Pteroptyx

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 : 101-104

Pteroptyx s. str. Olivier

Pteroptyx Olivier, 1902:72 ; 1907:55; 1909a:319; 1909b:lxxxii; 1910b:47; 1911b:16; 1913a:58. Olivier & Pic, 1909:139. McDermott, 1959:10 (partim); 1964:46 (partim); 1966:117 (partim). Ballantyne & McLean, 1970:223 (partim). Ballantyne, 1987a:117; 1987b:171; 2001:51. Ballantyne et al. 2011:8 (partim).

Nec Ballantyne in Calder 1998:180.

Type species. Olivier (1902) described Pteroptyx from 2 species viz. Luciola testacea Motsch. and Luciola malaccae Gorham. Lucas (1920) designated Luciola testacea as the type species of Pteroptyx (see Ballantyne in Calder 1998:179). Ballantyne and McLean (1970) were of the opinion, from an examination of a black and white photograph of the type species, that Luciola testacea lacked deflexed elytral apices and its taxonomic position is uncertain, and considered that Luciola malaccae should be advanced to the International Commission for Zoological Nomenclature for consideration as the type species of Pteroptyx s. str. This matter has been held in abeyance until a phylogenetic analysis established the position of all species with deflexed elytral apices.

Diagnosis. Pteroptyx s. str. is an oriental genus that belongs in a group of 7 genera with males characterized by: an elongate slender aedeagus with apices of LL concealed behind the median lobe when viewed from beneath, pronotal width less than width across elytral humeri, parallel-sided elytra, aedeagal sheath elongate slender, widest across middle, with posterior half of sternite not emarginate on either side, and tapering evenly towards a narrow entire apex. Males of Pteroptyx s. str. have a MFC, deflexed elytral apices, aedeagal sheath with bulbous paraprocts and bipartite LOs in V7. Males are distinguished from Australoluciola which has entire LOs in V7 and lacks the MFC; from Colophotia in lacking a median carina on V7, expanded and oblique PLP and elongate slender aedeagal sheath; from Pyrophanes and Poluninius gen. nov. which lack deflexed elytral apices; from Trisinuata gen. nov. and Medeopteryx gen. n which lack the MFC and bulbous aedeagal sheath paraprocts. It differs from Luciola indica in lacking the bulbous median lobe and non deflexed elytral apices. Two presently undescribed species were identified in this analysis ( Fig. 3 Node 15 blue numbers 5, 6) (one without deflexed elytral apices; Ballantyne in prep.).

List of species of Pteroptyx s. str.

- asymmetria Ballantyne

- bearni Olivier

- decolor Olivier

- gelasina Ballantyne

- maipo Ballantyne

- malaccae Olivier

- macdermotti McLean

- masatakai Kawashima

- sulawesiensis Kawashima

- tener Olivier

- truncata Ballantyne

- valida Olivier

Key to species of Pteroptyx from SE Asia using males

Modified slightly from, and with figure references to, Ballantyne et al. (2011). Interpretation of colour patterns in the elytra in ethanol preserved specimens where the underlying dark hind wing becomes visible may be difficult. Ballantyne (2008) suggested pulling the elytron aside and allowing it to dry.

1. V7 with slender incurving hair bearing lobes along its posterior margin between PLP and MPP (arrowed in Ballantyne et al. 2011 Figs 9, 10 ); LOs in V7 restricted to very small anterolateral plaques; PLP of V7 narrowed and considerably produced beyond the posterior margin of the MPP; T8 prolonged apically beyond MPP with margins converging posteriorly ( Ballantyne et al. 2011 Figs 9, 10 ).................................................................. macdermotti McLean V 7 without slender incurving hair bearing lobes along its posterior margin between PLP and MPP; LOs in V7 never restricted to very small anterolateral plaques; if posterolateral processes of V7 narrowed then not produced far beyond the posterior margin of MPP if at all; T8 not as above...................................................................... 2

2. T8 bearing slender elongate lobes along its posterior margin to either side of the posterior median emargination ( Ballantyne et al. 2011 Figs 7 , 13 ); flanges on ventral surface of T8 absent; basitarsus of legs 2 often excavated in its inner margin ( Ballantyne et al. 2011 Fig. 11 )................................................................................. 3 T8 without slender elongate lobes along its posterior margin to either side of the median posterior emargination (e.g. Ballantyne et al. 2011 Figs 8 , 17 , 19, 24 ); flanges may be present on ventral surface of T8.................................. 6 3. Posterior margin of T8 strongly asymmetrical especially when viewed from above; posterior margin of T7 broadly, shallowly and evenly emarginate; elytral apices broadly rounded (C> A or B); all FS simple; posterolateral corners of V7 produced and rounded; MPP with short narrow paired hooks with apices inclining inwards (Ballantyne 2001 Figs 1–3 )................................................................................................... asymmetria Ballantyne Posterior margin of T8 symmetrical or nearly so ( Ballantyne et al. 2011 Figs 7 , 19, 24 ); any asymmetry is in the paired lobes arising at each side of the MPP, and not an asymmetry of the entire posterior margin; posterior margin of T7 either scarcely emarginated and slightly bisinuate, or moderately deeply emarginated with acute posterolateral corners and straight anterior margin; elytral apices rounded or margin B obliquely truncate; FS1 slightly expanded in median area in decolor ; posterolateral corners of V7 produced and rounded, or not produced and angulate; posterior margin of T8 deeply emarginated in middle area with posterolateral corners produced and rounded, or barely and very narrowly emarginated in median line only.......... 4

4. Posterolateral processes of V7 angulate, not or scarcely produced posteriorly ( Ballantyne et al. 2011 Figs 7 , 12 ); posterior margin of V7 between PLP and MPP slightly sinuate; posterior margin of T7 with narrowed angulate corners and a small shallow median emargination (margin appears trisinuate); elytral apices obliquely truncate across most of their anterior (outer) margin (C); aedeagus dimensions B/A 0.6................................................................ tener Olivier Posterolateral processes of V7 rounded obtuse, and produced posteriorly; posterior margin of V7 between PLP and MPP with moderately deep and rounded emarginations; posterior margin of T7 deeply emarginated in middle area with posterolateral corners produced and rounded; posterior margin of T7 not appearing trisinuate; elytral apices C rounded or truncate; aedeagus dimensions B/A> 0.6.................................................................................. 5

5. Dorsal surface entirely pale coloured (dark markings at tip of elytra may be present); pronotum without any pink markings; head and anterior margin of scape pale yellow with labrum dark brown; elytral apices rounded; projections on either side of median emargination of T8 relatively broad and apically rounded; aedeagus dimensions B/A 0.75.......... decolor Olivier Elytra pale brown, semitransparent, with lateral margin paler than rest and pronotum often deep pinkish orange; if elytra pale then at least base and apex brown; head pale brown, with labrum slightly darker; elytral apex B obliquely truncate (Figs); projections to either side of median emargination of T8 slender and apically pointed; aedeagus dimensions B/A 0.85................................................................................................... bearni Olivier

6. Deflexed elytral apex shortened (wider than long) ( Ballantyne et al. 2011 Fig. 23 ); tibiae 3 not expanded; basitarsi 3 not swollen; fine ventrally directed flanges on ventral surface of T8 absent; PLP separated from MPP by moderately deep circular emarginations ( Ballantyne et al. 2011 Fig. 18 ); apices of PLP broad, flat, slightly obliquely truncate; posterior margin of T7 not emarginated and posterolateral corners not produced; lateral margins of T8 rounded.............. truncata Ballantyne Deflexed elytral apex not shortened (i.e. about as wide as long); tibiae 3 often expanded at apex and basitarsi 3 may be swollen ( Ballantyne et al. 2011 Fig. 26 ); fine ventrally directed flanges may be present on the ventral surface of T8 ( Ballantyne et al. 2011 Fig. 17 ); PLP separated from MPP by moderately deep circular emarginations or not; apices of PLP sometimes flat, slightly obliquely truncate; posterior margin of T7 usually emarginated and posterolateral corners produced; lateral margins of T8 rounded or straight and converging anteriorly............................................................. 7

7. Posterior end of elytra dimpled ( Ballantyne et al. 2011 Figs 20, 21 ); wide deep emarginations separating elongate narrow PLP from MPP; apices of PLP narrow and rounded; posterolateral corners of T8 angulate and lateral margins converge anteriorly; posterolateral corners of T7 narrowed and may project and are often visible from beneath in the emarginations between PLP and MPP ( Ballantyne et al. 2011 Fig. 19 )................................................... gelasina Ballantyne Posterior end of elytra not dimpled; either wide emarginations separating elongate PLP from MPP, or emarginations scarce; apices of PLP often slightly oblique, or PLP broadly rounded and scarcely produced; posterolateral corners of T8 angulate and lateral margins converge anteriorly or corners and lateral margins rounded; posterolateral corners of T7 not usually visible from beneath in the emarginations between PLP and MPP..................................................... 8

8. Posterolateral corners of V7 rounded or angulate, scarcely produced; MPP of V7 broad and apex almost squarely truncate or slightly rounded in ventral view and perpendicular to horizontal plane; median dorsal surface of MPP may be narrowly prolonged and apically emarginated.......................................................................... 9 PLP of V7 elongate, longer than wide, produced and apically obliquely truncated; MPP of V7 narrower and apex emarginated (emargination visible from beneath); median dorsal surface of MPP not developed ( Ballantyne et al. 2011 Figs 24, 25 )...................................................................... masatakai Kawashima , malaccae (Gorham) 1

9. Dorsal surface of MPP of V7 strongly prolonged and apically narrowly emarginated....................................................................... sulawesiensis Kawashima ; valida Olivier sensu Ballantyne (2001 Group 2:81) 2 Dorsal surface of MPP either without a median posterior projection or with a slight projection....................... 10

10. Elytral apices strongly deflexed ( Ballantyne et al. 2011 Fig. 32 ); posterior margin of outer area of elytral apex grooved; LO in V7 well separated in the middle ( Ballantyne et al. 2011 Figs 27, 31 ).... valida Olivier sensu Ballantyne (2001) Groups 1, 3 3 Elytral apices not strongly deflexed ( Ballantyne et al. 2011 Fig. 2 ); posterior margin of outer area of elytral apex not grooved; LO in V7 contiguous in the middle ( Ballantyne et al. 2011 Fig. 2 ) maipo Ballantyne

Footnotes 1. These species are not further distinguished here; Ballantyne (2001) identified malaccae in four distinct morphological groups from peninsular Malaysia, Sarawak, Thailand and Indonesia (Borneo); Kawashima (2003) did not align masatakai with any of these groups and malaccae and masatakai are distinguished here by the occurrence of masatakai on the island of Sulawesi. 2. Ballantyne (2001) distinguished a group of specimens of valida from peninsular Malaysia at Selangor, Sarawak and Indonesia (Bali) with a median posterior projection of the dorsal surface of MPP. 3. Ballantyne (2001) distinguished a single specimen from peninsular Malaysia as group 3.

Remarks. Two species of flashing fireflies in Malaysia both belonging to the genus Pteroptyx Olivier are similar, and their inconsistent identification in various publications, (not least those of the first author), is addressed and resolved here. The species are Pteroptyx tener Olivier , a mass synchronising species that is the basis for the tourist development involving firefly watching on the Selangor River in Western peninsular Malaysia, and Pteroptyx bearni Olivier , a non synchronising species sometimes found in the same habitats as tener . The problems in identification have arisen primarily because Pteroptyx bearni was first redescribed without examination or description of a type, which was not located in the initial investigations ( Ballantyne & McLean 1970). Subsequent publications, including depictions of this species on Malaysian stamps, assumed this characterisation to be definitive and are discussed below.

Ballantyne located in the Pic collection from the Natural History Museum Paris a single specimen which can be regarded as a surviving syntype of P. bearni and it is described here, and issues covering identifications of this species and P. tener are addressed.

Figures

FIGURE 1. Aquatica leii (Fu et Ballantyne) male. Taken at 10.33 am 12.vii.2011 in Wuhan by X. H. Fu.

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

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 %.

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 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.