Australoluciola, 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 : 39-44

Australoluciola gen. nov.

[ Figs 9–91 ]

Type species: Australoluciola australis (F.).

Diagnosis. Australoluciola gen. nov. is an Australian and New Guinean genus belonging in a group of seven genera characterized by: pronotal width less than width across elytral humeri, parallel-sided elytra, no MFC, an elongate slender aedeagus with LL largely concealed behind the ML when viewed from beneath; aedeagal sheath elongate slender, widest across middle, without bulbous paraprocts, and with both sides of posterior half of sheath sternite tapering evenly towards a narrow entire apex. Males distinguished from Colophotia in having no median carina on V7, expanded and oblique PLP and bipartite LOs in V7; from Pteroptyx in having no MFC, deflexed elytral apices, bulbous aedeagal sheath paraprocts and bipartite LOs in V7; from Pyrophanes (which has a MFC) and Poluninius gen. nov. (which has no MFC) in lacking both incurving lobes along V7 and bipartite LO; from Trisinuata gen. nov. by the entire LOs in V7 (those of Trisinuata gen. nov. are bipartite); from most Medeopteryx gen. nov. in lacking deflexed elytral apices. It differs from Luciola indica in lacking the bulbous median lobe and paraprocts on the aedeagal sheath.

All but three species have orange pronotum and dark brown elytra similar to many species of Medeopteryx gen. nov. Aus. aspera has median brown pronotal markings, and in Aus. flavicollis the elytral base and/or margins may be pale. Aus. japenensis sp. nov. alone has pale yellowish brown dorsal colour and dark elytral apices, characteristic of many SE Asian fireflies ( McDermott 1966). V7 may be trisinuate or with an MPP only, and PLP rounded, not produced; T8 has a well defined ventral groove margined by ridges and with short, wide and apically rounded flanges in Aus. aspera and Aus. pharusaurea sp. nov. only. Females macropterous and capable of flight. Larvae terrestrial, without laterally explanate tergal margins; laterotergites visible from above.

Male. Pronotum ( Figs 12 , 19, 25 , 58 ): dorsal surface without irregularities in posterolateral areas and longitudinal groove in lateral areas; punctation dense; anterior margin not explanate; either subparallel-sided, margins straight (A=B=C; e.g. Fig. 12 ), or lateral margins diverging posteriorly along their length (C>A, B), or lateral margins converging posteriorly (C<A, B); width <humeral width; anterolateral corners rounded obtuse or angulate; lateral margins without indentation at mid-point, and sinuousity in either horizontal or vertical plane; without indentation in lateral margin near posterolateral corner, and irregularities at corner; posterolateral corners usually angulate, rounded obtuse in Aus. australis ; posterolateral corners not usually projecting as far as median posterior margin; separated from it by scarce emarginations.

Hypomera: closed; median area of hypomeron not elevated in vertical direction; median area more widely flattened than elsewhere except in fuscamagna sp. nov. where anterior and posterior areas very narrowly flat; pronotal width/ GHW 1.2.

Elytron: punctation dense, not linear, not as large as that of pronotum, nor widely and evenly spaced; apices not deflexed; epipleuron and sutural ridge extend beyond mid-point, almost to apex but not extending 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 except in some fuscaparva sp. nov. where the humerus is narrowly visible; viewed from above anterior margin of 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, separated by <ASW or ASW (except for some flavicollis where ASD>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. Mouthparts: functional; apical labial palpomere strongly flattened, shaped like narrow triangle (narrowest at base and L 2–3 X W), with inner edge entire (e.g. Ballantyne 1988 Figs 10, 11 ), and at least half as long as apical maxillary palpomere. Antennae 11 segmented; length>GHW up to twice GHW; no segments flattened, shortened, or expanded; pedicel not produced; FS1 not shorter than pedicel.

Legs: with inner tarsal claw not split; without MFC; femora 3 swollen and curved and tibiae 3 curved in two species; no basitarsi expanded or excavated.

Abdomen (Figs 10,11,21,29,35,36,57,61,62,67,68,73,75,76,88): 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 single male of Aus. aspera , where V3 is strongly recurved ( Fig. 21 ); LO in V7 entire, either occupying most of V7, and reaching to sides but not posterior margin (Figs 11,20,21,28,35,61,68,73,75,87), or not reaching sides or posterior margin and occupying about half or less of V7 (e.g. Fig. 48 ); reaching into PLP where these are developed; posterior half of V7 not arched or swollen, muscle impressions not visible in this area; neither anterior nor posterior margin of LO emarginate; LO present in V6, occupying almost all V6. MPP present, symmetrical, apex rounded or truncate, either entire or shallowly emarginate, not laterally compressed, short or slightly longer than wide (L<W or L>W), not inclined dorsally nor engulfed by T8 apex, without dorsal ridge, median longitudinal trough. V7 without median carina, median longitudinal trough, anteromedian depression on face of LO, incurving lobes or pointed projections, median ‘dimple’, or reflexed lobes; posterior margin of V7 either trisinuate or with posterolateral corners rounded; if trisinuate then MPP longer than, and as wide as, PLP; if PLP not developed then posterolateral corners rounded and MPP well developed; PLP moderately produced, as wide as MPP, as long as wide or wider than long. T7 without prolonged anterolateral corners. T8: (Figs 29,30,36,62,76,77,88) well sclerotised, symmetrical, W=L, visible posterior area not narrowing abruptly, median posterior margin shallowly and narrowly emarginate; T8 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, margined by well defined symmetrical ridges; anterior end of ridges either not produced, with rounded or angulate outline; or with short wide apically acute flanges in aspera and pharusaurea ; without lateral depressed troughs, asymmetrical projections, median posterior ridge; concealed anterolateral arms of T8 as long as, or slightly shorter than 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 14,30,42,43,52,53,81,82,89) approx. 3 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 T 9, anterior margin without transverse band.

Aedeagus: ( Figs 15 –17,22–24,31–33,39–41,49–51,63–65,69,70,78–80) L/W2.9–6.0; LL lack lateral appendages; apices of LL not visible from beneath at sides of ML, LL/ML narrow; LL of equal length, slightly shorter than ML, contiguous or closely approaching along inner dorsal margins except in orapallida where inner margins diverge and apices are very narrow; LL 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; 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; bearing transverse ridge preapically on dorsal side; BP not strongly sclerotised, not hooded, not strongly emarginated along anterior margin.

Female ( Fig. 18 ). Macropterous and observed in flight in Aus. australis ( Ballantyne 1988 Fig. 12 ). 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 (Figs 34,54,55,91) consisting of 2 wide paired plates in Aus. australis, Aus. flavicollis and Aus. pharusaurea sp. nov.

Larva. Reliably associated for Aus. australis only; terrestrial; elongate, slender, spindle shaped ( Ballantyne & Buck 1979 Figs 32, 33 , 37 ; Ballantyne 1988) of the form of Pteroptyx valida and P. maipo larvae ( Ballantyne et al. 2011), without laterally explanate tergal margins and with laterotergites usually visible at sides; posterolateral corners of terga 1–8 rounded entire, of tergum 12 produced narrowly; median posterior margins of terga 1–11 with rounded projections beside mid line; without brush of hairs from apex of tibiotarsus; mandibles without inner teeth; antennal segment 3 short, sense cone adjacent to segment 3 short, wide.

Etymology. Australoluciola (feminine) is a compound noun indicating its origins (from the south) and previous affinities (seven of the species were described originally within the genus Luciola ).

Remarks. Australoluciola gen. nov. includes eight New Guinean and four Australian species, one of which, Aus. australis (F.), is found in mangroves along the E coast of Queensland, while Aus. orapallida (Ballantyne) is known from one locality in mangroves in Cape York Peninsula.

The other Australian species, Aus. flavicollis and Aus. nigra , have an extensive distribution along the eastern coast of Queensland occurring in open sclerophyll forest and rain forest remnants especially around Brisbane ( Ballantyne & Lambkin 2000). Three of the New Guinean species are distinguished by light patterns while nothing is known of their wider distribution.

List of species of Australoluciola gen. nov.

- anthracina (Olivier) comb. nov.

- (Olivier) comb. nov.

- australis (F.) comb. nov.

- baduria sp. nov.

- flavicollis (MacLeay) comb. nov.

- foveicollis (Olivier) comb. nov.

- fuscamagna sp. nov.

- fuscaparva sp. nov.

- japenensis sp. nov.

- (Olivier) comb. nov.

- orapallida (Ballantyne) comb. nov.

- pharusaurea sp. nov.

Key to species of Australoluciola gen. nov. using males

1. V7 trisinuate along its posterior margin (Figs 10,11,21,35,57,61,67,68,73,75)...................................... 2 V7 not trisinuate along its posterior margin (Figs 28,48,87); posterolateral corners rounded, not produced; median posterior projection always developed sometimes longer than wide, may be apically emarginated............................. 11

2. Hind femora swollen, hind tibiae curved (e.g. Fig. 37 )......................................................... 3 Hind femora not swollen, hind tibiae not curved............................................................. 4

3. Elytra dark brown......................................................................... baduria sp. nov. Elytra orange yellow with small dark area at apex............................................. japenensis sp. nov.

4. Pronotum all dark, or with dark markings around margins, or in centre........................................... 5 Pronotum pale coloured with no dark markings.............................................................. 8

5. Pronotum entirely dark marked; rest of dorsal body and all of ventral surface except for LOs dark brown (Figs 66,67,71,72,74) .................................................................................................... 6 Pronotum with dark markings around margins or in centre; ventral body may have paler markings..................... 7

6. Medium sized (6.2–6.6 mm long)........................................................ fuscamagna sp. nov. Small (3.5–4.0 mm long)................................................................ fuscaparva sp. nov.

7. Pronotum orange with orange margins and extensive median dark markings ( Fig. 25 ); elytra dark brown; ventral surface of body with pale areas including Los ( Fig. 20 ).................................................... aspera (Olivier) Pronotum orange with dark markings around margins; ventral body (apart from LOs) dark brown... anthracina (Olivier) var.

8. Ventral aspect of body entirely dark brown except: posterior margin of abdominal V5 may be pale or apices of femora 1 may be paler; head between eyes dark brown; T7, 8 dark brown (Figs 9,10,75,75)...................................... 9 Ventral surface of body always with some pale markings i.e. parts of head, thorax and/or portions of legs and/or basal segments of abdomen pale, head between eyes yellow or brown; T7, 8 pale (Figs 57,83–86)................................. 10

9. Large 9.9–10 mm long; known only from 3 males from Sea Falls Torricelli Mountains ( Figs 75–82 )........ maxima sp. nov. Smaller (<8 mm); know from SE New Guinea............................................ anthracina (Olivier) var.

10. New Britain; head between eyes yellow; elytral punctures small uniform ( Figs 83–91 ).............. pharusaurea sp. nov. Mainland New Guinea; head between eyes dark brown to black; punctures along some areas of elytra large and irregular (figs 56–65) foveicollis (Olivier)

11. Found in mangroves along east coast of Queensland as far south as Bundaberg; ventral body yellow except for white LOs; MPP short; head between eyes brown ( Figs 26–34 ).................................................. australis (F.) Open forest dweller; ventral body with extensive dark markings; MPP longer than wide, may be apically emarginate...... 12

12. Elytra dark brown with no pale margins........................................................ nigra (Olivier) Elytra dark brown with paler margins..................................................................... 13

13. Elytra brown at bases and apices, and lateral margins widely pale; most of ventral... surface of body pale; aedeagus with LL very narrow and widely separated dorsally ( Ballantyne & Lambkin 2000 Figs 19–22 )............ orapallida (Ballantyne) Elytra brown always with some orange markings, either across base, and/or sutural ridge, and/or lateral margin; most of ventral surface of body very dark brown; aedeagus with LL closely approaching dorsally ( Fig. 45 )........ flavicollis (MacLeay)

Table 8 lists species described originally as Luciola spp. , having orange pronota and dark brown, nonmargined, elytra, consistent with seven species assigned here to Australoluciola gen nov. Other genera/species with this colour pattern include Lloydiella spp. Ballantyne ( Ballantyne & Lambkin 2009), Atyphella immaculata Ballantyne ( Ballantyne & Lambkin 2000) , species of Medeopteryx gen nov., Pacifica salomonis (Olivier) comb. nov., certain Pyrophanes species , and some Curtos ( Jeng et al. 1998) .

pronota and dark brown/ black elytra.

Figures

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.

FIGURES 18–25. Australoluciola aspera, type female (18), Wewak male NHML 19–25. 18, 19 dorsal; 20, 21 ventral (21 abdomen only); 22–24 aedeagus dorsal (22), right lateral (23) and ventral (lateral projections of LL arrowed in 24); 25 pronotum dorsal. These figures share scale lines: 19, 20; 22–24.

FIGURES 26–34. Australoluciola australis, male (ANIC). 26, 27 dorsal and ventral (27 terminal abdomen removed); 28–30 abdomen, 28 V5–7 ventral; 28 segments 6–8 dorsal; 30 tergite 8 with aedeagus and sheath in place, ventral; 31–33 aedeagus left lateral (31), ventral (32) and dorsal; 34 reproductive system, Ross River female, from side, anterior end to right of page. Figure legends: 1 median oviduct plate; 2 posterior and 3 anterior bursa plates. These figures share scale lines: 28, 29; 31–33.

FIGURES 35–44. Australoluciola baduria paratype male 35 -37. Aus. japenensis paratype male 39-44, paratype female 38 (BPBM). 35, 36, 38, 44 abdomen dorsal (36), ventral 35, 38; 44 detail of posterior margin of terminal abdomen; 37 ventral right hind leg (coxa and trochanter not visible); 39-41 aedeagus ventral (39), left lateral (40) and dorsal (41) (anterior margin of ML arrowed); 42, 43 aedeagal sheath ventral (42) and dorsal. Figure legends: V valvifers of ovipositor; 1, 2 and 3 median oviduct plate, posterior and anterior bursa plates; These figures share scale lines: 35, 36; 39–41; 42, 43.

FIGURES 45–55. Australoluciola spp. 45, 54, 55 Australoluciola flavicollis, male 45, female 54, 55 (ANIC). 46 Luciola coarcticollis type male (MNHN). 47 Luciola gestroi type male (MNHN). 45–47 dorsal; 48 ventral V4–7 and T8; 49–51 aedeagus left lateral (49), ventral (50) and dorsal; 52, 53 aedeagal sheath with aedeagus ventral (52) and dorsal; 54, 55 female bursa from side (54) and beneath (55). Figure legends: 2 posterior plates; 3 anterior plates. These figures share scale lines:49– 51; 52, 53.

FIGURES 56–65. Australoluciola foveicollis male (56–60 Haveri, MNHN; 61–65 Mt Lamington SAMA). 56 dorsal; 57 ventral; 58 detail pronotum; 59, 60 lateral margins of elytron, posterior (59) and anterior half; 61, 62 abdomen ventral (61) and dorsal; 63–65 aedeagus right lateral (63), ventral (64) and dorsal. These figures share scale lines: 59, 60; 61, 62; 63–65.

FIGURES 66–74. Australoluciola spp. 66–71 Aus. fuscamagna sp. nov. holotype male (ANIC). 72–74 Aus. fuscaparva sp. nov. holotype male (ANIC). 66, 72 dorsal, 67 ventral; 68, 73 terminal abdomen with LO; 69, 70 aedeagus ventral (69) and left lateral; 71, 74 pronota, dorsal. These figures share scale lines 69, 70.

FIGURES 75–82. Australoluciola maxima sp. nov. holotype male (ANIC). 75–77 abdomen, ventral (75), dorsal (76), tergites 7, 8, dorsal (77); 78–80 aedeagus ventral (78), left lateral (79) and dorsal; 81, 82 aedeagal sheath ventral (81) and dorsal. These figures share scale lines: 78–80.

FIGURES 83–91. Australoluciola pharusaurea sp. nov. (83–89 holotype male ANIC; 90 paratype male G648 ANIC; 91 paratype Kerevat female ANIC). 83 dorsal; 84 pronotum dorsal; 85 anterior head; 86 left lateral apical palpomeres maxilla and labium, apical labial palpomere arrowed; 87, 88 abdomen ventral (87) and dorsal; 89 aedeagal sheath dorsal; 90 Tergite 7, 8 ventral; 91 bursa lateral. Figure legends: 2 posterior plate, 3 anterior plate.