Pyrophanes semilimbata (Olivier) Ballantyne & Lambkin, 2013

Ballantyne, Lesley A. & Lambkin, Christine L., 2013, Systematics and Phylogenetics of Indo-Pacific Luciolinae Fireflies (Coleoptera: Lampyridae) and the Description of new Genera, Zootaxa 3653 (1), pp. 1-162 : 111-116

Pyrophanes semilimbata (Olivier) comb. nov.

[ Figs 223 , 224,227]

Luciola semilimbata Olivier, 1883:76 ; 1913b: 417. Gorham, 1903: 326. Thancharoen et al. 2007:61.

Luciola venusta Olivier, 1883:76 ; 1902: 85 (synonymy). Thancharoen et al. 2007:61.

Luciola (Luciola) semilimbata Olivier. McDermott, 1966: 113 .

Types. Luciola semilimbata Olivier. Female. 'Indes Orientales' labelled 1. Female symbol; 2. Handwritten Luciola semilimbata Chr. Ind or; 3. Typed label Coll Chevrolet; 4. Handwritten Luciola semilimbata with Ern Oliv typed in corner; 5. Typed label Specimen typicum originale auctoris Ern Olivier (MNHN). Fig. 227 .

Luciola venusta Olivier. Male. Java ouest labelled 1. Typed label Specimen typicum originale auctoris Ern Olivier; 2. Handwritten Luciola venusta with Ern Oliv typed in corner (MNHN). Figs 223, 224 .

Diagnosis. The only known Pyrophanes species with hind femora not swollen and hind tibiae not curved; very similar in dorsal colouration to Pyro. appendiculata colouration differing in having only the lateral elytral margins pale ( appendiculata has both lateral and sutural margins pale, often widely so).

Remarks. Olivier (1883) described L. semilimbata from a female with elytral punctation in lines. Immediately below he described a male, from 'Java ouest' as L. venusta , with the last 3 abdominal ventrites white, and V7 trilobed. Olivier subsequently (1902) synonymised the two species. Both sexes conformed in the narrowly pale colour of the sutural ridge and lateral elytral margins. Olivier did not describe the elytral punctures in lines in the male. Gorham (1903) was unsure of the identification of specimens he described as venusta . Olivier (1913b) probably incorrectly recorded semilimbata (but not the sex of his specimens) from New Guinea, at Erima and Stephansort. Thancharoen et al. (2007 Fig. 5 ) distinguished 11 species, including semilimbata , where the original description indicated elytral punctures in lines.

Neither male nor female in MNHN has elytral punctation in lines. Males are very similar in dorsal colouration to Pyrophanes appendiculata and it is possible that Olivier misidentified some of his New Guinean material.

Trisinuata gen. nov.

[ Figs 228–265 ]

Type species: Trisinuata caudabifurca sp. nov.

Diagnosis. Trisinuata gen. nov. is a New Guinean genus that belongs in a group of 7 genera characterized by: an elongate slender aedeagus with LL concealed behind the ML when viewed from beneath, pronotal width less than width across the elytral humeri, parallel-sided elytra, aedeagal sheath elongate slender, widest across the middle, with posterior half of sternite tapering evenly towards a narrow entire apex; bulbous paraprocts absent. Males have bipartite LOs in V7, expanded horizontal PLP, trisinuate posterior margin of V7, and several species have deflexed elytral apices. Males are distinguished from Colophotia in lacking a median carina on V7, and oblique PLP; from Pteroptyx in lacking bulbous paraprocts and an MFC; from Pyrophanes and Poluninius gen. nov. in lacking incurving lobes along the posterior margin of V7; from Australoluciola gen. nov. which has entire LOs in V7 by the bipartite LOs; from Medeopteryx gen. nov. in having bipartite LOs in V7 with expanded PLP. It differs from Luciola indica in lacking the bulbous median lobe and paraprocts on the aedeagal sheath. Three species are dorsally brown; the remaining species have orange yellow pronota with dark brown elytra. Females associated by label data are macropterous. Larvae are not associated.

Male. Pronotum: dorsal surface without irregularities in posterolateral areas and longitudinal groove in lateral areas; punctation dense; anterior margin not explanate; subparallel-sided, with margins straight (A=B=C), except in papuana and similispapuae where C>A and B, with lateral margins diverging posteriorly along their length in papuana , and dimidiata where B>A, C ( Figs 239, 242 , 251 ); width <humeral width; anterolateral corners rounded obtuse; lateral margins without indentation at mid-point, or sinuousity in either horizontal or vertical plane; without indentation in lateral margin near posterolateral corner, and irregularities at corner; posterolateral corners angulate, inclined either at 90° or obliquely to the median line in papuae ; posterolateral corners either not projecting, or projecting as far as median posterior margin and separated from it by scarce emarginations.

Hypomera: closed; median area of hypomeron not elevated in vertical direction; median area more widely flattened than elsewhere; pronotal width/ GHW 1.2.

Elytron: punctation dense, not linear, not as large as that of pronotum, nor widely and evenly spaced; apices deflexed in four species, weakly so in minor ; non deflexed apices rounded; epipleuron and sutural ridge extend beyond mid-point, almost to apex but not as ridge around apex, neither thickened in apical half; no interstitial lines; elytral carina absent; in horizontal specimen viewed from below epipleuron at elytral base wide, covering humerus; viewed from above the anterior margin of the epipleuron arises level with or anterior to posterior margin of MS; epipleuron developed as a lateral ridge along most of length; sutural margins approximate along most of length in closed elytra; lateral margins parallel-sided.

Head: moderately depressed between eyes; well exposed in front of pronotum, not capable of complete retraction within prothoracic cavity; eyes moderately separated beneath at level of posterior margin of mouthpart complex; eyes above labrum close to moderately separated; frons-vertex junction rounded, without median elevation; posterolateral eye excavation not strongly developed, not visible in resting head position; antennal sockets on head between eyes, not contiguous, ASD <or subequal to ASW; clypeolabral suture present, flexible, not in front of anterior eye margin when head viewed with labrum horizontal; outer edges of labrum reach inner edges of closed mandibles or sometimes beyond. Mouthparts functional; apical labial palpomere strongly flattened, shaped like narrow triangle (narrowest at base and L 2–3 X W) and at least half as long as apical maxillary palpomere except in Tri. dimidiata sp. nov. where it is a short broad triangle (W/L=5/9), and 0.4 as long as apical maxillary palpomere, with inner edge entire. Antennae 11 segmented; length>GHW to twice GHW; no segments flattened, shortened, or expanded; pedicel not produced; FS1 not shorter than pedicel.

Legs: inner tarsal claw not split; without MFC; no femora or tibiae swollen or curved; no basitarsi expanded or excavated.

Abdomen ( Figs 228–234 , 240, 241, 243–250 , 253, 254 , 259–262 ): without cuticular remnants in association with aedeagal sheath; no ventrites with curved posterior margins nor extending anteriorly into emarginated posterior margin of anterior segment except in some Tri. caudabifurca sp. nov. where V3 is recurved; LO in V7 bipartite, inner margins not contiguous, reaching into PLP; posterior half of V7 not arched or swollen, muscle impressions visible between LO halves; LO present in V6, occupying almost all V6. MPP present, symmetrical, apex rounded or squarely truncate, either entire or shallowly emarginate, not laterally compressed, as long as or longer than width (L=W or L>W), not inclined dorsally nor engulfed by T8 apex, without dorsal ridge, median longitudinal trough; shorter than or subequal in length to PLP, MPP narrower than PLP except in Tri. dimidiata sp. nov. where they are subequal in width ( Fig. 245 ); emarginations between PLP and MPP obliquely truncate (deeper at inner corners nearest MPP) in Tri. dimidiata sp. nov. ( Fig. 245 ); V7 without median carina, median longitudinal trough, incurving lobes or pointed projections, median ‘dimple’, or reflexed lobes; posterior margin of V7 trisinuate with PLP moderately to considerably produced, as wide as or wider than MPP, longer than wide except in Tri. papuana where L=W ( Figs 253, 254 ). T7 without prolonged anterolateral corners. T8: sclerotised, symmetrical, W=L, visible posterior area not narrowing abruptly, median posterior margin shallowly and narrowly emarginate; widest across middle with lateral margins tapering evenly in both an anterior and posterior direction; without prolonged posterolateral corners, median posterior projections, not inclined ventrally nor engulfing posterior margin of V7 nor MPP, not extending conspicuously beyond posterior margin of V7. T8 ventral surface with well developed median longitudinal trough (except in Tri. similispapuae where it is quite short), margined by well defined symmetrical ridges; anterior end of ridges with flanges which are short to long, narrow to wide, and apically rounded or acute ( Figs 232 , 250 , 255 ); flanges symmetrical except in Tri. apicula sp. nov., papuae and similispapuae ( Fig. 261 ); without lateral depressed troughs, median posterior ridge; concealed anterolateral arms of T8 as long as visible posterior portion of T8, not laterally emarginated before their origins, not expanded dorsoventrally, expanded only in horizontal plane; without bifurcation of inner margin and ventrally directed pieces; lateral margins of T8 not enfolding sides of V7.

Aedeagal sheath ( Figs 238 , 257 ): approx. 3–3.5 times as long as wide; without bulbous paraprocts; symmetrical in posterior area where sheath sternite tapers evenly to a narrow rounded apex; anterior half of sternite relatively narrow, apically rounded; tergite without lateral arms extending anteriorly at sides of sheath sternite; tergite without projecting pieces along posterior margin of T9, anterior margin without transverse band.

Aedeagus ( Figs 237 , 258 , 259, 264 ): L/W 5/1; LL lack lateral appendages; LL apices not visible from beneath at sides of ML, LL/ML narrow; LL of equal length, slightly shorter than ML, contiguous along inner dorsal margins; separated longitudinally by most of their length; LL base width not = LL apex width which is narrower than that of ML; LL apices not expanded in horizontal plane; dorsal base of LL symmetrical, not excavated, may be slightly produced or not; LL without lateral hairy appendages along their outer ventral margins, not produced preapically nor narrowly on inner apical margin, apices of LL not inturned, nor out-turned; without projection on left LL; inner margins without slender leaf-like projection; ML symmetrical, without paired lateral teeth and tooth to left side, not strongly arched, apex not shaped like arrowhead, not bulbous, not inclined ventrally; BP not strongly sclerotised, not hooded, not strongly emarginated along anterior margin.

Female. Macropterous; associated by label data only for two species. Pronotum without irregularities in posterolateral areas; punctation moderate to dense; pronotal width less than humeral width; without indentation of lateral margin, irregularities at posterolateral corner; outline similar to that of male. Elytral punctation not as large as that of pronotum, nor evenly spaced; no interstitial lines; elytral carina absent. No legs or parts thereof swollen and /or curved. LO in V6 only, without any elevations or depressions or ridges on V7; median posterior margin of V7 widely emarginate, median area not broadly rounded; median posterior margin of V8 entire. Bursa plates consisting of 2 wide pairs in Tri. papuana and Tri. similispapuae ( Fig. 265 ).

Larva. Unknown.

Etymology. Trisinuata (feminine) is a noun latinised from the English trisinuate, describing the shape of the posterior margin of V7 where the expanded and lengthened PLP enhance the appearance of the trilobed posterior margin.

List of species of Trisinuata gen. nov.

- T. apicula sp. nov.

- T. caudabifurca sp. nov.

- T. dimidiata sp. nov.

- T. microthorax (Olivier) comb. nov.

- T. minor (Ballantyne) comb. nov.

- T. papuae (McDermott) comb. nov.

- T. papuana (Olivier) comb. nov.

- T. similispapuae (Ballantyne) comb. nov.

Key to species of Trisinuata gen. nov. using males

1. Elytral apices deflexed.................................................................................. 2 Elytral apices not deflexed.............................................................................. 6

2. Deflexed elytral apex scarcely wider than epipleuron ( Ballantyne & McLean 1970 Fig. 16m )..................................................................................................... minor (Ballantyne) comb. nov. Deflexed elytral apices well defined....................................................................... 3

3. Pronotum orange, lateral margin may be finely dark marked.................................................... 4 Pronotum brown...................................................................................... 5

4. Pronotum orange, without dark marginal markings; PLP apically emarginate ( Ballantyne & McLean 1970 Fig. 5 a–d )................................................................................ microthorax (Olivier) comb. nov. Pronotum orange with fine dark margins; PLP not apically emarginate................................ apicula sp. nov.

5. MPP of V7 elongate, slender, subequal in length and width to PLP or longer than PLP ( Figs 259, 260 ); aedeagus 2.5 times as long as broad; LL of aedeagus closely approximate dorsally; flanges of T8 asymmetrical ( Fig. 261 ; Ballantyne & McLean 1970 Fig. 17 ).......................................................... similispapuae (Ballantyne) comb. nov. MPP of V7 narrower and shorter than PLP; aedeagus 4.5 times as long as broad; LL of aedeagus approximate only at their apices ( Ballantyne & McLean 1970 Fig. 10 ); flanges symmetrical....................... papuae (McDermott) comb. nov.

6. Dorsal surface of body brown to dark brown ( Figs 251, 252 )............................ papuana (Olivier) comb. nov. Pronotum yellow, elytra dark brown....................................................................... 7

7. PLP no longer than MPP; ( Figs 243, 245 ).................................................... dimidiata sp. nov. PLP longer than MPP, considerably produced, longer than wide, and may incline ventrally ( Figs 228–231 , 240, 241 ).......................................................................................... caudabifurca sp. nov.

Figures

FIGURES 221–227. Pyrophanes spp. 221, 222 P. appendiculata (slides, USNM). 223, 224 Luciola venusta type male (MNHN). 225, 226 P. quadrimaculata type female (MNHN). 227 Luciola semilimbata type female (MNHN). 221 male, 222 female abdomens; 223, 225 dorsal; 224 ventral; 226 type labels. Figure legends: 1 anterior projection of aedeagal sheath sternite; 2 basal piece aedeagus; 3 lateral area anterior prolongation of T8; 4 posterior area of aedeagal sheath in region of paraprocts; 5 ventrolateral surface of T8 in spined area; 6 posterolateral area of T8; 7 anterior plates bursa; 8 posterior plates bursa; 9 median oviduct plate.

FIGURES 3–5. Majority rule tree for the complete analysis of the Luciolinae. Analysis consisted of consensus of 169 MPTs with a length of 5609 from a data set of 143 taxa and 436 morphological characters in (see Table 3). Figure 3, part 1: nodes 1– 34; 4, part 2 Nodes 35–52; 5, part 3 nodes 55–76. Nodes (red). New genera, new species, and still undescribed species in red and bold, with identification numbers blue. Black values above branches show the % of MPTs over 50 %.

FIGURES 228–238. Trisinuata caudabifurca sp. nov. holotype and paratype males (234, 237) (ANIC). 228–231 abdomen, ventral (228 arrow indicates muscles in area of MPP), dorsal (229), end on ventral surface uppermost (230), left lateral (231); 232 tergite 8 ventral (anterior arrows indicate flanges, posterior arows lateral ridges); 233, 234 apex of MPP; 235 semidiagrammatic left lateral view showing head, pronotum and elytron; 236 head and pronotum dorsal; 237 semidiagrammatic aedeagus left lateral (arrow to right indicates LL); 238 aedeagal sheath dorsal. Figure legends: BP basal piece; LO light organ; M muscles; ML median lobe; PLP posterolateral processes V7; T8 tergite 8. These figures share scale lines: 228–234.

FIGURES 239–250. Trisinuata spp. 239–241 Trisinuata caudabifurca sp. nov. (239 holotype male, 240, 241 paratype male Banz). 242–250 Trisinuata dimidiata sp nov. (242, 245, 246, 248, 249, 250 holotype male ANIC; 243, 244, 247 paratype male). 239, 242 body dorsal; 240, 243, 245 abdomen ventral; 241, 244, 246, 247 abdomen ddorsal (247 detali terminal two segments only); 248 V7 posterior margin from above; 249 apex of MPP of V7 posterior view dorsal surface uppermost; 250 T8 ventral (upper arrow flange; lower arrow ridge). Figure legends: M muscle; MPP median posteriorprojection of V7; PLP posterolateral projection of V7; T8 tergite 8. These figures share scale lines: 240, 241; 243, 244; 248, 249.

FIGURES 251–258. Trisinuata papuana Males (Komba SAMA) and type female (252). 251, 252 dorsal; 253, 254 abdomen, dorsal (253) and ventral; 255 T8 ventral; 256 aedeagal sheath dorsal (anterior margin of sheath sternite missing); 257, 258 aedeagus left lateral (257) and dorsal. These figures share scale lines: 253, 254; 257, 258.

FIGURES 259–265. Trisinuata similispapuae males and female (259–261, 263, 265 Baiyer River mating pair; 262 Mt Lamington males). 259, 263 abdomen ventral; 261 T8 ventral (anterior prolongation on left side missing; flanges arowed); 262 whole body, dorsal (262) and ventral; 264 aedeagus left lateral with extruded spermatophore; 265 bursa lateral. Figure legends: 1 median oviduct plate; 2 posterior bursa plates; 3 anterior bursa plates. These figures share scale lines: 259, 260.

FIGURES 9–17. Australoluciola anthracina type male, Cyclops (MNHN). 9, 10 dorsal (9) and ventral; 11 V6, 7 ventral; 12 pronotum dorsal; 13 type labels; 14 aedeagal sheath ventral; 15-17 aedeagus, dorsal (15), ventral (16) and lateral. These figures share scale lines: 15–17.