Colophotia Motschulsky 1853

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 : 66-68

Colophotia Motschulsky 1853

Figs 76, 77 , 206−229

Colophotia Dejean 1833: 103 (catalogue name only). Olivier 1885: 367; (in Baer) 1886: 132; 1907: 56; 1910a: 48: 1911a: 102; 1913b: 59. McDermott 1966: 116. Ballantyne, 1968: 106; 1987a: 234; 1987b: 173, 175–177. Ballantyne & McLean 1970: 234.

Colophotia Motschulsky 1853: 51 . Ballantyne & Lambkin 2009: 155, figs 98–105; 2013: 60–64, figs 92–108.

Type species: Lampyris praeusta Eschscholtz 1822 , designated by Motschulsky (1853).

Diagnosis. Colophotia is a S. E. Asian genus with a dense concentration of species in the Philippines. All but one are pale brownish or yellow dorsally usually with black elytral apices, and larvae are terrestrial. Species are never seen in high numbers, usually being taken singly when their flashing pattern is observed (Jusoh, Wong pers. comm.).

Colophotia belongs to a group of nine Luciolinae genera (the others being Australoluciola Ballantyne , Emarginata gen. nov., Inflata Boontop , Medeopteryx Ballantyne , Pteroptyx Olivier , Pyrophanes Olivier , Serratia gen. nov. and Trisinuata Ballantyne ) all of which have pronotal width less than width across the elytral humeri; parallel-sided elytra; aedeagal sheath elongate slender, widest across the middle, with each side of the posterior half of sternite tapering evenly towards a narrow entire apex. With the exception of Inflata Boontop , males of these genera are also characterized by: an elongate slender aedeagus with LL concealed behind the ML (when the specimen is viewed from beneath); the aedeagus of Inflata has a ML swollen in the middle and apically acute LL visible at their bases only from below). Colophotia males are distinguished from all other genera by a combination of the following characters:

1. an elongate slender aedeagus which is often 5 times as long as wide, or longer

2. the aedeagal LL are very short and scarcely separated along their median dorsal length (b/a is approximately 0.3)

3. an elongate slender aedeagal sheath which may be up to 7 times as long as wide

4. accommodating the elongate aedeagus and sheath, the paired anterolateral arms of T8 are longer than the posterior entire portion of T8

5. the anterolateral corners of T7 are prolonged (presumably to provide extra surface area for muscle attachment from the very elongated aedeagal sheath)

6. bipartite LO in V7 has an elongate median carina between the LO halves

7. antennal FS 7–9 are shorter than the remaining FS and FS 9 is usually apically acute

8. there is no MFC, no deflexed elytral apices, and no leg segments are swollen or curved or emarginated

9. the posterior margin of V7 is trilobed, with well-defined and oblique to vertical PLP, and a well-defined and often very large MPP which is apically emarginated, may be grooved along its median ventral length, and is prolonged into dorsally curving hooks.

Females are macropterous and the bursa in species investigated has wide paired plates. Larvae lack laterally explanate tergal margins, the laterotergites are clearly visible from above, and are very similar to described larvae of Pteroptyx spp.

Redescription of Male. Pronotum: not much wider than long; dorsal surface without irregularities in posterolateral areas (1) and longitudinal groove in lateral areas (2); punctation moderately dense (3); anterior margin not explanate (6); lateral margins diverging posteriorly along most of their length (C>A, B); width <humeral width; anterolateral corners rounded obtuse; lateral margins without indentation at mid-point, and sinuosity in either horizontal or vertical plane (15, 16); without indentation in lateral margin near posterolateral corner, and irregularities at corner (17, 18); posterolateral corners angulate, approximately 90° and inclined at 90° to the median line; 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; median area more widely flattened than elsewhere; pronotal width/ GHW 1.4–1.5.

Elytron: lateral margins parallel-sided; apices not deflexed; punctation dense (34, 35), not linear (36), not as large as that of pronotum, nor widely and evenly spaced; epipleuron and sutural ridge extend beyond mid-point almost to apex but not extending around apex, neither thickened in apical half; no interstitial lines; elytral carina absent; in horizontal specimen viewed from below epipleuron at elytral base wide, covering humerus or almost so; viewed from above anterior margin of epipleuron arises level with or 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.

Head: gently depressed between eyes; moderately well exposed in front of pronotum, and capable of partial retraction within prothoracic cavity; eyes with moderate separation beneath at level of posterior margin of mouthpart complex; eyes above labrum moderately to widely separated; 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, always separated by> ASW sometimes>2 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 or beyond. Mouthparts: functional; apical labial palpomere flattened, shaped like an elongate slender triangle (widest at base and L 2– 3x W), with inner edge entire, and at least half as long as apical maxillary palpomere. Antennae 11 segmented; length>GHW up to twice GHW; FS1 subequal in length to, or longer than, pedicel; no FS expanded or laterally produced; all FS including FS 1 longer than wide; FS 7–9 shorter than preceding FS and sometimes much darker in colour, with FS 9 apically acute.

Legs: with inner tarsal claw entire; without MFC; no leg segments swollen, curved or emarginated.

Abdomen: ( Ballantyne 1968 figs 11–14; Ballantyne & McLean 1970 figs d–g; Ballantyne & Lambkin 2009 figs 98–107; 2013 figs 92–108); without cuticular remnants in association with aedeagal sheath (131, 132); no ventrites with curved posterior margins, nor extending anteriorly into emarginated posterior margin of anterior segment (106–108). V6 with LO present, occupying almost ventrite surface. V7 with LO bipartite, occupying more than half the surface of ventrite, and reaching to sides and into the PLP; LO halves separated in middle, sometimes widely so; this area may appear slightly depressed; between LO halves a longitudinal median carina which may be low wide and rounded along its ventral length, or high narrow and acute; posterior margin of V7 trisinuate with a slightly asymmetrical MPP, apex of which is deeply emarginate, MPP as long as wide or longer than wide (L=W or L>W), at least subequal to or longer than PLP, and wider than PLP; MPP often with a deep median emargination margined laterally by strong ridges, either evenly or unevenly margined; MPP prolonged into paired hooks which incline dorsally but do not engulf T8, and may be asymmetrically inclined; hooks may arise from posterior apices of MPP and are visible from beneath, or from a preapical dorsal area when they are not visible from beneath; PLP well defined, either subequal in length to MPP or shorter, and inclined either obliquely or vertically so from below they appear narrower than the MPP. T8 well sclerotised, symmetrical, visible posterior area widest across median area (margins may appear convex sided); median posterior margin strongly emarginate; posterolateral corners may be strongly prolonged and incline dorsally in pinned specimen; not engulfing posterior margin of V7 nor MPP (133, 134), with well-defined median longitudinal trough margined laterally by well-defined ridges which are prolonged anteriorly into long apically rounded flanges; flanges may incline one to the anterior and the other posterior, or both slightly anteroventrally; concealed anterolateral arms of T8 always longer than visible posterior portion of T8, sometimes almost 3 times as long; not laterally emarginated before their origins, dorsoventral expansions absent, expanded only in horizontal plane; without bifurcation of inner margin and ventrally directed pieces (217); lateral margins of T8 not enfolding sides of V7 (190); T7 with prolonged anterolateral corners.

Aedeagal sheath: ( Ballantyne & Lambkin 2009 fig. 103; 2013 fig. 108) symmetrical; very long and narrow up to 7x as long as wide; without well-defined 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; tergite without projecting pieces along posterior margin of T9, anterior margin without transverse band.

Aedeagus: ( Ballantyne & Lambkin 2009 figs 101, 102; 2013 fig. 108) L/W 3/1 or greater, symmetrical with apices of LL not visible from beneath at sides of ML, LL/ML narrow; LL of equal length, much shorter than ML, contiguous along inner dorsal margins with apices very narrow and barely separated (b/ a= 0.3 approximately); dorsal base of LL symmetrical, evenly excavated; ML symmetrical; BP not strongly sclerotised, not hooded, not strongly emarginated along anterior margin.

Female ( Ballantyne & Lambkin 2013 figs 92–98, 104–108). Macropterous. Pronotum: without irregularities in posterolateral areas; punctation moderate to dense; pronotal width less than humeral width; without indentation of lateral margin, irregularities at posterolateral corner; outline similar to that of male. Elytra: subparallel-sided, punctation not as large as that of pronotum, nor evenly spaced; no interstitial lines; elytral carina absent. Legs: no legs or parts thereof swollen and /or curved. Abdomen: LO in V6 only, without any elevations or depressions or ridges on V7; median posterior margin of V7 widely emarginate. Bursa plates ( Ballantyne & Lambkin 2013 figs 94–98, 106) consisting of 2 wide paired plates; median oviduct plate small (reproductive system not investigated in all species).

Larva. (Associated by breeding for C. praeusta and C. brevis only). Terrestrial; elongate, slender spindle shaped ( Ballantyne & Lambkin 2009 fig. 517), of the form of Pteroptyx maipo and several Australian Australoluciola sp. larvae ( Ballantyne & Lambkin 2000; Ballantyne et al. 2011); tergal plates not produced at sides and laterotergites visible from above.

Remarks. Colophotia has been previously characterised in shortened form ( Ballantyne 1968; Ballantyne & McLean 1970; Ballantyne & Lambkin 2013), mainly because it has not been possible to locate all types. However since three species were scored in a matrix that now has over 400 characters of males, females, and larvae (Ballantyne et al. 2016) we believe sufficient information on several distinctive Colophotia species permits us to attempt a generic redescription, given that we have been able to locate a type series for one, and possible syntypes for another two. Of the seven species listed in Table 12 types cannot be located for three. Two species listed by McDermott (1900) viz. C. miranda Olivier , and C. truncata Olivier are treated as species incertae sedis. McDermott’s (1962) incorrect interpretation of the abdomen of C. praeusta was addressed by Ballantyne & Lambkin (2013: 64). We are presently unable to resolve some apparent inconsistencies concerning Luciola lata Olivier 1883: 79 the male of which Olivier appears to have described with a median carina on the last abdominal segment (“ultimo abdominis segment….in medio longitudinaliter carinata” and “dernier segment de l’abdomen…chargé sur son milieus d’une forte carène longitudinale obtuse”). McDermott (1966) synonymised the species with Luciola pallescens Gorham 1880 . From its size lata could well be a male of C. miranda , and both are treated here as species incertae sedis.

Ballantyne & Lambkin (2013) discussed the genus and their reasons for attributing it to Motschulsky. McDermott (1966) had attributed the genus to Dejean as do Bousquet & Bouchard (2013).

Figures

FIGURES 70–77. Key to genera characters 11. 70–72 Serratia sibuyania gen. et sp. nov.; 73–75 Emarginata trilucida gen. nov. et comb. nov.; 76, 77 Colophotia praeusta. 70, 72, 73 dorsal habitus (70 anterior body only); 74 ventral habitus; 71 dissected terminal abdomen ventral; 75 ventrite 7 and elytral apices from below; 76, 77 ventrites 6, 7 from left (76) and below (77). Figure legends: MC median carina; MPP median posterior projection; PLP posterolateral projections.

FIGURES 206−214. Colophotia bakeri syntype male (206−208); syntype female 209, 210, male 211−214. 206, 209 habitus dorsal; 207, 210 habitus ventral; 208 type labels; 211−214 abdomen: 211 dorsal ventrites 6 and 7 only after soaking in 70% ethanol; 212−214 whole abdomen: 212 ventral; 213 right lateral; 214 dorsal.

FIGURES 215−221. Colophotia bakeri male. 215−216 abdomen ventral (ventrites 4−7 only); 216 same left lateral; 217 ven- trites 6 and 7 right lateral (tergite 8 removed); 218−220 tergites 7, 8 with aedeagal sheath and aedeagus within sheath: 218 ventral, 219 dorsal. 220 left lateral; 221 tergite 8 with sheath removed.

FIGURES 222−229. Colophotia brevis type male (223, 224), Pahang male 222, 226−229. 222, 223 habitus ventral; 224 habitus dorsal; 226 abdomen dorsal; 227, 228 tergite 8: 227 ventral; 228 left lateral; 229 aedeagus dorsal.