Luciolinae, Lacordaire, 1857

Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, Zootaxa 4687 (1), pp. 1-174 : 11-25

Key to genera of Luciolinae using males

1. Aedeagus with LL visible at sides of ML when viewed either from above or below; aedeagal sheath usually asymmetrical with posterior half of sheath emarginated on one or both sides ( Figs 5−8 )............................................. 2 Aedeagus with distal half or more of LL concealed behind the ML when viewed from below; aedeagal sheath usually symmetrical with posterior half not emarginated on either side ( Figs 1−4 )........................................... 21

2. Clypeolabral suture absent; labrum well sclerotised, inflexible, inflexibly joined to head; rare New Guinean genus known from 4 specimens ( Fig. 9 ).................................................................... Missimia Ballantyne Clypeolabral suture present; labrum not well sclerotised, flexible, junction between labrum and rest of head flexible....... 3

3. Aedeagal sheath surrounded by a set of three sclerites encased in a ring of muscle in life; elytral punctation parallel over much of the surface; LO entire with anterior margin emarginated, often deeply ( Figs 10−12 )................ Sclerotia Ballantyne No sclerites surrounding aedeagal sheath; elytral punctation not parallel; LO either entire or bipartite; entire LO sometimes with median anterior emargination............................................................................ 4

4. Tergite 8 asymmetrical ( Figs 13−15 )...................................................................... 5 Tergite 8 symmetrical.................................................................................. 6

5. Tergite 8 asymmetrical because of emargination on left side ( Fig. 13 )....................... Asymmetricata Ballantyne Tergite 8 asymmetrical because of left pointing lobe arising beneath the tergite ( Figs 14, 15 , 233−237 )............................................................................................. Kuantana Ballantyne gen. nov.

6. Elytra with well-defined humeral carina; elytral punctation conspicuous, broad, deep and evenly spaced, as wide as pronotal punctation; LL usually appearing of uneven length ( Fig. 16 ).................................... Curtos Motschulsky Elytra without a well-defined humeral carina; elytral punctation not conspicuous, usually pin punctate and not appearing larger than pronotal punctation; LL of aedeagus usually of equal length................................................ 7

7. LO in V7 triangular in outline, outline following outline of V7, not reaching to sides or posterior margin of V7; aedeagus slightly asymmetrical with ML bent at apex and LL slightly uneven in length ( Figs 17, 18 )......... Triangulara Pimpasalee LO not triangular in outline; aedeagus symmetrical with LL of even lengths....................................... 8

8. LO in V7 usually retracted into anterior half or more, with posterior margin often strongly emarginated; posterior area of V7 often swollen or arched, always with underlying muscle impressions visible (e.g. Figs 19–21 ); PLP absent and lateral margins of V7 curving smoothly posteriorly, not subparallel sided; MPP strongly developed but never abruptly curved dorsally at its apex; aedeagal sheath asymmetrical, sternite posterior to tergite articulations emarginated on its right side ( Figs 5−8 )...... 9 LO usually occupying most of V7 and reaching to sides if not to posterior margin; if retracted to anterior area then posterior margin not emarginated, posterior area of V7 not swollen or arched, and muscle impressions not developed beneath cuticle; lateral margins of V7 often subparallel-sided but PLP not developed; MPP not strongly developed.................... 10

9. Species often with orange pronotum and dark brown elytra which may be margined ( Figs 22−25 ); ML with lateral teeth ( Fig. 26 ); T8 with membranous projections from base of anterolateral prolongations ( Fig. 27 ); ventral surface of T8 with asymmetrical ridge in median posterior area ( Figs 28, 29 ); aedeagal sheath with basal portion of sternite posterior to the tergite articulations subparallel-sided then emarginated on right side ( Fig. 29 ); LL not narrowed or out turned; Pacific Island species mainly from the Solomon Islands................................................................. Pacifica Ballantyne Species usually with cryptic colouration ( Figs 30−34, 37 ), brownish dorsum with brown medial markings on pronotum, at base of elytra adjacent to the MS, and at or near elytral apices; ML without lateral teeth ( Fig. 36 ); T8 without lateral projections as described above; aedeagal sheath sternite emarginated from the point of the tergite articulations; LL often narrowed at their apices and out-turned ( Figs 35, 36 ); New Guinean species................................. Pygatyphella (Ballantyne)

10. Aedeagal sheath symmetrical; sheath sternite elongate slender, either expanding gradually behind the tergite articulations ( Figs 39, 40 ), or slender and subparallel-sided for most of its length with expansion laterally near its apex ( Fig. 38 ); sheath sternite often terminated by paired lobes; MPP may be elongated and bent dorsally before its tip; T 8 may incline ventrally and partially envelop the apex of the MPP ( Figs 41−42 )................................................................ 11 Aedeagal sheath asymmetrical, with sternite emarginated on its right side from the point of the tergite articulations; or sheath symmetrical, broad, not emarginated on its right side; MPP usually short, apically rounded; if MPP longer then apex does not incline dorsally...................................................................................... 13

11. Posterior margin of V7 rounded; base of LL fused along their length with very short membranous projections at their posterolateral corners; inner ventral margins of LL with elongate ‘sausage’ shaped lobes ( Figs 44−49 )..................................................................................................... Abscondita Ballantyne et al. Posterior margin of V7 either rounded as above or with strongly developed MPP and median projection of T8 which may incline towards each other ( Figs 41, 42 ); basal portion of LL not fused along their length, with very elongated membranous apical portions; inner ventral margins of LL may have short, slightly rounded lobes ( Figs 50, 51 ).......................... 12

12. Posterior margin of V7 rounded, T8 without posterior median projection ( Fig. 43 )............. Pygoluciola sensu Fu et al. Posterior margin of V7 produced, usually inclining dorsally where it may be enveloped by the downturned tip of a median projection of T8 ( Figs 41, 42 )....................................................... Pygoluciola Wittmer s. str.

13. Pronotum subparallel-sided, with hypomeron widely flattened and upper and lower surfaces of the pronotum closely adpressed along its length ( Figs 52, 53 )................................................................. Emeia Fu et al. Pronotum usually with lateral margins divergent posteriorly; if subparallel sided then hypomeron not widely flattened; if surfaces adpressed then this only in posterior half of hypomeron………………………………………………………………………14

14. ML strongly laterally compressed, curved along its length with preapical portion produced and pointed; LL often expanded at their apices such that width across apices is sub equal to the width across bases; LL usually with elongate slender apically pointed lobes arising from their inner ventral surfaces....................................... Luciola Laporte s. str. ML not strongly laterally compressed, not strongly curved along its length, nor with a preapical point; LL not strongly expanded at their apices; no elongate slender apically pointed lobes arising from their inner ventral surfaces of LL............... 15

15. Aedeagal sheath sternite posterior to the tergite articulations with both sides irregularly emarginated; aquatic species from China and Japan ( Figs 54−56 )............................................................. Aquatica Fu et al. Aedeagal sheath sternite posterior to the tergite articulations with only right side smoothly emarginated ( Figs 57, 58 )....................................................................................... Atyphella “complex”...16

16. Aedeagal basal piece hooded; pronotum orange yellow; elytra dark brown, never margined; elytral margins slightly convex sided; New Guinean species, one species from Cape York Peninsula in Australia ( Figs 59−61 ) ……..… Lloydiella Ballantyne Aedeagal basal piece not hooded; pronotum either concolourous or with median dark markings; elytra may be dark brown, or variously pale margined, striped, or pale with or without apical dark markings; elytra parallel sided or convex sided; Australian, New Guinean and Pacific Island species.................................................................. 17

17. Orange pronotum with pin punctures widely separated in lateral areas; elytra convex sided, dark brown with lateral, apical and sutural margins orange; ML aedeagus constricted preapically ( Figs 62, 63 )........................ Convexa Ballantyne Pronotum either concolourous or with median dark markings; pronotal punctures close, not widely separated in lateral areas; elytra either convex or parallel sided; if species from the Solomon islands with pale elytral margins then sutural and apical margins dark; elytra either dark brown, or variously pale margined, striped, or pale with or without apical dark markings; Aus- tralian, New Guinean and Pacific Island species ………………………………………………………………………………18

18. Dorsal colouration pale brownish yellow or orange yellow, sometimes with dark brown elytral apices................. 19 Dorsal colouration never as above; pronotum often with dark medial markings; elytra either uniformly dark brown, dark brown with paler margins, or ‘striped’ (with paler longitudinal lines that may correspond to the interstitial lines)............... 20

19. Dorsum orange yellow with elytral apices black tipped; pronotal lateral margins either subparallel-sided or divergent; elytra parallel-sided; apical labial palpomere with inner margin dentate; one species from far north Australia ( Fig. 64 )....................................................................................... Aquilonia Ballantyne partim Dorsum brownish yellow with elytral apices black tipped or not; pronotal lateral margins divergent posteriorly; elytra slightly convex sided; apical labial palpomere with inner margin irregular, not dentate; two species known from islands off the eastern coast of the island of New Guinea ( Figs 65, 66 )....................................... Aquilonia Ballantyne partim

20. Orange pronotum and black elytra having pale lateral margins only (pale band does not extend around apex or along sutural margin); pronotal lateral margins divergent posteriorly; posterior area of hypomeron not closely adpressed; elytra parallel sided; MS black; LO in V7 entire; antennal sockets not contiguous; apical labial palpomere irregularly incised along its inner margin, not dentate; known from one large wide species (up to 14 mm long and 5.3 mm wide) which is widely distributed in the Solomon Islands ( Fig. 67 ).................................................................. Magnalata Ballantyne Dorsum variously coloured ( Figs 68, 69 ). If elytra dark brown with pale margins then both lateral and sutural margins pale; pronotal lateral margins divergent posteriorly or subparallel-sided; both anterior and posterior areas of hypomeron flattened and closely adpressed in Australian species, or only posterior area strongly adpressed; elytra parallel or convex sided; MS usually pale coloured; LO in V7 usually entire, bipartite in two species; antennal sockets sometimes contiguous; apical labial palpomere usually dentate with more than two teeth, irregular and not dentate in two species, ovoid entire in several species......................................................................................... Atyphella Olliff

21. Antennae with basal 4 flagellar segments gently serrate; LO in V7 entire with an anteromedian triangular emargination ( Figs 70, 71 )....................................................................... Serratia Ballantyne gen. nov. No antennal segments serrate; LO in V7 bipartite, if entire then without an anteromedian emargination................ 22

22. Posterior margin of elytra emarginate (usually only visible from beneath); posterior margin of V7 with well-defined MPP and rounded posterolateral corners ( Figs 73−75 )...................................... Emarginata Ballantyne gen. nov. Posterior margin of elytra not emarginated, may be deflexed.................................................. 23

23. Usually yellowish brown with or without black tipped elytra; LO in V7 bipartite with median longitudinal carina between LO halves; PLP oblique; MPP strongly developed, wider than PLP (when viewed from beneath), and projecting beyond them, MPP apically emarginated bearing well defined hooks which curve dorsally and may not be visible from beneath; FS 7−9 shorter than remainder ( Figs 72, 76, 77 )...................................................... Colophotia Motschulsky LO in V7 either entire or bipartite, if the latter then LO halves not separated by a median carina; PLP horizontal; MPP never strongly developed as described above; if MPP apically emarginated and prolonged into points then these short, may not be visible from beneath.................................................................................. 24

24. Median lobe of aedeagus swollen in basal 2/3; LL with broad bases visible from beneath in basal 1/3 only, and very narrow apices which closely approach dorsally at about midpoint of ML, and are not visible from beneath; having orange pronotum and dark brown elytra ( Figs 78−80, 83 )......................................................... Inflata Boontop Median lobe of aedeagus not greatly swollen in basal 2/3, approximately the same width along its length; LL visible at sides of ML for about 2/3 its length, with apices closely approximate behind and near apex of ML where they are concealed from below; colour variable, may be as above or with brown margined elytra, or yellowish brown dorsum with black tipped elytral apices. .................................................................................................. 25

25. Metafemoral comb present ( Figs 87, 88 ).................................................................. 26 Metafemoral comb absent............................................................................. 27

26. LO in V7 bipartite ( Fig. 82 )............................................................................ 28 LO in V7 entire ( Figs 81, 85, 86 )............................................................ Pteroptyx Partim

27. Elytral apices deflexed ( Fig. 81 )............................................................ Pteroptyx Partim Elytral apices not deflexed; incurving lobes present along posterior margin of V7 between PLP and MPP ( Fig. 82 ); aedeagal sheath with paraprocts (e.g. Figs 89−91 ).................................................... Pyrophanes Olivier

28. LO in V7 entire; two species without deflexed elytral apices; aedeagal sheath without paraprocts ( Figs 81 , 92, 93 ).......................................................................................... Medeopteryx Ballantyne LO in V7 bipartite; species with or without deflexed elytral apices ( Figs 94, 95 )................................... 29

29. Elytral apices not deflexed; yellowish brown dorsum with black elytral apices; bipartite LO halves in V7 closely approach in median line; incurving lobes along posterior margin of V7 fleshy and apically rounded; PLP short and rounded ( Fig. 94 )......................................................................................... Pteroptyx testacea Elytral apices deflexed or not; usually with orange pronotum and brown to black elytra; New Guinean species; bipartite LO halves well separated in median line, not closely approaching; incurving lobes absent; PLP usually horizontal and widely expanded ( Fig. 95 )...................................................................... Trisinuata Ballantyne

Figures

FIGURES 1−8. Key to genera characters 1. 1, 2 Australoluciola anthracina type male (MNHN); 3, 4 Aus. japenensis paratype male; 5, 6 Atyphella scabra male; 7, 8 Aty. leucura type male (MNHN). 1, 2, 5, 6 aedeagus ventral (1, 5), and dorsal (2, 6) (lateral lobes are arrowed). 3, 4, 7, 8 aedeagal sheath dorsal (3, 7), and ventral (4, 8). Oblique arrows in anterior section indicate point where tergite arms join the lateral margins of the sheath sternite; vertical arrow pointing to foot of page in 3 indicates anterior margin of sheath tergite; vertical and horizontal lines in 4 indicate length (vertical) and width (horizontal) of sheath sternite; lower paired arrows in 8 indicate emarginated margin of sheath sternite.

FIGURES 9−16. Key to genera characters 2. 9. Missimia flavida type male; 10 Luciola brahmina type male (MNHN); 11, 12 Luciola seriata type male (CMG); 13 Asymmetricata circumdata male; 14, 15 Kuantana menayah gen. et sp. nov.; 16 Curtos costipenis male. 9 anterior head (arrow indicates clypeus); 10, 11, 16 dorsal (arrows in 16 indicate humeral carina); 12 lateral tergite 8 aedeagal sheath and sclerites); 13 dorsal tergites 7 and 8; 14, 15, dorsal (14) and ventral tergite 8. Figure legends: AS aedeagal sheath; T7, 8 tergites 7, 8.

FIGURES 230−237. Kuantana menayah gen. et sp. nov. type male. 230, 232 habitus dorsal (232 slightly to side with right elytron pulled to side); 233−237 abdomen: 233 ventral. 234 dorsal, 235 right lateral; 236, 237 terminal abdomen tergites 7 8: 237 from beneath with aedeagus and sheath removed; 238 from above (left side of tergite 7 damaged in extraction). Scale lines are 1 mm.

FIGURES 17−21. Key to genera characters 3. 17, 18 Triangulara frontoflava; 19 Pygatyphella obsoleta; 20, 21 Pygat. huonensis. 17 ventral whole body; 18, 19, 21 ventral terminal abdomen; 20 right lateral terminal abdomen.

FIGURES 22−29. Key to genera characters 4. 22, 23, 26−28 Pacifica salomonis male; 24, 29 Pac. limbatipennis (Guadalcanal BPBM); 25 Pac. limbatifusca paratype male. 22, 24, 25 dorsal; 23 ventral; 26 aedeagus ventral; 27, 28 tergite 8 ventral and from right side showing ventral surface; upper arrows in 27, 28 indicate projections from anterior tergal arms; lower arrow in 27 indicates oblique ridge on ventral surface; 29 aedeagal sheath ventral (upper arrow indicates beginning of emargination, lower arrow indicates emargination.

FIGURES 30−37. Key to genera characters 5. 30, 34, 35 Pygatypella undulata; 31 Pygat. wisselmerenia; 32, 36 Pygat. obsoleta; 33 Pygat. peculiaris; 37 Pygat. kiunga paratype male. 30−34, 37 dorsal (34 diagrammatic representation of colour pattern); 35 aedeagal sheath, ventral; 36 aedeagus ventral, line outline.

FIGURES 38−43. Key to genera characters 6. 38, 43 Pygoluciola qingyu; 39, 40 Abscondita pallescens comb. nov.; 41, 42 Pygoluciola hamulata type male. 38−40 aedeagal sheath ventral (38, 39) and slightly right lateral (40) (arrows indicate position where lateral tergal arms join the median sternite); 41, 43 terminal abdomen ventral; 42 termnal abdomen left lateral. Figure legends: V7 ventrite 7; T8 tergite 8.

FIGURES 44–49. Key to genera characters 7. Luciola cerata lectotype aedeagus. 44, 47 dorsal; 45, 48 ventral (lobes arrowed); 46, 49 left lateral. Figs 47–49 outline diagrams of photos immediately above. Figure legend: ML median lobe; BP basal piece.

FIGURES 50–56. Key to genera characters 8. 50, 51 Pygoluciola satoi; 52, 53 Emeia pseudosauteri; 54, 55, 56 Aquatica ficta. 50, 51 aedeagus dorsal (50: arrow indicates separation in basal area of lateral lobes), and ventral; 52, 56 dorsal habitus; 53, 55 ventral habitus;; 54 aedeagal sheath ventral semidiagrammatic (arrows indicate pointed projections along sides of sheath sternite).

FIGURES 57–63. Key to genera characters 9. 57, 58 Atyphella aphrogeneia; 59 Lloydiella majuscula; 60, 61 Ll. japenensis; 62, 63 Convexa wolfi. 57, 58 aedeagal sheath dorsal (57) and ventral (arrows indicates emargination of right side of sheath sternite); 59 dorsal and ventral, and 63 dorsal habitus; 60–62 aedeagus, 60 slightly right lateral; 61, 62 ventral.

FIGURES 64–69. Key to genera characters 10. 64 Aquilonia costata; 65, 66 Aq. similismessoria comb. nov.; 67 Magnalata limbata; 68 Atyphella flammans; 69 Aty. aphrogeneia.

FIGURES 70–77. Key to genera characters 11. 70–72 Serratia sibuyania gen. et sp. nov.; 73–75 Emarginata trilucida gen. nov. et comb. nov.; 76, 77 Colophotia praeusta. 70, 72, 73 dorsal habitus (70 anterior body only); 74 ventral habitus; 71 dissected terminal abdomen ventral; 75 ventrite 7 and elytral apices from below; 76, 77 ventrites 6, 7 from left (76) and below (77). Figure legends: MC median carina; MPP median posterior projection; PLP posterolateral projections.

FIGURES 78–86. Key to genera characters 12. 78–80, 83 Inflata indica; 81 Medeopteryx cribellata; 82 Pyrophanes appendiculata; 84 Australoluciola flavicollis; 85 Aus. nigra; 86 Aus. anthracina. Figure legends: LL1 basal expanded portion of lateral lobes; LL2 apical narrowed portion of lateral lobes; ML median lobe.

FIGURES 87–95. Key to genera characters 13. 87–91 Pteroptyx macdermotti; 92, 93 Medeopteryx sp.; 94 Pteroptyx testacea; 95 Trisinuata caudabifurca. 87 left hind leg (without coxa) 88 right hind leg detail femur and tibia; 89–93 aedeagal sheath ventral (89, 91, 92), left lateral (90, 93) (paraprocts arrowed in 89–91); 94, 95 ventrites 6, 7 from beneath (incurving lobes arrowed in 95). Figure legend: LO light organs.