Medeopteryx, 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 : 76-84

Medeopteryx gen. nov.

[ Figs 139–184 ]

Type species: Pteroptyx effulgens Ballantyne.

Diagnosis. Medeopteryx gen. nov. is a new genus based mainly on New Guinean bent-winged fireflies formerly assigned to Pteroptyx , and is proposed for New Guinean and Australian species with an entire LO in V7, the posterior margin of which is trisinuate with PLP no longer or wider than the MPP. All but two species have deflexed elytral apices. It is most obviously distinguished from Pteroptyx s. str. in having an entire LO in V7, without a MFC and bulbous paraprocts on the aedeagal sheath.

Male. Pronotum: dorsal surface without irregularities in posterolateral areas and longitudinal groove in lateral areas; punctation dense; anterior margin not explanate; W/L 1.3–1.7; either subparallel-sided, margins straight (A=B=C), or lateral margins diverging with posterior rounded convergence (C>A, B); pronotal 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 usually angulate, 90º to median line, or rounded obtuse; posterolateral corners not usually projecting as far as median posterior margin and separated from it by scarce emarginations.

Hypomera: closed; median area 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 either deflexed or not; deflexed apex rounded, pointed or truncated, truncated apices short (Figs 141,143–146,153); epipleuron and sutural ridge extending beyond mid-point, to deflexed apex; no interstitial lines; elytral carina absent; in horizontal specimen viewed from below epipleuron at elytral base wide, covering humerus, viewed from above epipleuron arises anterior to or level with posterior margin of MS; epipleuron developed as a lateral ridge along most of length until deflexed apex; sutural margins approximate along most of length in closed elytra; lateral margins parallel-sided.

Head: moderately-deeply depressed between eyes; well exposed in front of pronotum, not capable of complete retraction within prothoracic cavity; eyes close to moderately separated beneath at level of posterior margin of mouthpart complex; eyes above labrum 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, usually separated by up to ASW or> ASW in the “ cribellata complex” ( Ballantyne 1987a:131); clypeolabral suture present, flexible, not in front of anterior eye margin when head viewed with labrum horizontal; outer edges of labrum reach to or beyond inner edges of closed mandibles. Mouthparts functional; apical labial palpomere strongly flattened, shaped like narrow triangle (narrowest at base), with inner edge entire; 2–3 X as long as wide. Antennae 11 segmented; length>GHW, usually 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; no femora or tibiae swollen or curved; no basitarsi expanded or excavated.

Abdomen ( Figs 139–146 ): without cuticular remnants in association with aedeagal sheath; V3 and/or V4 often with curved posterior margins and V4 extending anteriorly into emarginated posterior margin of V3 (this feature varies between different specimens in the one species and may be difficult to determine on dried pinned abdomens); posterior margin of V7 trisinuate; LO in V7 entire and occupying most of V7, reaching to sides but not posterior margin; posterior half of V7 not arched or swollen, muscle impressions not visible, anterior margin of LO emarginate only in clipeata sp. nov. ( Figs 147–151 ); LO present in V6, occupying all or almost V6. MPP present, symmetrical, apex emarginate, often shallowly so and emargination may not be visible from beneath; MPP not laterally compressed, short (L=W), not inclined dorsally nor engulfed by T8 apex, without dorsal ridge, median longitudinal trough; MPP longer than or subequal in length to, and wider than or subequal in width to, horizontal PLP. V7 often with median ‘dimple’ (depressed area Fig.143 arrowed) anterior to MPP, in which case the ventral surface of the MPP is elevated ( Ballantyne & McLean 1970 termed this a ‘hump’); V7 without median carina, median longitudinal trough, anteromedian longitudinal depression on face of LO, incurving lobes or pointed projections, or reflexed lobes on dorsal surface; dorsal face of light organ bearing paired cuticular strips running from the sides of the MPP over the surface of the LO and attaching anteriorly to longitudinal abdominal muscles, in antennata , corusca, cribellata, effulgens, elucens, flagrans, fulminea, platygaster , pupilla, similisantennata , similispupillae, sublustris, torricelliensis ( Ballantyne 1987a; 1993), or a wide single strip in clipeata sp. nov. ( Fig. 150 CS). T7 without prolonged anterolateral corners. T8 well sclerotised, symmetrical, W=L, visible posterior area not narrowing abruptly, median posterior margin emarginate, often widely so; usually 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 a well developed median longitudinal trough margined by well defined ridges; anterior end of ridges usually with flanges which may be symmetrical/asymmetrical, narrow or broad, and with rounded or pointed apices (Figs 142,174,175); without lateral depressed troughs, asymmetrical projections, median posterior ridge; concealed anterolateral arms of T8 either as long as visible posterior portion of T8 or slightly shorter, 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 174,176): approx. 3 times as long as wide; without 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 152,171–173, 180–182): L/W approx. 5/1; LL lack lateral appendages; apices of LL not visible from beneath at sides of ML, LL/ML narrow; LL of equal length, very slightly shorter than ML, contiguous or slightly separated longitudinally by most of their length along inner dorsal margins; 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; 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 ( Figs 154–167 ). Macropterous and some species observed in flight. 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 ( Figs 154–159 ). Bursa ( Figs 160–167 ) with paired wide plates; posterior oblique pair broad ventrally narrowing to pointed dorsal ends.

Larva. Not reliably associated.

Etymology. Medeopteryx (feminine noun; Greek mede = genitals, and pteryx = a wing, thus highlighting the shape of V7 and the deflexed elytral apices).

Remarks. Two groups within Pteroptyx Olivier have been recognized ( Ballantyne 1987a) and successive phylogenetic analyses of the Luciolinae support this move to further subdivide the bent-winged fireflies ( Ballantyne & Lambkin 2000, 2006, 2009).

List of species of Medeopteryx gen. nov.

- Species * have an exclusively Australian distribution

- amilae (Satô) comb. nov.

- antennata (Olivier) comb. nov.

- clipeata sp. nov.

- corusca (Ballantyne) comb. nov.

- cribellata (Olivier) comb. nov.

- effulgens (Ballantyne) comb. nov.

- elucens (Ballantyne) comb. nov.

- flagrans (Ballantyne) comb. nov.

- fulminea (Ballantyne) comb. nov.

- hanedai (Ballantyne) comb. nov.

- platygaster (Lea) comb. nov.

- pupilla (Olivier) comb. nov.

- similisantennata (Ballantyne) comb. nov.

- similispupillae sp. nov.

- sublustris (Ballantyne) comb. nov.

- tarsalis (Olivier) comb. nov.

- torricelliensis (Ballantyne) comb. nov.

Key to species of Medeopteryx using males

Modified from Ballantyne (1987a, 2001)

1. Elytral apices not deflexed.............................................................................. 2 Elytral apices deflexed................................................................................. 4

2. T7, 8 dark brown (( Fig. 179 )........................................................... similispupillae sp. nov. T7, 8 pale ( Fig. 170 )....................................................................... pupilla (Olivier)

3. Anterior margin of entire LO in V7 emarginated ( Fig. 147 )........................................ clipeata sp. nov. Anterior margin of LO in V7 not emarginated............................................................... 3

4. Elytral apices pointed, angle between junction of sides B & C <45°; e.g. Fig. 144 ( Ballantyne 1987a Figs 2g , 8h , 9g, 13o ).. 5 Elytral apices rounded or truncate, angle between junction of sides B & C> 45°, usually obtuse ( Ballantyne 1987a Figs 4b, g, 5g , 6i , 7m , 10h, 11h, 12g, n, 13h )......................................................................... 7

5 Head between eyes orange; endemic to New Britain ( Ballantyne 1987a Figs 5 , 8 )................. effulgens (Ballantyne) Head between eyes brown............................................................................. 6

6 Endemic to Cebu island, Philippines; light patterns unknown; flanges not determined...................... amilae (Satô) Endemic to mainland New Guinea; light patterns described in Ballantyne 1987a: 156); flanges wide, apices may be rounded in Kar Kar Island specimens ( Ballantyne 1987a Fig. 9 )......................................... fulminea (Ballantyne)

7 Elytra pale yellow with apex black; flanges broad, apically rounded ( Ballantyne & McLean 1970 Fig. 11 ).. tarsalis (Olivier) Elytra entirely medium to dark brown.................................................................... 8

8 Dimple (depressed area) present on V7; Figs 143, 145 ( Ballantyne & McLean 1970 Figs 7 , 11 ; Ballantyne 1987a Figs 2 , 7–11 ) ................................................................................................... 9 Dimple (depressed area) absent on V7 (e.g. Ballantyne 1987a, Figs 12, 13 )..................................... 12

9 Elytral apices truncate, (short, wider than long)................................................ platygaster (Lea) Elytral apices strongly deflexed (long, at least as long as wide)................................................ 10

10 Elytral apices with A and C subequal, slightly longer than B; dimple shallow, hump only gently elevated; median posterior projection of V7 apically scarcely emarginate; dorsally reflexed margins of Vs 6 and 7 entirely brown; flanges of T8 short, wide, apically rounded; ASD subequal to ASW ( Ballantyne 1987a Fig. 11 )........................ elucens (Ballantyne) Elytral apices with A, B and C subequal, or B and C subequal, slightly longer than A; dimple deep, or at least moderately developed, as is hump; MPP of V7 apically emarginate; dorsally reflexed margins of V6 and 7 white, or dorsally reflexed margin of V6 partly brown, and reflexed margin of V7 white; flanges elongate, narrow, apically acute; ASD> ASW......... 11

11 Elytral apices with A, B, C subequal; dimple deep; ( Fig. 143 ) ( Ballantyne 1987a Fig. 7 ).......... cribellata (Olivier) s. str. Elytral apices with B and C subequal, slightly longer than A; dimple only. moderately depressed ( Fig. 141 ) ( Ballantyne 1987a Fig. 10 )............................................................................ .. corusca (Ballantyne)

12 Pronotum with pronounced ridges and depressed areas in midlateral areas of disc ( Fig. 184 ); flanges short, wide, apically acute ( Ballantyne 1987a Fig. 12 )............................................................. sublustris (Ballantyne) Pronotum smooth over most of disc, without any pronounced ridges in midlateral areas........................... 13

13 Antennal FS 7–9 shorter than remainder.................................................................. 14 Antennal FS 7–9 not shorter than remainder.............................................................. 15

14 MPP of V7 apically emarginate in ventral aspect, and longer than wide across posterior face; apex of elytron moderately acute ( Ballantyne 1987a Figs 3, 4 )..................................................... similisantennata (Ballantyne) MPP of V7 entire in ventral aspect, and about as long as wide across posterior face; apex of elytron broadly rounded ( Ballantyne 1987a Figs 3, 4 ).................................................................... antennata (Olivier)

15 Pronotum with median dark spot on disc ( Ballantyne & McLean 1970 Fig. 15 ).................... hanedai (Ballantyne) Pronotum without dark markings on disc................................................................ 16

16 MPP of V7 apically expanded, bearing 2 short fine, dark, dorsally projecting hooks, and a fine lateral ridge; posterior margin of T8 (excluding median emargination) rounded ( Ballantyne & McLean 1970 Fig. 14 )........... torricelliensis (Ballantyne) MPP of V7 narrow, not expanded apically, not bearing hooks or fine lateral ridges as above; posterior margin of T8 (excluding median emargination) straight ( Ballantyne 1987a Fig. 13 )...................................... flagrans (Ballantyne)

Figures

FIGURE 2. Luciola cerata Olivier flashing patterns taken at Wushikeng, Taichung county, Taiwan, 7.43pm 24.iii.2009 by Jenzon Ho.

FIGURE 6. Strict consensus tree for 61 IP analysis; consensus of 163 MPT of length 2615 from phylogenetic analysis of 61 Indo-Pacific study group taxa. New genera, new species, and still undescribed species in red, bold. Bremer supports above branches.

FIGURE 7. Majority rule tree for female analysis. Tree shows the consensus of 1349 MPTs of length 3892 from the analysis of all characters for the 88 taxa with female traits scored. New genera, new species, and still undescribed species in red. Showing the % of MPT over 50 %, above branches.

FIGURE 8. Majority rule tree for larvae analysis. Tree shows the consensus of 4 MPTs of length 2245 from phylogenetic analysis of all characters for the 42 taxa scored for larval characters. New genera, new species, and still undescribed species in red. Showing the % of MPT over 50 %, above branches.

FIGURES 139–146. Medeopteryx spp. Males (ANIC). 139–142 M. corusca. 143 M. cribellata. 144 M. effulgens. 145 M. fulminea. 146 M. flagrans. 139, 140 abdomen, ventral (139, anterior margin of V4 arrowed) and dorsal; 141 apices of elytra ventral; 142 tergite 8 ventrolateral, flange arrowed; 143–146 terminal abdomen and elytral apices, ventral. These figures share scale lines: 139, 140.

FIGURES 147–153. Medeopteryx clipeata sp. nov. holotype male (BPBM). 147–151 abdomen, ventral (147), dorsal (148), from behind with ventral surface uppermost (149), dorsal V7 (150), left lateral (151); 152 aedeagus left lateral; 153 ventral apex right elytron. Figure legends: CS cuticular strips; MPP median posterior projection of V7; PLP posterolateral projections of V7; SPI spiracle; T8 tergite 8. These figures share scale lines: 147–151.

FIGURES 154–159. Medeopteryx spp. female abdomens ventral (not to scale) (ANIC). 154 M. antennata. 155 M. corusca (mating pair Catherine Mission). 156 M. cribellata (G381 mating pair). 157 M. effulgens (Kah New Ireland). 158 M. flagrans (G535). M. fulminea (Kar Kar Island).

FIGURES 160–167. Female Medeopteryx spp. bursa. 160 Medeopteryx antennata female of mating pair Sek Bridge (ANIC). 161, 162 M. corusca female of mating pair Kondiu (ANIC). 163 M. cribellata Kar Kar Island (ANIC). 164–167 M. similispupillae sp. nov. 164 bursa ventral; 165 reproductive system excluding ovaries lateral; 166 median oviduct plate ventral (anterior end to foot of page); 167 anterior plate lateral. Figure legends: 1 median oviduct plate; 2 posterior bursa plate; 3 anterior bursa plate; 4 spermatophore digesting gland.

FIGURES 168–176. Medeopteryx pupilla male Mt Lamington (AMUS). 168 dorsal; 169, 170 abdomen ventral (169) and dorsal; 171–173 aedeagus left lateral, dorsolateral and ventral; 174, 175 tergite 8 (174 ventral with aedeagal sheath; 175 from right side, dorsal surface uppermost, flanges arrowed); 176 aedeagal sheath dorsal. These figures share scale lines: 171–173.

FIGURES 177–184. Medeopteryx spp. Males. 177–182 M. similispupillae sp. nov. paratypes. 183 M. torricelliensis. 184 M. sublustris, diagrammatic representation of pronotal sculpturing. 177, 183 dorsal; 178, 179 abdomen, ventral (178) and dorsal; 180–182 aedeagus dorsal (180), left lateral (181) and ventral (182). These figures share scale lines: 180–182

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 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 %.