, 1939 are regarded as synonyms; male illustrated by Saunders (1861, as 8Elaphomyia brevicornis female9, a misidentification) and female wing illustrated by Perkins (1939, as 8N. longistigma9) ] Lubes) ppp a A re A A erh D. concisa (Walker, 1861) [Diplochorda turgida (Walker) (= N. longistigma Perkins: synonymised by McAlpine and Schneider (1978) after examination of their types) was listed as a separate species by Norrbom et al. (1999) but it is not known on what basis this was done; it had been treated previously as a synonym of D. concisa by both Osten Sacken (1881) and Hardy (1959) after examination of their types and, in the absence of further information, that arrangement is followed here. Walker9s (1861, 1865) descriptions are essentially identical, with that of the third antennal segment in D. concisa (8about one-third of the length of the 2nd9) clearly a lapsus; it is actually about 2.5 times the length of the second (D. Whitmore pers. comm.). This is a widespread species, known from Salawati Island, Ramoi (near Sorong) and Manokwari in West Papua, plus Kokoda and Mt Lamington in Papua New Guinea. ] Australian Entomologist, 2016, 43 (1) 25 Discussion Tribe Phytalmiini The composition of this tribe 4 genera Phytalmia, Diplochorda, Sessilina and Ortaloptera 4 was suggested by McAlpine and Schneider (1978) and accepted by Korneyev (1999), who noted that the female aculeus was only half the length of the oviscape (apomorphy). The presence of male genal processes in at least some species in all four genera, the apically acute but non-lobate wing cell bcu, the narrow and apically sharply acute pterostigma and the lack of a costal spine above the apex of vein Sc also support this association. At least two of the midtibial apical spines are well developed. Ortaloptera differs from the other genera in retaining two pairs of scutellar setae and a membranous metathoracic postcoxal bridge (plesiomorphies) and is presumably the most primitive of the tribe. In Sessilina, Diplochorda and Phytalmia there is only one pair of scutellar setae and the metathoracic postcoxal bridge is broadly sclerotised (apomorphies). In Diplochorda and Phytalmia the abdomen is petiolate and the anterior notopleural seta is weak or absent (apomorphies); the latter is well developed in Ortaloptera and Sessilina and the abdomen is not petiolate (plesiomorphies). Phytalmia, with its well developed genal processes and Diplochorda, with its strongly arched costa in males (both apomorphies), are the most speciose of the genera. In a phylogenetic study by Schutze et al. (2007), morphological and morphological + mitochondrial DNA data supported the monophyly of Phytalmia, whereas the mitochondrial data alone (COII, 16S and combined COII+16S) resulted in paraphyly with respect to Sessilina and/or Diplochorda, with each resulting cladogram different from the others. This casts considerable doubt on the reliability of molecular evidence, in isolation, as an indicator of phylogenetic relationships. All four genera occur in New Guinea, with single species of Phytalmia (P. mouldsi McAlpine & Schneider) and Diplochorda (D. australis Permkam & Hancock) occurring at Iron Range in northeastern Queensland. Tribe Phascini This tribe was regarded as distinct and defined by Korneyev (1994, 1999) to include five genera 4 Diarrhegmoides, Othniocera, Paraphasca, Phasca and Xenosophira, all with similar wing patterns and bare spermathecae with protruding nipple-like apices. Stigmatomyia, Gressittidium and Epinettyra were added subsequently by Hancock and Drew (2003) or Hancock (201 1b). Although Epinettyra has an atypical wing pattern, its spermathecae are similar to those of Othniocera (cf. Permkam and Hancock 1995 and Hardy 1986). The genera form an apparently closely related and monophyletic group but their interrelationships are difficult to determine. Korneyev (1999) noted that the Phascini and Phytalmiini both had the vanes of the phallapodeme fused into a Y-shaped structure (a homoplasious 26 Australian Entomologist, 2016, 43 (1) apomorphy that also occurs in most genera of Acanthonevrini and is thus possibly plesiomorphic within the subfamily). The distinctive, ivory-white medial vitta against a black scutum, seen in most species of Phascini (e.g. Phasca trifasciata Hardy: Hancock and Drew 2003), resembles that of Ortaloptera callistomyia (Fig. 2) and suggests a close relationship. Phascini differ from Phytalmiini and Epacrocerini in having only one of the midtibial apical spines well developed (apomorphy). All genera occur in New Guinea with the exception of Epinettyra, with its sole species E. setosa Permkam & Hancock known only from Iron Range and the Atherton Tableland in northeastern Queensland. Tribe Epacrocerini This tribe was regarded as distinct and defined by Korneyev (1994, 1999), based on the Epacrocerus group of Hardy (1982), to include four genera 4 Epacrocerus, Proepacrocerus, Tanymetopus and Udamolobium, with Sophiropsis later added by Hancock and Drew (2003). Proepacrocerus and Sophiropsis do not have the lobate second antennal segment seen in all the other genera but are associated on other characters, including the non-lobate wing cell bcu and the presence of one pair each of long frontal and orbital setae; these two genera are likely to be primitive within the tribe. Tanaodema Hardy is added to the group here; although unusual in many respects (Hardy 1987), it has the characteristic lobe on the inner margin of the second antennal segment (apomorphy), plus a non-lobate cell bcu and a greatly modified head, the latter apparently an extreme version of the modifications present in Tanymetopus and Udamolobium that are much less developed in Epacrocerus. In Udamolobium and Tanaodema the lobe of the second antennal segment is broadly rounded [anteriorly produced in Epacrocerus (sharply) and Tanymetopus (bluntly)], the wing pattern is somewhat reticulate posteriorly and the pterostigma is longer than cell c (apomorphies); these appear to be the most specialised genera. Several other apomorphies seen in Tanaodema 4 the narrow and sharply acute pterostigma, lack of a costal spine above the apex of vein Sc and presence of only one pair of scutellar setae 4 appear to be homoplasious with Phytalmiini. Korneyev (1999) noted that the eversible membrane of the ovipositor in Epacrocerini is impregnated by sclerotised, apparently dentiform structures, similar to that in tribe Phascini. The aculeus, with its blunt apex and four pairs of preapical setae, is very similar (although broader subapically) to that of Paraphasca, while the spermathecae (bare with a protruding, nipple-like apex, at least in Sophiropsis), closely resemble those of the Phascini, also suggesting a close relationship. As in Phytalmiini, at least two of the midtibial apical spines are well developed. The six genera are known only from New Guinea, with one species, Tanymetopus claripennis Hardy, extending as far east as New Britain.