Subfamily Limosininae Macquart, 1835

Key to species of the Pterogramma -like genera of the Pacific and Indomalayan regions

Couplets for the Pterogrammoides species are modified from, or quoted directly from, Papp (1989). Couplets for the Parapterogramma species originally described as Minialula are based on Hayashi (2013).

1. Mid tibia with a single dorsal bristle on distal half (as in Fig. 70A). Orbital bristles very small and fine (Fig. 70B). Wing long and narrow, crossveins in basal third of wing (Figs 70C, 72C). Abdominal sclerites strongly reduced or absent................................................2 ( Pterogrammoides Papp, 2008)

– Mid tibia with at least two dorsal bristles on distal half (as in Fig. 25A). Orbital bristles strong. Wing broad and ovoid, crossveins in middle of wing (Figs 40–41, 68). Abdominal sclerites desclerotized around edges but otherwise well-developed...................................................................................... 6

2. Scutum with distinct pattern of yellowish spots. Lateral sclerites of female T8 very large. One pair of orbital bristles; katepisternal bristle absent. Wing infuscate, darkest along veins .......................... ................................................................................................... Pt. poecilosomus Papp, 1972 (PNG)

– Scutum unicolourous, orange-brown to brown. Lateral sclerites of female T8 smaller. Two pairs of orbital bristles; katepisternal bristle present. Wing very lightly patterned ........................................ 3

3. Male S6–8 large, as wide as or wider than T1+2 (Papp 1989: figs 10, 26). Posteromedial process of surstylus thin, blade-like (Papp 1989: figs 15, 32). Postgonite strongly bent distally, apex truncate (Papp 1989: figs 14, 30)..................................................................................................................... 4

– Male S6–8 smaller, distinctly narrower than T1+2 (Papp 1989: figs 1, 16). Posteromedial process of surstylus stout, rounded (Papp 1989: figs 3, 17). Postgonite gently curved distally, apex tapered (Papp 1989: figs 5, 19)....................................................................................................................... 5

4. Length of male S6–7 1.3× epandrial width, epandrium with short setae (Papp 1989: fig. 10). “Postgonite curved in apical third” (Papp 1989: fig. 14). Surstylus with one large seta and one slightly smaller ventromedial tooth-like seta (Papp 1989: figs 13, 15) ............................................... .............................................................................................................. Pt. indicus Papp, 1989 (IND)

– Length of male S6–7 1.7× epandrial width, epandrium with longer setae (Papp 1989: fig. 26). “Postgonite curved in apical fifth” (Papp 1989: fig. 31). Surstylus with one very large ventromedial tooth-like seta and a much finer ventromedial seta (Papp 1989: figs 32–33)...................................... ................................................................................................................. Pt. thaii Papp, 1989 (THA)

5. Wing reduced, extending at most to end of T4. Postgonite “arcuately bent in its whole length, apex rounded, posterior margin serrate” (Fig. 71B). Surstylus anteroapically bare, with two stout apical setae, ventromedial tooth-like setae large and flat (Fig. 71A)............. Pt. baloghi Papp, 1972 (PNG)

– Wing extending beyond end of abdomen. Postgonite “more strongly bent, apex narrower”, posterior margin smooth (Papp 1989: fig. 19). Surstylus anteroapically densely setulose, without stout apical setae, ventromedial tooth-like setae small, conical (Papp 1989: figs 17–18) ..................................... ..................................................................................................... Pt. longipennis Papp, 1972 (PNG)

6. Head and thorax without a pattern of microtomentose areas, often bare and shiny (as in Fig. 51B). Anterior pair of dorsocentral bristles arising just before suture (as in Fig. 54A)................................ ...................................................................................................7 ( Pseudopterogramma Papp, 2008)

– Head and thorax with silvery microtomentum forming patterns (as in Fig. 2B). Anterior pair of dorsocentral bristles arising just behind suture (as in Fig. 3A) ........................................................... .....................................................................................................18 ( Parapterogramma Papp, 2008)

7. Postpedicel conical apically. Surstylus with stout tooth-like apical setae (Papp 2008: fig. 82). Female epiproct and hypoproct strongly reduced. M 4 (= CuA of most recent authors) usually ending at dm-m (= dm-cu), sometimes extending very slightly beyond (Papp 2008: fig. 318)............................................................................ Ps. metatarsalis (Papp, 2008) (SGP, THA)

– Postpedicel rounded apically. Surstylus never with stout tooth-like apical setae. Female epiproct and hypoproct well-developed and distinct. M 4 usually extending beyond dm-m (Fig. 68).................... 8

8. Hind tarsomeres greatly expanded and dorsoventrally flattened, fore tarsomeres slightly flattened (Fig. 43A) ................................................................................. Ps. aestivale (Richards, 1973) (AUS)

– Fore and hind tarsomeres cylindrical, sometimes swollen but not dorsoventrally flattened (as in Fig. 66A)............................................................................................................................................ 9

9. Wing largely hyaline, with a dark spot around R 1 or R 2+3 (Fig. 58B–C, G) .................................... 10

– Wing distinctly patterned, with light and dark areas (Fig. 58A, D–F, H–J) .................................... 12

10. Interfrontal bristles in four large pairs, approximately as thick as anterior orbital bristle (Fig. 48B). Hind tibia uniformly brown (Fig. 48A). Surstylus divided into anterior and posterior lobes (Fig. 49B)................. Ps. brevivenosum (Tenorio, 1967) (COK, FJI, PNG, PYF, TON, USA, WSM)

– Interfrontal bristles in three small pairs, distinctly weaker than anterior orbital bristle (Figs 46B, 60B). Basal third to half of hind tibia yellow, distinctly paler than apical half (Figs 46A, 60A). Surstylus with a single lobe, either curving anteriorly (Fig. 47B) or posteriorly (Fig. 61B) ...........11

11. Male mid tibia with a double row of stout apicoventral setae. M 4 extending distinctly beyond dm-m (Fig. 68B). Surstylus curved anteriorly, apex pointed (Fig. 47B). Postgonite very slightly sinuate, very narrow, apex simple (Fig. 47A). Basiphallus large (~0.6× length of postgonite)....................... ............................................................................. Ps. annectens (Richards, 1964) (IDN, PNG, SLB)

– Male mid tibia with a small midventral bristle and a small apicoventral bristle. M 4 not developed past dm-m (Fig. 68G). Surstylus curved posteriorly, apex rounded (Fig. 61B). Postgonite curved, broader (length ~5.5 × medial width), apex with several spine-like inner lobes (Fig. 61A). Basiphallus small (~0.3 × length of postgonite)................................................................ Ps. kasut sp. nov. (IDN, SGP)

12. Hind tibia with a large dorsal preapical seta (as in Fig. 66A) ......................................................... 13

– Hind tibia with uniformly short dorsal setae (as in Fig. 54A)......................................................... 16

13. Fore and mid tibiae brown with central third yellow, sometimes yellow area indistinct (Fig. 51A). Male S5 with a posteroventral flange on left third (Fig. 52C). Posterior lobe of surstylus elongate and sinuate, relatively parallel-sided throughout (Fig. 52B). Postgonite with three distinct, strong bends, strongly tapered after apical bend (Fig. 52A) ...................................................................................... ......................................................... Ps. conicum (Richards, 1946) (AUS, FSM, GUM, PLW, PNG)

– Fore and mid tibiae solid brown, without any yellow banding (as in Fig. 57A). Male S5 simple, without any additional posteroventral lobes (Fig. 58C; Papp 2008: figs 121, 130). Posterior lobe of surstylus triangular (Fig. 58B) or rectangular (Papp 2008: figs 126, 132). Postgonite gently sinuate and gradually tapered...................................................................................................................... 14

14. Scutum yellow-orange, contrasting strongly with dark brown pleuron (Fig. 57A). Male S5 asymmetrical, strongly reduced on right side and rectangular on the left; setae limited to a transverse row of eight posteromedial setae (Fig. 58C). Posterior lobe of surstylus distinctly smaller than anterior lobe (Fig. 58B) ............................................................................... Ps. fijiensis sp. nov. (FJI)

– Scutum brown, concolourous or nearly so with pleuron (as in Fig. 66A). Male S5 almost symmetrical, both sides rectangular and well-developed; uniformly short-setose (Papp 2008: figs 121, 130). Posterior lobe of surstylus distinctly larger than anterior lobe (Papp 2008: figs 126, 132)............. 15

15. Posterior lobe of surstylus with a short (length 1.0× height) apicoventral point, anterior lobe smooth ventrally (Papp 2008: fig. 132). Postgonite slightly broadened subbasally (Papp 2008: fig. 133)...... ............................................................................................... Ps. insulare (Papp, 1972) (IDN, PNG)

– Posterior lobe of surstylus with a long (length 2.1 × height) apicoventral point, anterior lobe with a triangular posteroventral lobe (Papp 2008: fig. 126). Postgonite almost parallel-sided, gradually tapered and not broadened subbasally (Papp 2008: fig. 128) .............................................................. ..................................... Ps. siamense Papp, 2008 (IDN, IND, MYS, NPL, PHL, SGP, THA, VNM)

16. Mid tibia brown with two yellow rings. R 4+5 almost straight (Richards 1973: fig. 63). Surstylus with very few setae, limited to ventral surface (Richards 1973: fig. 69) .................................................... ................................................................................................. Ps. deemingi (Richards, 1973) (PNG)

– Mid tibia solid yellow-brown, without distinct rings. R 4+5 curved (Figs 68E, H). Surstylus with many setae, some on lateral surface (Figs 55B, 64B) ............................................................................... 17

17. Fore and hind tibiae solid yellow (Fig. 54A). R 2+3 distinctly sinuate (Fig. 68E). Male S5 uniformly short-setose (Fig. 55C). Surstylus triangular, about as tall as epandrium (Fig. 55B). Postgonite strongly bent forwards, thick and asymmetrically bifid apically (Fig. 55A) ...................................... .............................................................................................. Ps. fenestrata (Richards, 1973) (AUS)

– Fore and hind tibiae distinctly banded (Fig. 63A). R 2+3 almost straight (Fig. 68H). Male S5 with a transverse row of very long medial setae (Fig. 64C). Surstylus bent at an obtuse angle, about a quarter the height of epandrium (Fig. 64B). Postgonite gently bisinuate and slender (Fig. 64A) ............................................................................... Ps. ketambensis sp. nov. (IDN, MYS)

18. Costa ending at apex of R 4+5. One postpronotal bristle. Mid tibia with two posterodorsal bristles on proximal half. Male S6 with a knob-like lobe (Papp 2008: fig. 98)................................................ 19

– Costa usually extending far beyond apex of R 4+5 (only slightly beyond in Pa. magnum sp. nov.). Two postpronotal bristles. Mid tibia with one posterodorsal bristle on proximal half. Male S6 simple, without a knob-like lobe .................................................................................................................. 21

19. Male mid tibia with a long, hair-like apicoventral seta (Hayashi 2011: fig. 2). Anterior lobe of surstylus bent at ~90° angle in relation to posterior lobe (Papp 2008: fig. 99) ................................... ........................................................................................ Pa. poeciloptera (Papp, 2008) (JPN, TWN)

– Male mid tibia with stout apicoventral bristle. Anterior lobe of surstylus bent at an angle of <90° in relation to posterior lobe (Hayashi 2013: figs 6, 9) ......................................................................... 20

20. Male S5 with uniform setae and a slight posteromedial emargination (Hayashi 2013: fig. 5). Postgonite bent about 70° at midlength, apex pointed (Hayashi 2013: fig. 7) .................................... ...................................................................... Pa. continentalis (Hayashi, 2013) (NPL, THA, VNM)

– Male S5 bare medially with a short posteromedial lobe (Hayashi 2013: fig. 8). Postgonite bent about 30° at midlength, apex rounded (Hayashi 2013: fig. 10).......... Pa. tropicalis (Hayashi, 2013) (IDN)

21. Orbital plate with strong silver-gold microtomentose line connecting orbital and inner vertical bristles (as in Fig. 2B)...................................................................................................................... 22 – Orbital plate with silver-gold microtomentum limited to bases of orbital and inner vertical bristles (as in Fig. 8B) .................................................................................................................................. 27

22. Costa extending slightly beyond (1–2 × costal width) end of R 4+5; M 1 distinct beyond dm-m (Fig. 40G). Scutum with dark medial longitudinal stripe (Fig. 17B). Posterior lobe of surstylus rectangular, much larger than anterior lobe (Fig. 18B).......................... Pa. magnum sp. nov. (AUS)

– Costa extending well beyond (5–6× costal width) end of R 4+5; M 1 indistinct beyond dm-m (Fig. 40A– F, H). Scutum with pale medial longitudinal stripe (as in Fig. 2B). Posterior lobe of surstylus various but usually smaller than anterior lobe; if larger than or comparable to anterior lobe, posterior lobe triangular (Fig. 24B)........................................................................................................................ 23

23. R 2+3 straight for most of its length, usually bent towards costa apically (Figs 40A, 41A) .............. 24

– R 2+3 gently to strongly curved for entire length (Figs 40E–F, 41C)................................................. 25

24. Male mid tibia with a stout apicoventral bristle and two rows of stout ventral setae. M 4 indistinct beyond dm-m (Fig. 40A). Male S5 projecting posteromedially and with setae restricted to posterolateral margins (Papp 2008: fig. 116) ................ Pa. asiaticum Papp, 2008 (IDN, SGP, THA)

– Male mid tibia with a small stout apicoventral bristle only. M 4 short but tubular beyond dm-m (Fig. 41A). Male S5 rectangular and desclerotized posteromedially with relatively uniformly distributed setae (Fig. 24D) ...................................................................... Pa. ofiensis sp. nov. (PNG)

25. Head yellow, scutum yellow with distinct brown spots, lower two-thirds of anepisternum and below dark brown (Fig. 14). Male mid tibia with a long apicoventral bristle. Male S5 with thickened, tooth-like setae along posterior margin (Fig. 15D). Surstylus trilobed, with a small inner median lobe and larger triangular anterior and posterior lobes (Fig. 15B) ............... Pa. hieroglyphicum sp. nov. (FJI)

– Head caramel-brown, scutum brown with dark and light longitudinal stripes, sides of thorax brown (Figs 11A, 28A). Male mid tibia with a short apicoventral bristle, with or without two rows of stout ventral setae. Male S5 various, relatively bare (Fig. 12D) or densely setose laterally (Fig. 29D), but never with tooth-like posterior setae. Surstylus various but always bilobed, never with a small inner medial lobe (Figs 12B, 29B)............................................................................................................ 26

26. M 4 indistinct beyond dm-m; CuA+CuP separated from wing margin by 0.7× its length (Fig. 40F). Male S5 with relatively sparse posterolateral setae (Fig. 12D). Postgonite significantly reduced, 0.5 × length of distiphallus (Fig. 12C). Spermathecal bulb spherical (Fig. 13D–E) ........................... ........................................................................................................... Pa. gonitellum sp. nov. (AUS)

– M 4 tubular beyond dm-m, extending 0.7–0.8 × length of dm-m; CuA+CuP separated from wing margin by 0.3–0.4× its length (Fig. 41C). Male S5 with densely long-setose posterolateral lobes (Fig. 29D). Postgonite not reduced, 1.3 × length of distiphallus (Fig. 29C). Spermathecal bulb cylindrical with deep basal and apical invaginations (Fig. 30D–E) .............. Pa. pollex sp. nov. (FJI)

27. M 4 distinct and tubular beyond dm-m, lower corner of dm usually angular ................................... 28

– M 4 indistinct beyond dm-m, lower corner of dm usually relatively rounded.................................. 32

28. Scutum shiny yellow-orange, without any microtomentose stripes; frons yellow-orange, concolourous with scutum (Fig. 3B). Posterior lobe of surstylus with long ventral setae, almost as long as lobe itself (Fig. 4B).............................................................................................. Pa. bicolor sp. nov. (FJI)

– Scutum dull brown with pale brown and grey microtomentose stripes; frons reddish caramel-brown, distinctly different from scutal colour (as in Fig. 8B). Posterior lobe of surstylus with shorter ventral setae, half as long as lobe at most (Figs 9B, 26B, 35B, 38B).......................................................... 29

29. Scutum with dark central microtomentose stripe (Fig. 8B). R 2+3 almost straight and reaching costa at an angle of 25–30°, resulting in a very narrow r 1; CuA+CuP short, indistinct beyond level of M 4 (Fig. 40D). Male mid tibia with a single, long apicoventral bristle. Male S5 with an elongate posterolateral extension on left side (Fig. 9D)................................ Pa. digitisternum sp. nov. (AUS)

– Scutum with pale central microtomentose stripe (as in Fig. 37B). R 2+3 distinctly curved and reaching costa at an angle of 60–80°, resulting in a much broader r 1; CuA+CuP longer, distinct beyond level of M 4 as a coloured pseudovein (Fig. 41B, E–F). Male mid tibia with a small apicoventral bristle, with or without 1–2 rows of stout ventral setae. Male S5 roughly symmetrical, without an elongate posterolateral extension (Figs 26D, 35D, 38D) ............................................................................... 30

30. Microtomentum of orbital plate silver-white, strongly reflective; scutum dark brown with conspicuous greyish microtomentum (Fig. 34B). Male S5 relatively simple with only scattered setae along posterior margin (Fig. 35D). Epandrium with uniformly sized setae; male cercus with 1–2 setae (Fig. 35B). Anterior lobe of surstylus largely bare and preapically constricted (Fig. 35B) .................................................................................................. Pa. simplex sp. nov. (FJI)

– Microtomentum of orbital plate silver-gold; scutum pale brown to brown (Figs 25B, 37B). Male S5 with large, densely setose posterolateral arms (Figs 26D, 38D). Epandrium with one distinctly enlarged lateral seta on either side of anal fissure; male cercus with 8–10 setae (Figs 26B, 38B). Anterior lobe of surstylus with several ventral setae and evenly tapered apically (Figs 26B, 38B) ... ......................................................................................................................................................... 31

31. Male mid tibia with a weak apicoventral bristle and a row of stout ventral setae. Male S5 with a more rectangular or ovoid posteroventral emargination, lateral arms with shorter setae (<0.4× lateral S5 length) (Fig. 38D). Anterior lobe of surstylus rectangular, posterior lobe flattened, curved outwards, and longer than anterior lobe (Fig. 38B). Postgonite tapered apically; epiphallus absent (Fig. 38C).................................................................................................... Pa. tokotaai sp. nov. (FJI)

– Male mid tibia with a stout apicoventral bristle only. Male S5 with a more triangular posteroventral emargination, lateral arms with longer setae (> 0.5× lateral S5 length) (Fig. 26D). Anterior lobe of surstylus subtriangular, posterior lobe stouter and shorter than anterior lobe (Fig. 26B). Postgonite bifurcate apically; epiphallus present (Fig. 26C)..................................... Pa. peckorum sp. nov. (FJI)

32. Scutum uniformly chocolate brown, lacking microtomentose stripes (Fig. 6B). Male mid tibia with a stout apicoventral bristle only. Postgonite boot-shaped with many ventral tooth-like knobs; distiphallus with sclerotized tooth-like lateral projections (Fig. 7C) ... Pa. dentatum sp. nov. (PNG)

– Scutum with distinct microtomentose stripes (Figs 20B, 31B). Male mid tibia with a long apicoventral bristle, with or without a row of stout ventral setae. Postgonite various but ventral surface smooth, without tooth-like knobs; distiphallus with membranous lateral lobes only, lacking sclerotized lateral projections (Figs 21C, 32C)............................................................................................................. 33

33. R 2+3 almost straight, reaching costa at an angle of 40–45°, resulting in a narrower r 1 (Fig. 41D). Acrostichal setulae in 6–8rows between anterior dorsocentrals.Male mid tibia with a long apicoventral bristle only. Male S5 broad with sparse setae and a pair of triangular posterior projections surrounding a small posteromedial emargination (Fig. 32D). Epandrium with uniformly sized setae; anterior lobe of surstylus much larger than posterior lobe (Fig. 32B) ......... Pa. pugioformis sp. nov. (AUS, PNG)

– R 2+3 strongly curved, reaching costa at an angle of 60–70°, resulting in a broader r 1 (Fig. 40H). Acrostichal setulae in 9–10 rows between anterior dorsocentrals. Male mid tibia with a row of stout ventral setae in addition to a long apicoventral bristle. Male S5 with large, densely setose posterolateral arms surrounding a very large posteromedial emargination (Fig. 21D). Epandrium with a distinctly enlarged seta on either side of anal fissure; anterior lobe of surstylus slightly smaller than posterior lobe (Fig. 21B)....................................................................... Pa. matau sp. nov. (FJI)