Creophilus flavipennis Clarke, Dave J., 2011

Clarke, Dave J., 2011, Testing the phylogenetic utility of morphological character systems, with a revision of Creophilus Leach (Coleoptera: Staphylinidae), Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 163 (3), pp. 723-812 : 772-775

4. CREOPHILUS FLAVIPENNIS (HOPE) COMB. NOV., STAT. NOV.

( FIGS 1I, 4E , 10A , 21 , 22 )

Staphylinus flavipennis Hope, 1831: 32 . Type locality: Nepal; Herman, 2001b: 3502 (as nomen dubium).

Creophilus villipennis Kraatz, 1859: 67 . Type locality: ‘ India boreali’; Fauvel, 1875: 54; Bugnion, 1911: 135–152, figs 1–10; Bernhauer & Schubert, 1914: 399; Cameron, 1932: 261; 1933 d: 392; Scheerpeltz, 1933: 1413; Cerruti, 1951: 17; Scheerpeltz, 1976: 92; Smetana & Davies, 2000: 38; Herman, 2001b: 3325; Smetana, 2004: 672; new synonymy.

Emus violaceus Fauvel, 1878a: 248 . Type locality: ‘ Moluques, Bouro’ ; 1879: 94; new synonymy.

Emus villipennis ; Fauvel, 1878a: 248; Fauvel, 1879: 94 (as E. villipenni ).

Emus insularis Fauvel, 1879: 94 . Type locality: ‘Batcian’; new synonymy.

Creophilus insularis ; Bernhauer & Schubert, 1914: 399; Bernhauer, 1926: 349; Scheerpeltz, 1933: 1412; Herman, 2001b: 3326.

Creophilus violaceus ; Bernhauer & Schubert, 1914: 399; Herman, 2001b: 3326.

Type material: Staphylinus flavipennis Hope. Lectotype (here designated). ♀, ‘[circular with red border] Type| H. T./ flavipennis Hope. / Hardwicke | Bequest./ flavipennis. Hope.| 4241/ FMNH-INS 0000 016 808/ [red] LECTOTYPE | Staphylinus | flavipennis Hope, 1831 | designated by| D. J. Clarke 2008’ (in BMNH). Only one specimen was in Hardwicke’s collection.

Creophilus villipennis Kraatz. Lectotype (here designated). ♂, ‘1307/ [yellow] India sept.| Bacon leg./ Coll. Kraatz/ [red] Syntypus / coll. DEI| Eberswalde/ coll. DEI| Müncheburg/ FMNH-INS 0000 016 810/ [red] LECTOTYPE | Creophilus | villipennis Kraatz, 1859 | designated by| D. J. Clarke 2008’ (in DEI). Specimen and terminalia mounted on card; left antennomeres 7–11 and protarsal claw missing; right elytron with pinhole close to scutellum. Paralectotypes (3). All with labels ‘[red] Syntypus / Coll. Kraatz/ coll. DEI| Eberswalde/ coll. DEI| Müncheburg/ [yellow] PARALECTOTYPE | Creophilus | villipennis Kraatz, 1859 | designated by| D. J. Clarke 2008’: ♂, ‘[yellow] India sept.| Bacon leg./ Creophilus | villipennis| Kr./ FMNH-INS 0000 016 809’ (in DEI); ♀, same collecting data and ‘FMNH-INS 0000 016 811’ (in DEI); ♀, ‘Decan/ Saunders/ Creophilus | villipennis Kr. / FMNH-INS 0000 016 812’ (in DEI).

Emus violaceus Fauvel. Holotype (by monotypy). ♂, ‘ Molucche | Buru X | Beccari 1875/ Typus / violaceus| Fauvel / Emus | violaceus FvL.| ♂ / Museo Civico | di Genova / FMNH-INS 0000 016 807/ [red] HOLO- TYPE| Emus | violaceus Fauvel, 1878 | teste D. J. Clarke 2008’ (in MCSN). Fauvel’s (1878a) original description was based on ‘ Un seul ♂ ’.

Emus insularis Fauvel. Lectotype (here designated). ♂, ‘Batchian/ V. insularis| FvL./ [red text in black box] Ex-Typis/ R.I.Sc.N.B. 17.479| Coll. et det. A. Fauvel/ FMNH-INS 0000 016 805/ [red] LECTO- TYPE| Emus | insularis Fauvel, 1879 | designated by| D. J. Clarke 2008’ (in IRSNB). Specimen with right mandible extended. Paralectotype. ♀, ‘Ceram/ Coll. et det. A. Fauvel| Creophilus | insularis| Fauv.| R. I.Sc.N.B. 17.479/ [red text in black box] Ex-Typis/ FMNH-INS 0000 016 804/ [yellow] PARALECTO- TYPE| Emus | insularis Fauvel, 1879 | designated by| D. J. Clarke 2008’ (in IRSNB); specimen missing right middle leg.

Other material examined: 241 specimens. See supporting information, Appendix S1.

Diagnosis: With characters of the maxillosus -group; eyes occupying most of sides of head (e.g. Fig. 21A ); elytra variably yellowish and/or brownish-black; vestiture of head, pronotum, most of elytra, legs, apicolateral regions of tergites IV– V, apical fringes of tergites III – VI and most of ventral side whitish-grey to golden-brown; tergal chaetotaxic formula = 6-6-6-6-4-6.

Description: Measurements (N = 10♂, 10♀). Forebody length: ♂ 6.1–10.3 mm, ♀ 6.7–8.9 mm. See supporting Table S4 for comparison of ranges of male and female ratios. Head. Head ( Fig. 21A ) transverse, only slightly widened posteriorly, suborbicular in smaller specimens; HW/HL = 1.25–1.53; basal margins densely setose; dorsal punctuation fine, moderately dense; eyes very large ( EYL /HL = 0.59–0.81), protruding, occupying sides of head, lateral margins of head obscured by eye in dorsal view, HL1/HL2 large, much greater in females than in males (♂ = 1.92–8.00, ♀ = 7.67–12.00); antennae as in Figure 21C , moderately clavate; antennomeres 1–6 entirely black, 7–11 greyish-black, 11 frequently reddish-brown; apex of antennomere 11 slightly to moderately emarginate medially, each pair of apical setae widely separated, one on each side of apex; mandibles as in Figure 21B , longer than head in large males, shorter in females ( ML /HL ♂ = 0.89–1.47, ♀ = 0.75–0.96), right mandibular teeth subequal in size, T 1 reduced in smaller specimens. Thorax and abdomen. Pronotum ( Fig. 21G ) slightly transverse ( PW / PL = 1.07–1.22), moderately to strongly narrowed posteriorly; PL 1.12– 1.35 ¥ ESL; basolateral margins very shallowly emarginate in front of indistinct hind angles; with dense and long whitish peripheral setae and coarse long whitish vestiture on anterolateral declivities; scutellum with yellowish-brown vestiture; elytra uniformly blackish-brown to almost yellowish, or blackishbrown anteriorly fading to yellowish-brown posteriorly, often with faint violet sheen; vestiture dense, evenly golden-yellow or darker with indistinctly lighter setae at middle (as in Fig. 1I); wings fully developed, clear yellowish-brown, without black spot in medial field between MP 3 and MP 4 veins; pterothorax and legs covered with golden or whitish vestiture; dorsal abdominal vestiture mostly black, with golden brown or whitish maculation restricted to apicolateral regions of tergites (and parasclerites) IV– V, and apical fringes of tergites III – VI; vestiture of sternites III – VI mostly golden-brown, VII and VIII mostly black in females, in males each with medial patch of golden vestiture; abdominal tergite VII with well-developed palisade fringe. Male genitalia. Aedeagus as in Figure 21F ; median lobe apex elongate, deeply incised proximal to lateral recurved teeth, the latter blunt, reduced ( Fig. 21F , lit); with paired apicolateral sclerites (as) articulated to internal edge of median orifice, not fused to it; median orifice internally grooved on dorsal side and with unique internal teeth ( Fig. 21J , arrow). Paramere as in Figure 21D . Internal sac inverted as in Figure 21F , everted as in Figure 21E ; ventromedian spiculose strip (vr) much shorter than ventral sclerite (vs), the latter partly divided at middle ( Fig. 21I ). Female internal genitalia. Internal female genitalia as in Figure 21H ; vaginal plate (vp) with median sclerotized strip (ss), produced posteriorly into unique apically bifid process; vaginal fold with two large pigmented regions dorsal to vaginal plate ( Fig. 21H , arrow). Chaetotaxy. Elytral discal series with 6–8 macrosetae; procoxa with two macrosetae at distal margin; tergal chaetotaxic formula = 6-6-6-6-4-6, inner lateral macrosetae absent from tergite VII.

Variation: Philippine and eastern Indonesian specimens are overall pitchy black, distinctly darker, with almost black elytra and whitish-grey vestiture at the middle of the elytra, gradually fading yellowish posteriorly. These specimens have a distinctly greater contrast between integument and vestiture coloration and match the type specimens of C. insularis (Fauvel) . Indian and South East Asian specimens have distinctly lighter (almost uniformly yellowish) elytra and elytral vestiture.

Comparison: Creophilus flavipennis can generally be distinguished from other species of the C. maxillosus - group by the enormous eyes alone ( Figs 1I, 21A ), but the vestiture patterning is also diagnostic. The unique male and female genitalia are of particular interest for diagnosing this species, as these characters are constant throughout the range of the species.

Distribution ( Fig. 22 ): Oriental, widespread: Nepal, India, Sri Lanka, Myanmar, China (one specimen), Laos, Thailand, Vietnam, Malaysia, Indonesia, Philippines, and east to Tanimbar Islands ( Indonesia). Smetana (2004) records it additionally from Kashmir in India. Herman (2001b) records C. insularis (= C. flavipennis ) from New Guinea (evidently based on Scheerpeltz, 1933), but I could not confirm that it occurs there.

Biology and ecology: Many specimens have been taken at light ( UV, mercury vapour), others from Malaise traps and various carrion. Habitat: lowland rainforest to alpine habitats. Altitude: 61–4500 m. Phenology: throughout the year. Other biology and life-history characteristics are unknown. Larvae and pupae are unknown.

Remarks: Staphylinus flavipennis was listed by Herman (2001b: 3502) as a nomen dubium. Hope (1831) provided a valid description: ‘Ater, capite thoraceque nigro, elytrisque villoso aureis abdomineque nigro. Long. lin. 7 ½; lat. 2.’, agreeing both with the lectotype of S. flavipennis and the type series of C. villipennis . The name S. flavipennis is therefore a senior subjective synonym and should be adopted as the valid name because (1) under Article 23.1 it has priority over C. villipennis , and (2) the name C. villipennis , as a junior synonym, does not meet the usage requirement of Article 23.9.1.2 ( ICZN, 1999).

Fauvel (1878a) described the large male holotype of Emus violaceus as follows: ‘denticulo laterali postico angulum ipsum formante’, referring to the basolateral pronotal denticle typical of large males of the C. maxillosus -group, and ‘capite inter antennas anguste profundeque bifossulato’, referring to two accentuated anterior tentorial pits between the antennal fossae – an occasional aberration seen in other species. In describing Emus insularis, Fauvel (1879) primarily relied on allometric characters of both sexes: ‘capite multo minore, in utroque sexu simili... mandibulis duplo brevioribus... linea basale collo profunde transversim impressa... thorace... haud perspicue denticulato’. All are typical of small C. flavipennis specimens. In all other respects C. insularis and C. violaceus type specimens are typical of C. flavipennis .

Five specimens from the Philippines were identified as ‘ Creophilus semipubescens ’. This is presumably a Bernhauer manuscript name as I have seen no published reference. One specimen in NMW ( FMNH-INS 0000 106 557) bears a ‘cotypus’ label, another in ZMHB ( FMNH-INS 0000 016 806) bears a ‘paratypus’ label. Three specimens in FMNH were identified by Bernhauer as C. semipubescens ( FMNH-INS 0000 016 564–565, 016 791). The first of these specimens bears a label in Bernhauer’s handwriting ‘ Creophilus semipubescens Brh Typ. ’.

Figures

Figure 4. Male copulatory pieces of selected species in the Creophilus-complex, dorsal view. In each figure the large paired dark structures are the sclerotized arms of the copulatory piece. A, C. variegatus; B, C. maxillosus maxillosus; C, C. m. villosus; D, C. galapagensis; E, C. flavipennis; F, C. incanus; G, C. oculatus; H, C. erythrocephalus; I, C. imitator; J, C. lanio; K, C. albertisi; L, Hadrotes crassus; M, Thinopinus pictus; N, Hadropinus fossor; O, Liusus hilleri. Abbreviations: iedg, apical inner edge of arms of copulatory piece; por, parallel oblique ridges of copulatory piece (see also Fig. 10); xbar, ‘cross-bar’ of appendix. Additional abbreviations as in Figure 3. Scale bar = 1 mm.

Figure 10. Scanning electron micrographs of copulatory pieces. A, Creophilus flavipennis; B, C. maxillosus villosus; C, C. erythrocephalus. Abbreviations as in Figures 3 and 4. Scale bars = 0.1 mm.

Figure 21. Creophilus flavipennis, details of morphology (setae of head, mandibles, antenna, and pronotum omitted). A, head, dorsal (small male); B, mandibles, dorsal (bases omitted); C, right antenna, dorsal (only apical setae shown); D, paramere, dorsal; E, apical part of median lobe with fully everted internal sac, right lateral; F, aedeagus with details of internal sac, right lateral (parameral setae omitted); G, pronotum (small male); H, female internal genitalia, ventral – arrow indicates subsclerotized or pigmented regions; I, ventral sclerite of internal sac, ventral (lower) and apical (upper); J, median lobe apex, ventral – arrow indicates internal tooth flanking the groove of the median orifice. Abbreviations: T1, T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars: I = 0.1 mm; rest = 1 mm.

Figure 22. Distribution of all georeferenced specimens of C. flavipennis (circles).