Pteroptyx Olivier 1902

Pteroptyx Olivier 1902 View in CoL

Pteroptyx Olivier, 1902: 72 View in CoL ; 1907: 55; 1909a: 319; 1909b:lxxxii; 1910: 47; 1911: 16; 1913b: 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 View Cited Treatment (partim). nec Ballantyne in Calder 1998: 180.

Type species: Luciola testacea Motschulsky designated by Lucas 1920. ( ZMMU) .

Diagnosis. Pteroptyx 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 have a MFC, deflexed elytral apices in all but Pt. surabayia sp. nov., Pt. sayangia sp. nov. and Pt. galbina sp. nov., aedeagal sheath with bulbous paraprocts, and bipartite LOs in V7 except in Pt. surabayia sp. nov., Pt. sayangia sp. nov. and Pt. galbina sp. nov. Males are distinguished from Australoluciola which has entire LOs in V7 and no MFC and no deflexed elytral apices; from Colophotia which has a median carina on V7, expanded and oblique PLP and elongate slender aedeagal sheath, no MFC and no deflexed elytral apices; from Pyrophanes and Poluninius both of which have incurving lobes along the posterior margin of V7 (bulbous in Poluninius ), bipartite LOs in V7 and an MFC but no deflexed elytral apices; from Trisinuata and Medeopteryx which have no MFC and no bulbous aedeagal sheath paraprocts. Pteroptyx differs from Luciola indica which has no deflexed elytral apices and entire LOs in V7 most obviously in being without the bulbous median lobe.

Ballantyne & McLean (1970) redescribed the genus from 17 species (three were species incertae). Ballantyne (1987a) included 21 species, and Ballantyne et al. (2011) added a further new species Pt. maipo Ballantyne. Ballantyne & Lambkin (2013) redefined the breadth of the genus, listing and keying 12 species of Pteroptyx that was restricted to oriental species having a MFC. At the same time Ballantyne & Lambkin erected two other genera ( Medeopteryx and Trisinuata ) to accommodate species without a MFC which also have deflexed elytral apices in the male. With the extended morphological concept of the genus it is now appropriate that it should be fully redescribed here.

Redescription of Pteroptyx Olivier. Male. Pronotum ( Figs. 77 View FIGURES 77−87 , 88, 93, 101, 103): dorsal surface without irregularities in posterolateral areas and longitudinal groove in lateral areas (1, 2); punctation dense (sparse in Pt. macdermotti ); anterior margin not explanate (6); either subparallel-sided, margins straight (A=B=C in Pt. sayangia sp. nov.), or with some divergence in anterior 1/3 with remainder subparallel-sided (B=C), or lateral margins diverging posteriorly along their length (C>A, B in Pt. macdermotti ), or lateral margins widest across middle (B>A, C); width <humeral width; anterolateral corners rounded obtuse or angulate; lateral margins without indentation at mid-point, and sinuousity in either horizontal or vertical plane (16); without indentation in lateral margin near posterolateral corner (17), and irregularities at corner (18); posterolateral corners rounded or angulate, rounded corners slightly obtuse or subequal to 90°, angulate corners subequal to 90°; 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 (16); median area more widely flattened than elsewhere.

Elytron ( Figs. 77, 78 View FIGURES 77−87 , 88, 89, 93, 94, 102, 103, 113, 114): punctation dense, not linear (35, 36), not as large as that of pronotum, nor widely and evenly spaced (34); apices usually deflexed, not deflexed in three species where the V7 LO is also entire; deflexed elytral apices can be rounded or truncate, with the deflexed portion short in Pt. maipo and Pt. truncata ; epipleuron and sutural ridge extend beyond mid-point, almost to apex but not extending around apex when apex not deflexed, neither thickened in apical half; no interstitial lines; elytral carina absent (54); in horizontal specimen viewed from below epipleuron at elytral base wide, covering humerus (44); viewed from above anterior margin of epipleuron arises 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: usually moderately depressed between eyes, with minimal depression in Pt. macdermotti , Pt. truncata ; 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 except for wide separation in Pt. malaccae ; eyes above labrum close to moderately separated, SIW/GHW moderate, wide in Pt. macdermotti , close in Pt. truncata ; 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; clypeolabral suture present, flexible, not in front of anterior eye margin when head viewed with labrum horizontal; outer edges of labrum usually reach beyond inner edges of closed mandibles except in Pt. maipo and Pt. galbina sp. nov. Mouthparts: functional; apical labial palpomere flattened, shaped like narrow triangle (narrowest at base and L 2–3 X W except in Pt. maipo where L=W), with inner edge entire, and at least half as long as apical maxillary palpomere except in Pt. maipo where it is slightly less than ½ the length. Antennae 11 segmented; length>GHW up to twice GHW; pedicel produced at outer apical angle in Pt. gelasina , Pt. macdermotti ; FS1 expanded at apex in Pt. gelasina , Pt. macdermotti and Pt. malaccae , and expanded in median area in Pt. macdermotti ; FS4 expanded at outer apical angle in Pt. macdermotti ; FS1 not shorter than pedicel.

Legs ( Figs. 78 View FIGURES 77−87 , 89, 94, 102, 108, 116): with inner tarsal claw not split (95); with MFC (presence or absence not reliably determined in Pt. testacea ); no femora or tibiae swollen or curved (97̄99), tibiae 3 sightly expanded towards apex in Pt. malaccae ; basitarsus 1 excavated on inner margin in Pt. tener .

Abdomen ( Figs. 79−82 View FIGURES 77−87 , 90, 94−96, 102, 104, 105, 114, 115): without cuticular remnants in association with aedeagal sheath (132); posterior margin of V3 usually recurved (106̄108) (not recurved in Pt. sayangia sp. nov., Pt. g albina sp. nov.); posterior margin of V4 recurved except in Pt. sayangia sp. nov., Pt. galbina sp. nov., Pt. maipo and some Pt. surabayia sp. nov. (neither character reliably established in Pt. testacea ); anterior margin of V4 extending forwards into posterior margin of V3 where this is recurved; LO in V7 usually bipartite, entire in three species which also have non deflexed elytral apices (apparently entire in Pt.. testacea Figs. 114, 115 View FIGURES 113-116 ); entire LOs occupying most of V7, and reaching to sides but not always to posterior margin; sometimes reaching MPP and PLP; LO with anterior and posterior margins straight; LO present in V6, occupying almost all V6. MPP present, symmetrical, apex rounded or truncate (broad and apically acute with minute median emargination in Pt. testacea ), usually shallowly emarginate (deeply so in Pt. macdermotti ), not laterally compressed (162), short or slightly longer than wide (L<W or L>W), very wide in Pt. macdermotti , not inclined dorsally nor engulfed by T8 apex (169), without dorsal ridge (172), median longitudinal trough (176). V7 without median carina (114), median longitudinal trough (117), anteromedian depression on face of LO, incurving lobes (121) or pointed projections (127), median ‘dimple’ (128), or reflexed lobes (129); posterior margin of V7 trisinuate; MPP shorter than, as long as, or longer than PLP (PLP not obviously developed in Pt. testacea and posterolateral corners rounded); wider than or as wide as PLP; PLP usually moderately produced (greatly so in Pt. macdermotti , scarcely projecting in Pt. tener ), narrower than or as wide as MPP; PLP L>W, L=W or W>L. T7 without prolonged anterolateral corners. T8 well sclerotised, symmetrical, W=L, visible posterior area not narrowing abruptly except in Pt. macdermott i where lateral margins converge posteriorly, median posterior margin shallowly and narrowly emarginate (deeply so in Pt. macdermotti ); T8 usually widest across middle; without prolonged posterolateral corners (191), not inclined ventrally nor engulfing posterior margin of V7 nor MPP (185, 191), not extending conspicuously beyond posterior margin of V7 except in Pt. macdermotti ; T8 ventral surface with well developed median longitudinal trough, margined by well defined symmetrical ridges; anterior end of ridges either not produced, or with symmetrical short narrow acute flanges in Pt. maipo and Pt. valida ; without lateral depressed troughs (206), asymmetrical projections (207), median posterior ridge (208); concealed anterolateral arms of T8 as long as, or slightly shorter than visible posterior portion of T8, not laterally emarginated before their origins (214), not expanded dorsoventrally (216), expanded only in horizontal plane; without bifurcation of inner margin (217) and ventrally directed pieces; lateral margins of T8 not enfolding sides of V7 (189); posterior margin emarginated, sometimes with elongate slender projections at each side.

Aedeagal sheath ( Figs. 83, 84 View FIGURES 77−87 , 100, 110): symmetrical; approx. 3 times as long as wide; with 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 (252); tergite without projecting pieces along posterior margin of T9 (256), anterior margin without transverse band (257).

Aedeagus ( Figs. 85 View FIGURES 77−87 ̄87, 91, 97̄99, 109, 111, 112): symmetrical; L/W2.9–6.0; LL lateral appendages absent (306); 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; LL separated longitudinally by at least half their length, sometimes less; LL base width wider than, not = LL apex width which is narrower than that of ML; LL apices not expanded in horizontal plane (294); dorsal base of LL symmetrical, may be excavated or slightly produced; LL without lateral hairy appendages along their outer ventral margins (304), not produced preapically nor narrowly on inner apical margin (307), apices of LL not inturned, nor out-turned (314, 316); without projection on left LL (319); 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 (323−325).

Female ( Figs. 106, 107 View FIGURES 101-112 ): Macropterous and observed in flight in certain species. Pronotum without irregularities in posterolateral areas; punctation moderate to dense; pronotal width less than humeral width; without indentation of lateral margin or 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 paired plates or paired single plates in Pt. maipo , Pt. valida and Pt. surabayia sp. nov. ( Fig. 107 View FIGURES 101-112 ).

Larva. Reliably associated by breeding for Pt. maipo , Pt. tener and Pt. valida only; terrestrial; elongate, slender, spindle shaped ( Ballantyne et al. 2011; Fu et al. 2012b), 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 midline; without brush of hairs from apex of tibiotarsus; mandibles without inner teeth; antennal segment 3 short, sense cone adjacent to segment 3 short, wide.

List of species of Pteroptyx

Pteroptyx asymmetria Ballantyne 2001

Pteroptyx bearni Olivier 1909

Pteroptyx decolor Olivier 1911

Pteroptyx galbina Jusoh sp. nov.

Pteroptyx gelasina Ballantyne 2001

Pteroptyx gombakia Ballantyne sp. nov.

Pteroptyx maipo Ballantyne 2011

Pteroptyx malaccae ( Gorham 1880)

Pteroptyx macdermotti McLean 1970

Pteroptyx masatakai Kawashima 2003

Pteroptyx sayangia Ballantyne sp. nov.

Pteroptyx sulawesiensis Kawashima 2003

Pteroptyx surabayia Ballantyne sp. nov.

Pteroptyx tener Olivier 1907

Pteroptyx testacea (Motsch. 1854)

Pteroptyx truncata Ballantyne 2001

Pteroptyx valida Olivier 1909

Key to species of Pteroptyx View in CoL from SE Asia using males. Modified from Ballantyne & Lambkin (2013)

1. Without deflexed elytral apices........................................................................... 2

- With deflexed elytral apices............................................................................. 4

2. Pronotal anterolateral and posterolateral corners angulate with lateral margins subparallel-sided (A=B=C)........................................................................................... sayangia View in CoL sp. nov. (Figs. 93, 94)

- Pronotal corners rounded, lateral margins subparallel-sided in posterior 2/3 (B=C)................................... 3

3. LO in V7 not extending into either MPP or PLP; MPP not medially emarginated........ surabayia View in CoL sp. nov. ( Figs. 102, 104 View FIGURES 101-112 )

- LO in V7 extending into both MPP and PLP; MPP medially emarginated.................. galbina View in CoL sp. nov. ( Figs. 78, 79 View FIGURES 77−87 )

4. Posterior margin of V7 with incurving lobes between the MPP and PLP (Fig. 90)................................... 5

- Posterior margin of V7 without incurving lobes between the MPP and PLP........................................ 6

5. V7 with slender incurving hair bearing lobes along its posterior margin between PLP and MPP (arrowed in Ballantyne et al. 2011: Figs. 9, 10 View FIGURES 3-10 ); 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 View FIGURES 3-10 )................................................................. macdermotti McLean View in CoL

- V7 with broad flat hairless lobes along its posterior margin between the PLP and MPP; LO in V7 entire and occupying most of the area of V7; PLP short; T8 not prolonged apically beyond the MPP (Fig. 90)...................... gombakia View in CoL sp. nov.

6. T8 bearing slender elongate lobes along its posterior margin to either side of the posterior median emargination ( Ballantyne et al. 2011: Figs. 7 View FIGURES 3-10 , 13 View FIGURES 11–17 ); flanges on ventral surface of T8 absent; basitarsus of legs 2 often excavated in its inner margin ( Ballantyne et al. 2011: Fig. 11 View FIGURES 11–17 ).................................................................................7

- 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 View FIGURES 3-10 , 17 View FIGURES 11–17 , 19, 24 View FIGURES 18-25 ); flanges may be present on ventral surface of T8................................ 10

7. 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 View FIGURE 1 View FIGURE 2 View FIGURES 3-10 ).................................................................................................. asymmetria Ballantyne View in CoL

- Posterior margin of T8 symmetrical or nearly so ( Ballantyne et al. 2011: Figs. 7 View FIGURES 3-10 , 19, 24 View FIGURES 18-25 ); 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; FS 1 slightly expanded in median area in decolor View in CoL ; 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..........8

8. Posterolateral processes of V7 angulate, not or scarcely produced posteriorly ( Ballantyne et al. 2011: Figs. 7 View FIGURES 3-10 , 12 View FIGURES 11–17 ); 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 View in CoL

- 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...................................................................................9

9. 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 ( Ballantyne & McLean 1970 Fig. 8 h, i View FIGURES 3-10 )................................................................................ decolor Olivier View in CoL

- 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; projections to either side of median emargination of T8 slender and apically pointed; aedeagus dimensions b/a 0.85 ( Ballantyne & Lambkin 2013 Figs. 191−198)...................................................................... bearni Olivier View in CoL

10. Deflexed elytral apex shortened (wider than long) ( Ballantyne et al. 2011: Fig. 23 View FIGURES 18-25 ); 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 View FIGURES 18-25 ); 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 View in CoL

- 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 View FIGURES 26-32 ); fine ventrally directed flanges may be present on the ventral surface of T8 ( Ballantyne et al. 2011 Fig. 17 View FIGURES 11–17 ); 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............................................................ 11

11. Posterior end of elytra dimpled ( Ballantyne et al. 2011 Figs. 20, 21 View FIGURES 18-25 ); 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 View FIGURES 18-25 ).................................................. gelasina Ballantyne View in CoL

- 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.................................................... 12

12. 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......................................................................... 13

- 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 View FIGURES 18-25 )...................................................................... masatakai Kawashima View in CoL , malaccae (Gorham) View in CoL 1

13. Dorsal surface of MPP of V7 strongly prolonged and apically narrowly emarginated..................................................................... .. sulawesiensis Kawashima View in CoL ; valida Olivier sensu Ballantyne (2001 View in CoL Group 2: 81) 2

- Dorsal surface of MPP either without a median posterior projection or with a slight projection........................ 14

14. Elytral apices strongly deflexed ( Ballantyne et al. 2011 Fig. 32 View FIGURES 26-32 ); posterior margin of outer area of elytral apex grooved; LO in V7 well separated in the middle ( Ballantyne et al. 2011 Figs. 27, 31 View FIGURES 26-32 )... valida Olivier sensu Ballantyne (2001) Groups View in CoL 1 & 3 3

- Elytral apices not strongly deflexed ( Ballantyne et al. 2011: 2); posterior margin of outer area of elytral apex not grooved; LO in V7 contiguous in the middle ( Ballantyne et al. 2011 Fig. 2 View FIGURE 2 )..................................... maipo Ballantyne View in CoL

Notes. 1) These species are not further distinguished here; Ballantyne (2001) identified Pt. malaccae View in CoL in four distinct morphological groups from peninsular Malaysia, Sarawak, Thailand and Indonesia (Borneo); Kawashima (2003) did not align Pt. masatakai View in CoL with any of these groups and Pt. malaccae View in CoL and Pt. masatakai View in CoL are distinguished here by the occurrence of Pt. masatakai View in CoL on the island of Sulawesi. 2) Ballantyne (2001) distinguished a group of specimens of Pt. valida View in CoL 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 Pt. valida View in CoL group 3.