Pygatyphella (Ballantyne)

Ballantyne, Lesley A. & Lambkin, Christine, 2009, Systematics of Indo-Pacific fireflies with a redefinition of Australasian Atyphella Olliff, Madagascan Photuroluciola Pic, and description of seven new genera from the Luciolinae (Coleoptera: Lampyridae), Zootaxa 1997, pp. 1-188 : 77-80

Pygatyphella (Ballantyne)

( Figs 23–26, 60–72, 255–487)

Luciola (Pygatyphella) Ballantyne, 1968:109 .

Type species: Atyphella obsoleta Olivier

Diagnosis. With the exception of one species (Pygat. wisselmerenia ) all species have the LO retracted into the anterior area of V7 and the posterior area arched and often swollen. Many species have a distinctive dorsal colouration that Ballantyne (1968) considered resembled bird droppings.

Male. Pronotum: dorsal surface lacking irregularities in posterolateral areas and longitudinal groove in lateral areas; punctation dense. Anterior margin not explanate.

Pronotum wider across posterior area than rest; pronotal width greater than humeral width. Anterolateral corners rounded obtuse; lateral margins in anterior half divergent posteriorly; lateral margins in posterior half diverge then converge with rounded or angulate convergence (both types of convergence may be seen in one individual, and between individuals in same species) except in Pygat. uberia sp. n., where lateral margins diverge along length; indentation at mid-point absent; lacking sinuousity in either horizontal or vertical plane; indentation in lateral margin near posterolateral corner present or absent (may differ from right to left sides of pronotum and between individuals in same species); irregularities at corner present or absent; posterolateral corners rounded or angulate; rounded corners obtuse, angulate corners approximately 90° or less (never very acute), and inclined obliquely to the median line; posterolateral corners project as far as, or beyond median posterior margin, separated from it by shallow emargination.

Hypomera closed. Median area of hypomeron not elevated vertically; anterior area of hypomeron not flat to side of head; posterior area widely and strongly flattened and strongly adpressed; median area of hypomeron shows marked transition between anterior and posterior areas in New Guinea species; pronotal width/ GHW index 1.6.

Elytron punctation dense, not linear, not as large as that of pronotum, nor widely and evenly spaced; apices not deflexed; epipleuron and suture extend beyond mid-point, do not extend as a ridge around apex and neither is expanded in apical half; 0 or 2 interstitial lines, if developed do not exceed suture; viewed from beneath with specimen horizontal epipleuron at elytral base narrow, covering humerus, and viewed from above arises anterior to posterior margin of the MS; epipleuron developed as lateral ridge along most of length; sutural margins approximate along most of length in closed elytra; lateral margins parallel-sided or convex.

Head moderately, often strongly depressed between eyes; moderately exposed in front of pronotum, or capable of complete retraction within the prothoracic cavity; eyes close to moderately separated beneath at level of posterior margin of mouthpart complex; eyes above labrum close except in pulcherrima where SIW/ GHW is 0.18; frons-vertex junction rounded, not well–defined; lacking median elevation; posterolateral eye excavation not strongly developed and not visible in resting head position; antennal sockets on head between eyes, contiguous or separated by less ASW; clypeolabral suture present, flexible, not in front of anterior eye margin when viewed with labrum horizontal; outer edges of labrum reach inner edges of closed mandibles. Mouthparts functional; apical segment of labial palpi non–lunate, strongly flattened, in the shape of a wide triangle, with inner edge dentate, with 3 or more ‘teeth’. Antennae 11 segmented; length>GHW to twice GHW; no segments flattened, shortened, or expanded; pedicel not produced; FS1 not shorter than pedicel; FS always at least 2 X as long as wide.

Legs with inner tarsal claw not split; lacking MFC; no femora or tibiae swollen or curved; no basitarsi expanded or excavated.

Abdomen lacking cuticular remnants in association with aedeagal sheath; no ventrites with curved posterior margins nor extending anteriorly into emarginated posterior margin of anterior segment; LO in V7 entire, or bipartite; entire LO occupies most (in marginata ), or half or less of V7, often restricted to anterior half of V7 or less, often medially emarginated across its posterior margin, sometimes deeply so; bipartite LO (in uberia sp. n. only) not restricted to anterolateral plaques; entire LO reaching to sides or not, not reaching to posterior margin; bipartite LO occupies> half V7; anterior margin of entire LO in V7 not emarginate; posterior half of V7 arched (except in wisselmerenia sp. n.), may be swollen, transverse muscle impressions usually visible in this area; LO in species with swollen posterior half of V7 emarginated posteriorly; posterolateral corners not produced, distinct or obliterated; if present rounded or angulate; LO present in V6, occupying almost all V6. MPP present, if symmetrical then apex rounded, pointed, squarely truncate, or bisinuate (in okapa sp. n.); if asymmetrical then obliquely truncate; not medially emarginated, not laterally compressed, short in pulcherrima , tagensis ; usually L=W, L>W (in uberia sp. n.), not strongly inclined dorsally; not engulfed by the apex of T8 except in undulata ; MPP with basal emarginations in uberia sp. n.; dorsal ridge of MPP often present, narrow to wide, to left of middle, or median in karimui sp. n, ridge lacking median elevations. V7 lacking median carina, median longitudinal trough, anteromedian depression on face of LO, PLP (except in okapa sp. n. where they are slightly produced, narrow and horizontal), incurving lobes or pointed projections, median ‘dimple’, or reflexed lobes; V7 dorsal surface with paired lobes in peculiaris only. T7 lacking prolonged posterolateral corners. T8 strongly sclerotised, not subparallel-sided, margins usually converge gently towards posterior end; symmetrical, W=L or L>W of visible posterior portion, which does not narrow abruptly; lacking prolonged posterolateral corners, median posterior emargination, median posterior projections, not inclined ventrally nor engulfing the posterior margin of V7 nor the MPP, except in undulata , where the posterior half of T8 is not strongly narrowed, with down turned portion a thickened wide shelf; T8 not extending conspicuously beyond posterior margin of V7 horizontally; T8 ventral surface lacking flanges, lateral depressed troughs; median longitudinal trough absent or present; if present margins fine, slightly thickened or well developed; median longitudinal trough if developed longitudinal except in nabiria sp. n.; trough very short in japenensis sp. n.; trough margins symmetrical except in okapa sp. n.; various asymmetrical projections in anterolateral and posterolateral left and right present in uberia ; median posterior ridge present or not, usually curved, may be straight ( plagiata ); concealed anterolateral arms of T8 present, not laterally emarginated before their origins, narrow, either as long as visible posterior portion or narrow in tomba sp. n., and expanded dorsoventrally except in tomba sp. n. and wisselmerenia sp. n.; apices lacking bifurcation of inner margin; bases with ventrally directed pieces present or absent.

Aedeagal sheath never> 4 times as long as wide; lacking paraprocts; asymmetrical in posterior area with sheath sternite either emarginated on right side from point of attachment of tergite, or subparallel-sided for a third its length past articulation with sheath tergite, and then emarginated on right side, or subparallel-sided along its length in uberia sp. n.; sternite not angulate on L or R sides; posterior margin entire, rounded, not emarginated on either side preapically except in uberia where it is unevenly emarginated preapically on the right side; anterior half of sternite broad, apically rounded; tergite lacking lateral arms that extend widely anteriorly at the sides of the sheath sternite; tergite not subdivided, lacking projecting pieces along posterior margin of tergite 9; anterior margin of tergite lacking transverse band (anterior margin sometimes more heavily sclerotised than.

Aedeagus L/W 3/1 or shorter; LL lack lateral appendages, visible from beneath at sides of ML except in uberia sp. n. where they may not both be visible in same plane, LL/ML wide to moderate; LL of equal length, slightly shorter than ML except in uberia sp. n. where ML much longer than LL; LL diverging along inner margins, and separated there by> half their length; LL base width not=LL apex width which is subequal to or narrower than that of ML; LL apices not expanded horizontally except in eliptaminensis ; dorsal base of LL either symmetrical, or if asymmetrical, strongly produced to right; if symmetrical not excavated, median margin prolonged and broadly rounded; LL lacking lateral hairy appendages along their outer ventral margins, which may be produced preapically and also either narrowly or widely on their inner apical margin obliquely truncate along their preapical inner margins in tagensis , huonensis and lacking strongly developed tooth at anterior end of truncation; apices of LL out–turned in peculiaris ; lacking projection on left LL only; inner margins lacking slender leaf-like projection; ML symmetrical, if asymmetrical then curved to left horizontally; paired lateral teeth may be present; lacking tooth to left side, not strongly arched, and apex not in shape of arrowhead, not bulbous, not inclined ventrally; BP not very narrow, not strongly sclerotised, not hooded, usually in two pieces and not strongly emarginated along anterior margin.

Female. Macropterous and either assumed capable of flight or observed in flight. Pronotum shaped as for male; lacking irregularities in posterolateral areas; punctation moderate to dense; pronotum> humeral width; indentation of lateral margin often present (may differ from one side of pronotum to the other in the one individual and between individuals in the one population), irregularities near posterolateral corner present or absent. Elytral punctation not as large as that of pronotum nor evenly spaced, punctation dense or sparse. Head not strongly reduced but can be retracted within prothoracic cavity, and antennae on head between eyes. Elytra with 0, 2, interstitial lines; elytral carina absent. No legs or parts thereof swollen and /or curved. LO in V6 only, lacking any elevations or depressions or ridges on V7 except for depressions in posterolateral areas in some species which probably reflect torsion of underlying D–V muscles.

Larva not associated.

Remarks. Pygatyphella (Ballantyne) exists in two clearly defined forms referred to in descriptions as Pygatyphella A or B:

Pygatyphella A: Dorsal colouration usually cryptic, very pale brown with dark markings on pronotum, MS, MN and elytral base and apex; pronotum often with angulate convergence along lateral margins, and small depression often present just anterior to posterolateral corner; lacking irregularities at the corners; posterolateral corners may project beyond median posterior margin; at least two interstitial lines usually well–defined; elytral margins convex or parallel-sided; posterior area of ventrite 7 may reach into LO (which is medially emarginated); apex of MPP rounded, pointed, or squarely or obliquely truncate; MPP lacking median dorsal ridge (except in karimui sp. n.), lacking curved, slightly off-centre posterior ridge on ventral surface of T8; dorsal surface of posterior area of MPP faced with cuticle that is not attached to the ventral surface of V7 and ends just behind the area of muscle attachment posterior to the LO; the anterior margins of this cuticle are reflexed in Pygat. peculiaris ; median area of this facing cuticle lacking longitudinal split; T8 anterolateral prolongations lacking ventral projections at their base; aedeagal sheath not subparallel-sided in basal third; aedeagus L/W usually>3; LL/ML not wide; ML sometimes asymmetrical in a horizontal plane only, never with lateral teeth; anterior margin of LL often asymmetrically produced. Within Pygatyphella A, a group of species similar to Pygat. obsoleta in colour and terminal abdomen morphology is referred to in descriptions as the ‘‘obsoleta complex’’.

Pygatyphella B: Dorsal colouration with orange pronotum (sometimes with median dark mark), and dark brown elytra which may be pale margined except in Pygat. russellia , which is pale dorsally; pronotum never with angulate convergence along lateral margins, or small depression present just anterior to corner; with irregularities at corners; posterolateral corners always rounded obtuse, not projecting strongly if at all beyond median posterior margin; no interstitial lines well–defined; elytral margins convex-sided; posterior area of ventrite 7 never reaching into LO (LO not medially emarginated); apex of MPP rounded or squarely or obliquely truncate; MPP with dorsal longitudinal ridge; dorsal surface of the posterior area of the MPP faced with cuticle that is not attached to the ventral surface of V7 and ends just behind the area of muscle attachment posterior to the LO; the anterior margin of this cuticle is emarginate and the ridge that continues along the ventral surface of the MPP is a continuation of this emargination (e.g. Figs 342–349, especially Fig. 358); ventral surface of T8 with usually curved, slightly off–centre ridge close to posterior margin; T8 outlines in Pygat. limbatipennis , limbatifusca and salomonis characterised in Figs 438 –442; T8 with pale partly membranous ventral projections from the bases of the anterolateral prolongations; aedeagal sheath sternite posterior to the lateral tergite articulations subparallel-sided in basal 1/3 to 1/2; aedeagus L/W<3; LL/ML wide; ML never asymmetrical, always with lateral teeth; anterior margin of LL never asymmetrically produced.