Atyphella Olliff, 1890

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 : 34-40

Atyphella Olliff, 1890

( Figs 9–11, 35, 78, 79, 83–85, 118 –179)

Atyphella Olliff, 1890:645 . Lea, 1909:110. Olivier, 1909b:lxxxii (Partim); 1910b:40; 1911:171; 1913:417 (Partim). Ballantyne, 1987b:172, 175–77, 181, 183–5. Calder, 1998:176 (Partim). Ballantyne & Lambkin, 2000:22 (Partim); 2006 (Partim):30.

Luciola (Luciola) Laporte. Sensu McDermott, 1964:45 ; 1966:99.

Luciola (Atyphella) (Olliff) . Ballantyne, 1968:108. Ballantyne & McLean, 1970:23.

Type species. Atyphella lychnus Olliff, 1890 .

Diagnosis. Males of Australian species, and A. scabra , with anterior hypomeron flat to neck, and posterior area flattened and strongly adpressed; non-Australian species lack the flat anterior hypomeron; epipleuron either continuing around elytral apex as a ridge ( Figs 9–11, 79, 83, 118, 119, 122–125, 127–129, 169, 170) or not; frons-vertex junction rounded, or angulate ( Figs 84, 85); mouthparts functional or not; apical segment of labial palpi either strongly flattened, with inner margin dentate, or ovoid with inner margin entire; aedeagal sheath with posterior area of sternite emarginate on its right side from attachment of tergite. Distinguished most obviously from Aquilonia and Gilvainsula by their distinctive pale dorsal colouration; lacking the asymmetrical tergite 8 of Asymmetricata ; differing from Convexa and Magnalata which have distinctive dark elytra often with pale margins; lacking the dorsal colour pattern, shape of aedeagus and hooded basal piece of Lloydiella , and lacking the arched V7 with LO retracted to anterior half or less characteristic of Pygatyphella . Females ( Figs 130–136) either macropterous or exhibiting varying degrees of brachelytry. Larvae with laterally explanate tergal margins concealing laterotergites from above.

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

Pronotum ( Figs 9, 11, 84, 85, 118, 122–129, 137–139, 142, 144, 147, 150–152, 154, 157 –159,166–169, 177, 178) usually wider across posterior area than rest, subparallel-sided in inconspicua , some immaculata ; pronotal width usually greater than humeral width, subequal to humeral width in aphrogeneia , guerini, inconspicua , lewisi and similis . Anterolateral corners rounded obtuse; lateral margins in anterior half divergent, or subparallel-sided in inconspicua ( Fig. 125); lateral margins in posterior half either subparallel-sided in inconspicua and some immaculata ( Fig. 127), or diverge along their length in similis some dalmatia sp. n. ( Fig. 147) and some immaculata , or diverge then converge with rounded convergence (( Figs 78, 84, 123, 129, 147, 158, 159, 166, 169, 177); lacking indentation at mid-point, or sinuousity in either horizontal or vertical plane; subparallel-sided margins straight; lacking indentation in lateral margin near posterolateral corner, and irregularities at corner; posterolateral corners rounded or angulate; rounded corners obtuse, <90°; in dalmatia sp. n.; angulate corners approximately 90° and inclined at right angles, or obliquely, to median line; posterolateral corners either not projecting as far as median posterior margin ( aphrogeneia ), or projecting as far as or beyond it and separated from it by a shallow emargination except in some aphrogeneia .

Hypomera closed. Median area of hypomeron not elevated vertically; anterior, and posterior areas of hypomeron narrowly to widely flat to side of head in Australian species and A scabra ( Fig. 84, 178); anterior area of hypomeron not flattened in remainder; posterior area of hypomeron widely flattened in remainder ( Fig. 84) except for some aphrogeneia and inconspicua where it is narrow; flattened areas closely adpressed except for some aphrogeneia ; pronotal width/ GHW index 1.6 except for inconspicua (1.2), and 1.4–1.5 in some aphrogeneia and immaculata .

Elytron punctation usually dense ( Figs 123, 125, 126–129, 137, 138, 144, 150, 153, 154, 167, 177), sparse in aphrogeneia ( Fig. 118); punctation not linear, not as large as pronotum, nor widely and evenly spaced; apices not deflexed; epipleuron and suture extend beyond mid-point, may extend as ridge around apex without any further expansion of either, or apical half of both epipleuron, sutural ridge, and elytral apex expanded in aphrogeneia ; ridge around apex of elytron may be visible from beneath; 0, 2, 3 or 4 interstitial lines, inner two exceed height of suture in scabra only; elytral carina absent; in horizontal specimen viewed from beneath epipleuron at elytral base wide, covers humerus from below; in specimen viewed from above arises anterior to posterior margin of MS; epipleuron developed as lateral ridge along most of length; sutural margins approximate along most of their length in closed elytra except in aphrogeneia where they diverge in apical half; lateral margins parallel-sided or convex.

Head moderately to strongly depressed between eyes; moderately exposed in front of pronotum, not capable of complete retraction within prothoracic cavity except in lychnus , monteithi and scintillans ; eyes close to moderately separated beneath at level of posterior margin of mouthpart complex; eyes above labrum close (moderately separated in some dalmatia sp. n.,); frons-vertex ‘junction’ rounded or acute ( Fig. 84), with median elevation in conspicua only ( Ballantyne & Lambkin 2000 Fig. 2f); posterolateral eye excavation not strongly developed, not visible in resting head position; antennal sockets on head between eyes, contiguous or separated by up to ASW;,clypeolabral suture present, flexible, not in front of anterior eye margin when head viewed with labrum horizontal; outer edges of labrum reach inner edges of closed mandibles. Mouthparts functional or not; apical segment of labial palpi non–lunate, either strongly flattened, shaped either like a wide to narrow triangle, with inner edge irregular ( guerini ) or dentate; or narrow triangle with L>W; or not flattened and ovoid, as long as or longer than wide. Antennae 11 segmented, sometimes less; length subequal to GHW – twice GHW; no segments flattened, shortened, or expanded; pedicel not produced; FS1 not shorter than pedicel; in very short antennae FS may be subequal in length and width.

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 usually entire, bipartite in scabra (Fig. 178, 179; restricted to anterolateral plaques), kirakira sp. n. ( Fig. 142, 143), and possibly lewisi where it is not possible to determine its extent; either reaching both to sides and posterior margin in olivieri and testaceolineata , or reaching only sides; entire LO in V7 occupies most of V7; bipartite LO occupies> half V7; neither anterior nor posterior margin of entire LO in V7 emarginate; posterior half of V7 not arched or swollen, muscle impressions not visible in this area; LO present in V6, entire and occupying almost all V6 except in scabra where it is restricted to anterolateral plaques. MPP present, symmetrical, apex rounded, (except narrow and pointed, or rounded in scabra ), entire, not laterally compressed, short, not inclined dorsally nor engulfed by T8 apex, lacking dorsal ridge and median longitudinal trough. V7 lacking median carina, median longitudinal trough, anteromedian depression on face of LO, incurving lobes, pointed projections, median ‘dimple’, or reflexed lobes and PLP (inconsistent development of this area in scabra could be postmortem change). T7 lacking prolonged posterolateral corners. T8 not strongly sclerotised, often subparallel-sided, symmetrical, W=L or W>L, visible posterior area does not narrow abruptly, lacking prolonged posterolateral corners, median posterior emargination, median posterior projections, not inclined ventrally nor engulfing posterior margin of V7 nor MPP, extending conspicuously beyond posterior margin of V 7 in scabra only; T8 ventral surface lacking flanges, lateral depressed troughs, median longitudinal trough, asymmetrical projections, median posterior ridge; concealed anterolateral arms of T8 either absent (in scabra ), or very short and narrow, or elongate but not as long as visible posterior portion of T8, and narrow; not laterally emarginated before their origins, not expanded dorsoventrally, apices lacking bifurcation of inner margin, bases lacking ventrally directed pieces; lateral margins of T8 not enfolding sides of V7.

Ballantyne and Lambkin 2000, 2001, 2006 Present composition

Luciolinae Luciolinae

Luciola s str. (Clade 1) includes: Missimia gen. n.

• L. italica (type species) Presently unassigned (Node 1 Luciola cruciata – L. owadai )

• L. dejeani Presently unassigned (Node 2 L. leii – L. ficta )

• Lampyroidea sp. Presently unassigned (Node 4 L. substriata J – L. substriata A)

• Bourgeoisia sp. Pygoluciola Wittmer (Node 9 Pyg hamulata – Pyg wittmeri )

• L. (Hotaria) parvula Photuroluciola Pic

Luciola (Clade 2) Curtos Motschulsky (Node 37 okinawanus – costipennis )

• Luciola sp. (Australian species) Presently unassigned (Node 35 L. australis– L. pupilla )

• Pteroptyx Pteroptyx Olivier sensu latu (Node 28 P. platygaster– P effulgens )

• Pyrophanes Colophotia Dejean (Node 24 concolor – plagiata )

• Colophotia Pyrophanes Olivier (Node 21 beccarii – quadrimaculata )

Atyphella Olliff Pteroptyx s. str. (Node 18 L. sp MFC– truncata )

Pygoluciola Wittmer Luciola Laporte s str. (Node 6 dejeani – antipoda )

Curtos Motschulsky Aquilonia gen. n. (Node 5 costata )

Gilvainsula gen. n. (Node 47 messoria – similismessoria ) Magnalata gen. n. (Node 55 carolinae – rennellia )

Asymmetricata gen. n.. (Node 52 circumdata – ovalis )

Convexa gen. n.. (Node 51 wolfi )

Lloydiella gen. n. (Node 50 uberia – wareo )

Atyphella Olliff (Node 78 aphrogeneia – similis )

Pygatyphella (Ballantyne) (Node 3 wisselmerenia – nabiria )

Aedeagal sheath ( Figs 121, 145, 148, 170, 171, 181–183) never> 4 times as long as wide; lacking paraprocts; asymmetrical in posterior area where sheath sternite emarginated on right side from point of attachment of tergite; sternite not angulate on L or R sides, not subparallel-sided, posterior margin apically entire and rounded except for short rounded median projection in scabra (Figs 182, 183); not emarginated on either side preapically; anterior half of sternite broad, apically rounded; tergite lacking lateral arms extending anteriorly at sides of sheath sternite; tergite not subdivided, lacking projecting pieces along posterior margin of tergite 9; anterior margin tergite 9 lacking transverse band.

Aedeagus ( Ballantyne & Lambkin 2000 Fig. 5 in part) Figs 120, 145, 146, 148, 149, 160–165, 172–174, 184, 185) L/W 3/1 or shorter; LL lack lateral appendages, visible from beneath at sides of ML, LL/ML moderate; LL of equal length, subequal to or slightly longer or shorter than ML, diverging or not along inner margins, separated there by> half their length; LL base width not = LL apex width which is subequal to or narrower than ML; LL apices not expanded horizontally; dorsal base of LL symmetrical, not excavated, if median margin prolonged then either broadly rounded, broadly truncate, pointed entire, or pointed, medially emarginate; LL lacking lateral hairy appendages along outer ventral margins, which are not produced preapically, nor narrowly on inner apical margin, and not obliquely truncate along their preapical inner margins except for palauensis and dalmatia sp. n.; apices of LL not inturned, may be out–turned with apex rounded; lacking projection on left LL only; inner margins lacking slender leaf-like projection. ML symmetrical, lacking paired lateral teeth and tooth to left side, not strongly arched, apex not shaped like 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 ( Figs 130–136, 155, 156). Pronotum lacking irregularities in posterolateral areas; punctation moderate to dense, sparse in lewisi and conspicua ; pronotum> humeral width in macropterous females; lacking indentation of lateral margin, and irregularities at posterolateral corner. Elytral punctation not as large as pronotum nor evenly spaced. Atyphella females of four forms ( Figs 130–136): 1. Macropterous, capable of flight; head of winged female form. 2. Fore wings cover most if not all of (gravid) female abdomen; hind wings shortened, may be about ¾ as long as fore wings ( flammans and lychnus ) ( Figs 130, 131), head of wingless female form, pronotum with laterally divergent margins in at least anterior half. 3. Fore wings cover body but hind wings absent ( atra ); head, eyes and pronotal shape as for 2. 4. Brachelytral females with very short or vestigial hind wings ( Figs 132–136), head and eyes same form as 2, pronotum subparallel-sided in conspicua, inconspicua and lewisi , pronotal margins diverge posteriorly in scintillans and similis . 0, 2, 3 or 4 interstitial lines (0, 2 or 4 in macropterous females; 3 in atra which lacks hind wings); 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.

Larva ( Ballantyne & Lambkin 2000 Figs 7, 12, 15). Terrestrial; tergal plates sclerotised to margins, lateral tergal margins explanate, finely margined or thickened, may be ridge-like, covering laterotergites from above; arrangement of plates on ventral aspect of thorax and abdomen like that described ( Ballantyne & Lambkin, 2000; Ballantyne & Menayah, 2002). Protergum L=W or W>L in aphrogeneia and guerini ; tubercles sometimes present along anterior margin, posterolateral corners rounded; median line sometimes with ridged margins; lateral margin of protergum thickened in olivieri , guerini ; punctures in anterior half of terga 2–10 sometimes larger than rest; posterolateral corners of terga 1–8 rounded entire, of tergum 12 either produced narrowly or not; median posterior margins of terga 1–11 lacking either rounded or pointed projections; lacking brush of hairs from apex of tibiotarsus; mandibles lacking inner teeth; antennal segment 3 short, sense cone adjacent to segment 3 short, wide; terrestrial, with laterosternites on abdominal segments 1–8 bearing spiracles.

Remarks. Atyphella was restored to generic level by Ballantyne (in Calder 1998), its status confirmed by cladistic analysis, and revised and redescribed from its Australian examples by Ballantyne and Lambkin (2000). The internal composition was further expanded in Ballantyne and Lambkin (2001, 2006).

Ballantyne and Lambkin (2000) included 23 species in Atyphella Olliff , 16 Australian and seven from the island of New Guinea. Atyphella s. str. as defined here comprises 24 species mainly from Australia and includes 14 of those 16 Australian species, and one of the New Guinean species ( leucura ), and is expanded to include A. testaceolineata , and two new species. Its present composition and the new assignments for species removed from Atyphella are shown in Table 4.

As Australian species# were dealt with in detail in Ballantyne and Lambkin (2000) they are treated here in an abbreviated fashion.