Luciolinae

Fu, Xinhua, Ballantyne, Lesley & Lambkin, Christine, 2012, Emeia gen. nov., a new genus of Luciolinae fireflies from China (Coleoptera: Lampyridae) with an unusual trilobite-like larva, and a redescription of the genus Curtos Motschulsky, Zootaxa 3403, pp. 1-53 : 7-8

Key to genera and species groups of Luciolinae using males

This key reflects the results obtained in the phylogenetic analysis herein (see Figs 7 – 9 ) and is expanded from Ballantyne and Lambkin (2009). All species groups related to the genus Luciola ( Fig. 7 ) and suggested by the present phylogenetic analysis are keyed. Genera related to Atyphella Olliff are treated in the form recognised by Ballantyne and Lambkin (2009). The Luciolinae genera recognised here are outlined in Table 3.

1. Clypeolabral suture absent; labrum well sclerotised, inflexible, inflexibly joined to rest of head; in specimen held horizontally and viewed from beneath humerus visible, not covered by epipleuron; posterolateral corners of pronotum very acute and pointed ( Ballantyne & Lambkin 2009 Figs 21, 22 , 27, 29, 30 , 32–34 )............................ Missimia Ballantyne

- Clypeolabral suture present ( Ballantyne & Lambkin 2009 Figs 28 , 31), labrum not well sclerotised, flexible, junction between rest of head and labrum flexible; in specimen held horizontally and viewed from beneath humerus not visible, is covered by epipleuron; posterolateral corners of pronotum not very acute and pointed.........................................2

2. When viewed from the side midlateral margins of pronotum elevated (appear sinuate from side); asymmetrical ML of aedeagus with acute apex, finely serrate along dorsal edges, bearing infolding flaps behind apex; left LL with preapical flap (right LL lacking flap); from beneath LL just visible at sides of ML, both apices not usually visible in same plane, separated in apical 1/3 only; aedeagal sheath sternite posterior to tergite articulations subparallel-sided in basal 1/3, then unevenly emarginated on both sides and narrowing to a slender obliquely truncate apex ( Ballantyne & Lambkin 2009 Figs 38–55 ; Fu & Ballantyne 2008 Figs 41–55 )............................................................................ Photuroluciola Pic

- When viewed from the side midlateral margins of pronotum not elevated; if ML of aedeagus asymmetrical then lacking apex as above; no serrations along ML apex nor infolding flaps behind apex; neither LL with preapical flap; viewed from beneath LL either concealed behind ML at their apices or widely visible at the sides of the ML; aedeagal sheath subparallel-sided in basal portion or laterally emarginate; if subparallel-sided not emarginated on both sides towards its apex nor narrowing to an obliquely truncate apex ( Ballantyne & Lambkin 2009 Figs 35, 36 , 37 )............................................ 3

3. LO absent, or present only in V6; eyes with strong posterior emargination; dark almost black dorsal colouration ( Ballantyne & Lambkin 2009 Fig. 59).... Bourgeoisia antipodum (Bourgeois) , B. hypocrita (Olivier) , undescribed New Caledonian species

- Eyes with or without strong posterior emargination; LO present in V7 (may be entire or bipartite); dorsal colouration variable, seldom as above...................................................................................... 4

4. LL of aedeagus widely visible at sides of ML when aedeagus viewed from beneath; pronotum wider than or subequal to humeral width, rarely less than ( Ballantyne & Lambkin 2009 Figs 35 , 73, 76)...................................... 5

- LL of aedeagus not visible at sides of ML when aedeagus viewed from beneath; pronotum always narrower than humeral width ( Ballantyne & Lambkin 2009 Figs 36 , 75)............................................................ 22

5. Elytral humeral carina absent; elytral punctation not conspicuous, not evenly spaced, not as wide and deep as that of pronotum; MPP short, squat, barely produced; aedeagus with LL as wide at their bases as at their apices; LL bearing elongate slender flattened lobes along inner (ventral) margins; ML strongly laterally compressed, curving ventrally with preapical point (as in Ballantyne & Lambkin 2009 Fig. 514) ( Ballantyne 1968 Figs 162, 169)............................. Luciola s. str. Partim1

- Elytral humeral carina present ( Ballantyne & Lambkin 2009 Figs 57, 58) or absent; elytral punctation either not conspicuous, not evenly spaced, not as wide and as deep as that of pronotum, or conspicuous, evenly spaced and similar in size to that of pronotum ( Ballantyne & Lambkin 2009 Figs 57, 58); MPP always obviously developed, sometimes L>W with apex inclined dorsally ( Ballantyne & Lambkin 2009 Figs 60, 61); aedeagal LL always narrower at apices than at bases; LL lacking elongate slender flattened lobes on inner (ventral) margins; ML not strongly laterally compressed nor with preapical point......... 6

6. Elytral humeral carina developed; punctation of elytra conspicuous, evenly spaced, as wide and deep as that of pronotum; LL of uneven length ( Ballantyne & Lambkin 2009 Figs 57, 58)..................................... Curtos Motschulsky

- Elytral humeral carina not developed, punctation of elytra not any larger and deeper than that of pronotum, usually smaller and narrower; LL of even length............................................................................. 7

7. Pronotum subparallel-sided, with hypomeron widely flattened and surfaces closely adpressed along its length..................................................................................... Emeia gen. nov. Fig. 8 Number 19

- If pronotum subparallel-sided then hypomeron not widely flattened with surfaces closely adpressed along its length....... 8

8. Aedeagal sheath surrounded by a set of 3 sclerites (2 lateral and one ventral; Fig. 43 ) ( Jeng et al. 2003 Fig. 14 ); at least some punctures in elytron in semiparallel rows; anterior margin of entire light organ emarginated, often deeply ( Figs 44, 45 )...... Luciola carinata Gorham , L. substriata Gorham , L. aquatilis Thancharoen , L. seriata Olivier , L. cingulata Olivier , L. brahmina Bourgeois

- Aedeagal sheath lacking sclerites surrounding it; elytral punctation not in lines; anterior margin of entire light organ not emarginated.............................................................................................. 9

9. Light organ triangular in outline, straight sides tapering posteriorly to a narrow rounded apex. ( Fig. 46 ).................................................................................... undescribed Thailand species (see Table 2 )

- Light organ not triangular in outline...................................................................... 10

10. Posterior half of aedeagal sheath (posterior to tergite articulations at sides) irregularly emarginate along both sides ( Jeng et al. 2003 Figs 29–31 ); aquatic species from China and Japan; Fu et al. 2010 Figs 5 , 13, 14 , 22–24 , 36 , 37 )........................................................................................... Aquatica Fu, Ballantyne & Lambkin

- If posterior half of aedeagal sheath (posterior to tergite articulations at sides) emarginate this is evenly emarginate on one side only; posterior margin often not emarginate................................................................ 11

11. MPP of V7 elongate and inclining dorsally; median posterior margin of T8 narrowly produced and inclining ventrally where it may envelop the MPP ( Ballantyne & Lambkin 2006 Figs 25 , 26 ; 2009 Fig. 60)... Pygoluciola Wittmer Partim Fig. 8 Node 51

- MPP developed, may be elongate (L>W), not inclining strongly dorsally; median posterior margin of T8 not narrowly produced and inclining ventrally........................................................................... 12

12. Aedeagal sheath sternite with narrow elongate anterior portion and expanded posterior area; lateral arms of tergite widely visible at sides of sheath sternite and extending forward ( Fu & Ballantyne 2008 Figs 17, 19 ; Ballantyne 2008 Fig.9 )......... 13

- Aedeagal sheath sternite lacking narrow elongate portion and expanded posterior area; if lateral arms of tergite visible at sides of sternite then close to sternite margins or not developed as above; aedeagus with basal piece not usually bilobed or asymmetrical............................................................................................... 14

13. Basal section of lateral lobes of aedeagus fused for almost all of their length dorsally; inner ventral margins of LL with elongate cylindrical lobes ( Jeng et al. 2003 Figs 27, 28 ; Fu & Ballantyne 2008 Fig. 20 ); apical section of lateral lobes very short................ Luciola dejeani Gemminger , L. dubia Olivier , L. praeusta Kiesenwetter , L. cerata Olivie r, L. aegrota Olivier

- Basal section of lateral lobes of aedeagus not fused; inner ventral margins of LL lacking elongate cylindrical lobes; apical section of LL always longer than ML, often very pale and membranous; aedeagus with strongly asymmetrical bilobed basal piece; MPP scarcely produced; V7 almost entirely occupied by LO; posterolateral corners of T8 either very narrowly pointed, and slightly down turned ( Fu & Ballantyne 2008 Figs 17, 19 ; Ballantyne 2008 Fig. 9 ) or not................................................................................................ Pygoluciola Wittmer Partim Fig. 8 Node 48

14. Median lobe of aedeagus lacking lateral teeth..............................................................15

- Median lobe of aedeagus with lateral teeth; Pacific Ocean species restricted to the Solomon Islands ( Ballantyne & Lambkin 2009 Figs 364–376)............................................ Pygatyphella (Ballantyne) Partim Fig. 9 Node 54

15. Pronotum wider than width across elytral humeri; often with small indentation along posterolateral margin, near corner ( Ballantyne & Lambkin 2009 Fig. 64); anterior hypomeron never strongly flattened, often delimited from posterior hypomeron by sloping area especially if head can be withdrawn into prothoracic cavity; posterior area of hypomeron widely flattened and closely adpressed; elytra never with ridge continuing around apex ( Ballantyne & Lambkin 2009 Figs 69–71); dorsal colouration often with median dark pronotal mark, dark markings on either or both of MS and MN, and at elytral base dark markings either of triangular form adjacent to MS or restricted to basal half of suture only ( Ballantyne & Lambkin 2009 Figs 64, 69–71); apical segment of labial palpi like a wide flattened triangle with 3 or more teeth on inner margin; posterior half of V7 usually arched, sometimes swollen ( Ballantyne & Lambkin 2009 Figs 61, 62) (not arched in Pygat. wisselmerenia); MPP always developed, usually longer than wide, with apex rounded, or squarely or obliquely truncate; LO usually retracted into anterior half or less of V7 (except in Pygat. wisselmerenia where it occupies most of V7), sometimes bipartite; T8 symmetrical, not

Figures

FIGURE 7. Majority rule tree, consensus of 169 MPT of length 5609 from phylogenetic analysis of 143 taxa and 436 morphological characters in Appendix 2. Part 1 Nodes 1 – 34. Node numbers red, bold, italics. Undescribed species in red, bold with identification numbers blue, bold, italics.

FIGURE 8. Majority rule tree, consensus of 169 MPT of length 5609 from phylogenetic analysis of 143 taxa and 436 morphological characters in Appendix 2. Part 2 Nodes 35 – 52. Node numbers red, bold, italics. Undescribed species in red, bold with identification numbers blue, bold, italics.

FIGURE 9. Majority rule tree, consensus of 169 MPT of length 5609 from phylogenetic analysis of 143 taxa and 436 morphological characters in Appendix 2. Part 3 nodes 55 – 76. Node numbers red, bold, italics. Undescribed species in red, bold with identification numbers blue, bold, italics.

FIGURES 16 – 25. Emeia pseudosauteri male 16 – 22, female 23 – 25. 16, 17 aedeagal sheath (right margin of posterior section of sheath sternite arrowed in 16). 18, 19 aedeagus. 20 – 21 entire abdomen. 22 T 8. 23 genitalia, anterior end to top of page. 24, 25 female after prolonged soaking in ethanol, illuminated from below (extent of hind wing arrowed in 24). (16, 18, 20 22, 24 ventral, 17, 19, 21, 25 dorsal). (Scale bar = 1 mm).

FIGURES 26 – 30. Emeia pseudosauteri fifth instar larva SEMs. 26, 27 head. 28 maxillary palp. 29 apex apical maxillary palpomere. 30 body. (27, 28 ventral, 26, 30 dorsal).

FIGURES 32 – 36. Habitus Curtos costipennis. 32, 33 male, 34, 35 female, 36 larva. (32, 34, 36 dorsal, 33, 35 ventral) (Scale bar = 5 mm).

FIGURES 37 – 42. Curtos costipennis larval head SEMs. 37 anterior; 38, 39 entire head; 40 detail anterior margin; 41 outer margin tip of mandible; 42 left antenna (37 dorsal surface uppermost; 38 dorsal; 39, 40 42 ventral).

FIGURES 43 – 49. Characters and states. Adult males and females. 43 Luciola aquatilis Thancharoen: T 8, aedeagal sheath, and sclerites surrounding sheath (T 8 = tergite 8; 1, 2, 3 = sclerites right side, ventral and left side in order; AS = aedeagal sheath; AE = aedeagus). 44 Luciola cingulata Olivier holotype male terminal abdomen (MCSN). 45 Luciola near substriata terminal abdomen (ANIC). 46, 47 undescribed species 8 Figure 2 number 20: 46 male abdomen; 47 female genitalia anterior end to right (styli arrowed). 48, 49 Male of undescribed species New Caledonia 2 Figure 7 number 18: 48, 49 head (ANIC). (43 – 47 ventral; 48 left lateral; 49 dorsal). (Scale bar = 1 mm)

FIGURES 50 – 55. Characters and states. Larvae. 50, 51 Luciola hypocrita Olivier: 50 metatergum and abdominal terga 1 – 7; 51 thoracic terga 1 – 3 (ANIC). 52, 53, 54, 55 undescribed species New Caledonia 1 Figure 7 number 17: 52 abdominal terga 1 – 5 (projections along posterior margin at sides of median line arrowed); 53 thoracic terga 1 – 3; 54 abdominal segments 1 – 5 (posterior projections of laterotergites, laterosternites and median sternal plates arrowed in order from outside in). 55 brush of hairs at apex of tibiotarsi legs 2, 3 (QMBA). (50 – 53 dorsal; 54 ventral; 55 lateral). (Scale bar = 1 mm).

FIGURES 10 – 15. Characters and states. Adult males. 10 – 12 Luciola indica Motschulsky male. 13 – 15 undescribed species 7 (Figure 7 number 4) male. 10, 11 whole body, (character 119); 12 aedeagus (character 271); 13, 14 terminal abdomen; 15 T 8 (10, 13, 15 ventral, 11, 12, 14 dorsal (characters 121 – 123).

FIGURES 1 – 6. Habitus Emeia pseudosauteri. 1, 2 male; 3, 4 female; 5, 6 larva (1, 3, 5 dorsal; 2, 4, 6 ventral) (Scale bar = 5 mm).