Pygoluciola Wittmer 1939

Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, Zootaxa 4687 (1), pp. 1-174 : 120-122

Pygoluciola Wittmer 1939

Figs 38, 41−43 , 50, 51 , 364−462

Pygoluciola Wittmer 1939: 21 . Ballantyne 2008: 1 . Ballantyne & Lambkin 2006: 21; 2009: 107; 2013: 108. Ballantyne Lambkin Boontop et al. 2015: 8. Ballantyne Lambkin Luan et al. 2016: 204. Fu & Ballantyne 2008: 1 . Fu Ballantyne & Lambkin 2010: 2; 2012: 6. Wattanachaiyingcharoen & Nak-Eiam 2012: 24. Yiu 2017: 105.

Luciola subgenus Pygoluciola (Wittmer) . McDermott 1966: 115; Ballantyne 1968: 119; Ballantyne & Lambkin 2000: 82; 2001: 361; Ballantyne & McLean 1970: 233.

Type species. Pygoluciola stylifer Wittmer 1939 , by monotypy ( RMNH) .

Diagnosis. Repeated with slight modification from Nada & Ballantyne (2018). Many Pygoluciola are difficult to diagnose to genus using only external morphology and may require dissection. All species of Pygoluciola have a similar distinctive pattern of aedeagal sheath and aedeagus: the aedeagal sheath has an elongate narrow sternite which extends well beyond the lateral tergite articulations and may bear paired lobes at its apex (four species have the sheath sternite terminated by a boomerang shaped lobe). The aedeagal LL are divisible into two sections, a basal well sclerotized portion which has an asymmetrical anterior margin and a short median dorsal separation, and an apical portion which is membranous and elongated usually well beyond the rounded apex of the median lobe.

Using only external morphology, Pygoluciola exists in at least three different forms which can be diagnosed individually:

1. The form described originally by Wittmer (1939) has V7 with a median posterior prolongation which often curves dorsally, and may be engulfed by a similar prolongation of the midposterior margin of T8, which curves ventrally.

2. In P. dunguna Nada and P. tamarat Jusoh sp. nov. the median posterior margin of V7 is prolonged and inclines slightly upward where the tip of the MPP abuts against the underside of the narrow T8.

3. In both P. phupan sp. nov. and P. qingyu (Fu et Ballantyne) there is no MPP and the posterior margin of V7 is evenly rounded; the posterior margin of T 8 in both is slightly downturned, narrowly in P. qingyu (at right angles to the longitudinal axis of the body), more well defined in P. phupan sp. nov.

LOs in V7 are often retracted from the lateral and posterior margins and these lateral margins may be uprolled. T8 is always narrower than T7, often with parallel sides, elongate slender anterolateral prolongations, and entire posterior margin; dorsoventral muscles joining the lateral margins of T7 to the ventrite below create depressed areas at the sides of T7. Known females are macropterous and bursa plates appear to be hooked with a single point of attachment to the inside walls of the bursa and up to three points of irregular length which incline in an anterior, posterior and inner direction. Larvae associated by breeding for P. qingyu only are terrestrial with strong well sclerotized dorsal plates, and mandibles with two inner teeth. Larvae associated with catches of adult P. dunguna may be riparian or even aquatic, having many blunt spines arising from the margins of all tergal plates but the terminal one. They also have mandibles with two inner teeth.

Remarks. Wittmer (1939) described the genus Pygoluciola from only one species Pygoluciola hamulata Wittmer , having pronounced modifications of the terminal abdomen where both the median posterior areas of V7 and T8 are prolonged and may incline, V7 dorsally and T8 ventrally, and in dried pinned specimens they may envelop each other. Pygoluciola was submerged under the genus Luciola by McDermott (1966) and treated as a subgenus by Ballantyne (1968) and Ballantyne & Lambkin (2001), until Ballantyne & Lambkin (2006) reassigned it to generic status. Subsequently Ballantyne (1968), Ballantyne & Lambkin (2001, 2006), and Ballantyne (2008) described five more species.

Pygoluciola Wittmer was thought to be a rare genus as it was poorly represented in collections ( Ballantyne 2008; Ballantyne & Lambkin 2001, 2006). We now know that it exists in several forms as described above. The first as described by Wittmer (1939) is the rarer form, and consists of six species (see Table 24 ) which are easily recognised by the modifications to the terminal abdomen in males;

Fu & Ballantyne (2008) first realised that Pygoluciola could exist in a second form where the pronounced abdominal modifications described above did not exist. Species having this form are only recognisable on dissection, where the typical Pygoluciola patterns of genitalia are seen: aedeagus with LL considerably longer than ML and membranous in their apical half; basal portion of LL well sclerotised and not fused along all of their dorsal length; aedeagal sheath symmetrical, with narrow elongated anterior portion of sheath sternite expanding in its posterior area and often terminated by paired lobes; lateral arms of tergite of sheath visible beside the sheath sternite. Fu and Ballantyne (2008) described a distinctively coloured species from China as P. qingyu . Nak-Eiam in Thailand has discovered several species with similar genitalic characters to P. qingyu , and it seems that Pygoluciola is much more widespread than previously thought (Nak-Eiam 2015). Wattanachaiyingcharoen & Nak-Eiam (2012) distinguished mainland (Asian) species and island (Borneo and Mindanao) and indicated that the pronounced terminal abdomen modifications seemed to be characteristic of the island species. They felt the long separation of the mainland and islands mentioned may have led to the separation of the different forms of this genus. Fu (2014: 65, 67, 68) identified three possible Pygoluciola sp. from China which will be addressed elsewhere. He associated a larva for the unidentified species on page 65, and listed as ‘ Luciola sp.’ on page 46 a species which we would assign to Pygoluciola , it having the ‘boomerang’ shaped piece at the posterior end of the aedeagal sheath sternite. Of the four species we characterise here with that feature, three have pale margined elytra, while the dorsal colour of the fourth, P. abscondita comb. nov., is much paler than that illustrated by Fu.

Figures

FIGURES 38−43. Key to genera characters 6. 38, 43 Pygoluciola qingyu; 39, 40 Abscondita pallescens comb. nov.; 41, 42 Pygoluciola hamulata type male. 38−40 aedeagal sheath ventral (38, 39) and slightly right lateral (40) (arrows indicate position where lateral tergal arms join the median sternite); 41, 43 terminal abdomen ventral; 42 termnal abdomen left lateral. Figure legends: V7 ventrite 7; T8 tergite 8.

FIGURES 50–56. Key to genera characters 8. 50, 51 Pygoluciola satoi; 52, 53 Emeia pseudosauteri; 54, 55, 56 Aquatica ficta. 50, 51 aedeagus dorsal (50: arrow indicates separation in basal area of lateral lobes), and ventral; 52, 56 dorsal habitus; 53, 55 ventral habitus;; 54 aedeagal sheath ventral semidiagrammatic (arrows indicate pointed projections along sides of sheath sternite).

FIGURES 364−373. Pygoluciola ambita comb. nov. males. 366, 367 type male Luciola ambita (MNHN); 364, 365, 368−373 Java male (Prague). 364 habitus dorsal; 365, 366 habitus ventral; 368, 369 abdomen: 368 dorsal; 369 ventral; 370−372 ae- deagus: 370 ventral; 371 dorsal; 372 left lateral; 373 aedeagal sheath ventral with aedeagus contained. Scale lines are 1 mm.

FIGURES 374−381. Pygoluciola bangladeshi sp. nov. type males. 374, 376 habitus dorsal (376 pronotum and anterior portion of elytra only); 375 habitus ventral; 377 aedeagal sheath ventral (arrows indicate hairy lobes arising from boomerang shaped piece at apex of sheath sternite; 378, 379 aedeagus: 378 dorsal, 379 ventral; 380, 381: abdomen: 380 ventral; 381 dorsal. Scale lines are 1 mm.

FIGURES 382−393. Pygoluciola calceata comb. nov. 382, 383, 384, 388 holotype male (MNHN); 385−387, 389−393 Nilgiri Hills male (NHML). 382, 384 habitus dorsal (384 pronotum only); 385, 386 abdomen: 385 ventral; 386 dorsal; 387 tergite 8 ventral with aedeagal sheath and aedeagus contained; 388 type label; 389, 390 aedeagal sheath ventral: 389−393 aedeagus: 391 ventral; 392 left lateral; 393 dorsal.

FIGURES 394−404. Pygoluciola insularis comb. nov. 394, 395, 400 type female (MNHN); 396−399, 401−404 Andaman Islands males (NHML). 394, 397 habitus ventral; 395, 396 habitus dorsal; 398, 399 abdomen: 398 ventral; 399 dorsal; 400 type label; 401 aedeagal sheath ventral; 402−403 aedeagus: 402 ventral; 403 dorsal; 404 tergite 8 ventral with aedeagal sheath and aedeagus contained.

FIGURES 405−417. Pygoluciola matalangao sp. nov. type males. 405 habitus dorsal; 406 habitus ventral; 407 head anterior; 408, 409 dorsal pronotum; 410 abdomen ventral tergite 8 removed; 411 terminal abdominal segments dorsal, tergite 8 removed (arrows indicate dorsally reflexed margins of ventrite 7); 412 tergite 8 ventral; 413−415 aedeagus: 413 ventral; 414 left lateral; 415 dorsal; 416, 417 aedeagal sheath: 416 dorsal; 417 ventral.

FIGURES 418−421. Pygoluciola matalangao sp. nov. females. 418 habitus dorsal; 419 habitus ventral; 420, 420 pronotum and base of elytra, dorsal.

FIGURES 422−429. Pygoluciola nitescens comb. nov. male. 422, 423 Borneo coll. Chaper (MNHN); 424−429 Sarawak coll Polunin (ANIC). 422, 424 habitus dorsal; 423 habitus ventral; 425, 426 terminal abdomen: 425 ventral; 426 dorsal; 427−429 aedeagal sheath: 427 dorsal; 428 ventral (outline of lobes arising from boomerang shaped piece at apex of sheath sternite arrowed); 429 left lateral.

FIGURES 430–436. Pygoluciola nitescens comb. nov. 430−432 male (Sarawak coll. Polunin;ANIC); 433−436 female (Ranau; NHML). 430−432 aedeagus: 430 ventral; 431 dorsal; 432 left lateral; 433 habitus dorsal; 434 habitus ventral; 435 abdominal ventrite 8, ventral; 436 reproductive system, ovipositor to right, spermatophore digesting gland and spermatheca to left (arrow indicates area of bursa and possible thin bursa plates). Scale lines are 1 mm in 430, 435 and 2 mm in 433.

FIGURES 437−448. Pygoluciola phupan sp. nov. type males. 437, 438 habitus dorsal; 439, 440 tergite 8: 439 ventral; 440 right lateral (projections arrowed); 441 habitus ventral; 442− 444 abdomen: 442 ventral; 443 dorsal; 444 right lateral; 445, 446 aedeagus: 445 ventral; 446 dorsal (transmitted light); 447, 448 aedeagal sheath: 447 dorsal; 448 ventral. Scale lines are 1 mm unless otherwise indicated.

FIGURES 449−455. Pygoluciola tamarat sp. nov. type males. 449 habitus dorsal; 450 habitus ventral; 451 ventrite 7 dorsal surface; 452, 453 aedeagus: 452 ventral; 453 dorsal; 454 aedeagal sheath ventral; 455 tergite 8 dorsal. Scale lines are 1 mm.

FIGURES 456−462. Pygoluciola vitalisi comb. nov. males. 456 habitus dorsal; 457 habitus ventral; 458, 461, 462 aedeagus: 458 left lateral; 461 ventral; 462 dorsal; 459, 469 aedeagal sheath: 459 ventral (with aedeagus); 460 ventral without aedeagus. Scale lines are 1 mm.

Tables

TABLE 24. List of species of Pygoluciola Wittmer (*References: 1 = Ballantyne 1968; 2 = Ballantyne & Lambkin 2001; 3 = Ballantyne 2008; 4 = Fu & Ballantyne 2008; 5 = Ballantyne & Lambkin 2013; 6 = Ballantyne & Lambkin 2006; 7 = Yiu 2017; 8 = Nada & Ballantyne 2018. # = species keyed in Ballantyne 2008.)

Species
													of species *
	Australia	Malaysia peninsula	Island of Borneo	Myanmar	China	Sri Lanka	Indonesia Java Sumatra	Andaman Islands	India	Bangladesh Philippines Location of types		Females associated	Larvae associated References for illustrations
abscondita ( Olivier 1891) comb.				*							MNHN
nov. ambita ( Olivier 1896) comb. nov.							*				MNHN
bangladeshi Ballantyne sp. nov.										*	ANIC	yes
calceata ( Olivier 1905) comb. nov.						*			*		MNHN
cowleyi ( Blackburn 1897)	*										NHML		5
dunguna Nada 2018		*									FRIM	yes	8
guigliae ( Ballantyne 1968)#			*								BPBM	yes	1 6
hamulata ( Olivier 1885) #			*								CMG		1 3
insularis ( Olivier 1883) comb. nov.							*	*			MNHN
kinabalua (Ballantyne & Lambkin			*								NHML	yes	2
2001)# matalangao sp. nov. Ballantyne											ANIC	yes
nitescens ( Olivier 1905) comb. nov.			*						*		MNHN	yes
phupan Ballantyne sp. nov.											NHML
qingyu Fu & Ballantyne 2008					*						BEIJING	yes	4 7
satoi ( Ballantyne 2008) #										*	ZRC		3
stylifer Wittmer 1939 #			*								LEIDEN		6
tamarat Jusoh sp. nov.											MZUM
vitalisi ( Pic 1934) comb. nov.									*		MNHN
wittmeri ( Ballantyne 1968)#			*								BPBM		1