Creophilus incanus, (KLUG)

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 : 766-768

2. CREOPHILUS INCANUS (KLUG)

( FIGS 1G, 2K, 4F , 16 , 17 )

Staphylinus (Emus) incanus Klug, 1834: 137 . Type locality: Madagascar.

Staphylinus incanus ; Erichson, 1839: 350; Vinson, 1956: 51; Vinson, 1967: 320.

Staphylinus (Creophilus) Coquerelii Fauvel, 1866: 317 . Type locality: Réunion: Bourbon; 1867: 67; 1868: 71 (same as 1867 paper); Vinson, 1956: 51 (synonym of C. incanus ); Lecoq, 1990: 206 (as coquereli, synonym of C. incanus ).

Creophilus incanus ; Fauvel, 1875: 55; Alluaud, 1900: 87; Bernhauer & Schubert, 1914: 398; Lecoq, 1990: 206, 210, fig. 39; Herman, 2001b: 3317.

Creocephalus incanus ; Fairmaire, 1893: 523 (error for Creophilus , see Blackwelder, 1952: 111).

Creophilus Coquereli ; Alluaud, 1900: 88; Bernhauer & Schubert, 1914: 398 (synonym of C. incanus ).

Type material: Staphylinus (Emus) incanus Klug. Lectotype (here designated). ♂, ‘[orange] Type/ incanus Kl. / 5884/ [green] incanus| Kl.| Madag Goud./ [blue] Hist.-Coll. ( Coleoptera )| Nr. 5884| Staphylinus incanus Kl. | Madagasc., Goudot| Zool. Mus. Berlin / [red] SYNTYPUS | Staphylinus | incanus Klug, 1832 | labeled by MNHUB 2004/ FMNH-INS 0000 016 781/ [red] LECTOTYPE | Staphylinus (Emus) | incanus Klug, 1832 | designated by| D. J. Clarke 2008’ (in ZMHB). Specimen missing left metatarsus and right metatarsal claws. Paralectotype. ♀, ‘[orange] Type/ [blue] Hist.-Coll. ( Coleoptera )| Nr. 5884| Staphylinus incanus Kl. | Madagasc., Goudot| Zool. Mus. Berlin / [red] SYNTYPUS Staphylinus | incanus Klug, 1832 | labeled by MNHUB 2004/ FMNH-INS 0000 016 782/ [yellow] PARALECTOTYPE | Staphylinus (Emus) | incanus Klug, 1832 | designated by| D. J. Clarke 2008’ (in ZMHB).

Staphylinus (Creophilus) Coquerelii Fauvel. Syntypes (?; not seen). Vinson (1956) notes ‘type: Réunion, probably in Paris Museum’ . Fauvel did not indicate the number of specimens examined.

Other material examined: 170 specimens. See supporting information, Appendix S1 .

Diagnosis: With characters of the maxillosus -group; integument and elytra uniformly black; antenna distinctly clavate; antennomere 11 convex apically ( Fig. 16B ); basolateral pronotal impressions marked by deep punctule ( Fig. 16G ); basolateral pronotal macroseta absent; elytral vestiture with indistinct whitish- or silvery-grey patch ( Figs 1G, 16H ); tergal chaetotaxic formula 6-6-6-0(2)-4-6.

Description: Measurements ( N = 10♂, 10♀). Forebody length: ♂ 5.8–11.0 mm, ♀ 6.1–8.0 mm. See surpporting Table S 4 for comparison of ranges of male and female ratios. Head. Head weakly trapezoidal and widest posteriorly to subrectangular, particularly in females; HW/ HL = 1.43–1.70; basal margins sparsely setose; dorsal punctation sparse; eyes large ( EYL / HL = 0.48–0.65), dorsolateral in large males, more lateral in females and smaller males, not protruding, lateral margins of head visible in dorsal view only in large males, obscured by eye in smaller specimens; HL 1/ HL 2 much greater in females than males (♂ = 1.21–2.40, ♀ = 2.60–3.25); antennae as in Figure 16B , distinctly clavate; antennomeres 1–6 brownish-black, 7–11 greyish-black, 9–10 narrowed anteriorly, thickened posteriorly; apex of antennomere 11 convex medially; each pair of apical setae moderately widely separated, one on each side of apex; mandibles as in Figure 16A , longer than head in large males, subequal to head in females and small males ( ML / HL ♂ = 0.88–1.48, ♀ = 0.94–1.06), T 1 largest, T 3 hardly visible in dorsal view. Thorax and abdomen. Pronotum ( Fig. 16G ) slightly transverse ( PW / PL = 1.07–1.24), moderately to distinctly narrowed posteriorly; PL 1.24–1.56 ¥ ESL; basolateral margins very shallowly emarginate, hind angles distinct, obtuse; disc surface faintly sinuous laterally, with sparse, black, peripheral setae and very sparse short vestiture on lower anterolateral declivities; basolateral impressions sparsely setose, accentuated by deep punctule; scutellum with uniformly black vestiture; elytra uniformly black to brownish-black, vestiture sparse, with poorly delimited central white patch ( Fig. 16H ); wings fully developed, clear yellowish-brown, without black spot in medial field between MP 3 and MP 4 veins; vestiture of pterothorax mostly whitish, that of legs mostly black; dorsal abdominal vestiture mostly black but with variable maculation of whitish vestiture concentrated on tergites and parasclerites III – V ( Fig. 1G); vestiture of sternites III – VI mostly whitish, interrupted by more or less circular black patches laterally, vestiture on sternites VII –IX mostly black; whitish abdominal vestiture frequently with bluish-grey tinge; abdominal tergite VII with well-developed palisade fringe. Male genitalia. Aedeagus as in Figure 16F ; median lobe apex narrowly subacuminate ( Fig. 16C ); paired apicolateral sclerites (as) articulated to internal edge of median orifice, not fused to it. Paramere as in Figure 16D . Internal sac inverted as in Figure 16F , everted as in Figure 16E ; ventral sclerite (vs) deeply notched apically ( Fig. 16J ); ventral process (vp) short, with ventromedian spiculose strip (vr) shorter than ventral sclerite (vs). Female internal genitalia. Internal female genitalia as in Figure 16I ; vaginal plate (vp) broadly rounded to truncate posteriorly, with median sclerotized strip (ss); surface of vaginal fold ( Fig. 16I , vf) with embedded minute tooth-like scales (as in Fig. 9M). Chaetotaxy. Basolateral pronotal macrosetae absent; elytral discal series with 3–4 macrosetae; tergal chaetotaxic formula = 6-6-6-0(2)-4-6, inner lateral macrosetae absent from tergite VII, tergite VI usually without macrosetae, outer lateral macrosetae occasionally present.

Variation: Five specimens from Réunion and three from Madagascar have mostly black vestiture on elytra, tergites, and ventral side, with only paired groups of white setae in marginal fringes of tergites III – VI. Aedeagi of males from Réunion and Mauritius were identical to those from Madagascar.

data). Phenology: February–December. Vinson (1956) records one specimen ‘on decaying Clemora beetles’ ( Scarabaeidae : junior synonym of Phyllophaga Harris ), another ‘trapped in an Aristolochia flower’. He notes the species is ‘most probably a predator of dipterous larvae in carrion’.

Remarks: A female specimen from Réunion in Bernhauer’s collection ( FMNH) bears the label ‘ nigerrimus Typus’ in Bernhauer’s handwriting. Another ‘ nigerrimus ’ specimen (syntype?) is possibly in NMW ( A. F. Newton, pers. comm.). Further collecting on Réunion may reveal subspecific differences from Madagascan specimens.

Comparison: Creophilus incanus resembles C. galapagensis and C. maxillosus villosus in having the whitish dorsal abdominal vestiture concentrated on tergites IV– V. It is distinguished from both most easily by the convex apex of the terminal antennomere and the posteriorly thickened preceding two antennomeres ( Fig. 16B ).

Distribution ( Fig. 17 ): Madagascar, Mauritius, Mayotte ( Alluaud, 1900), and Réunion.

Biology and ecology: Rare in collections. Two specimens were taken from cow dung. Habitat: unknown. Altitude: 800 m (only one specimen with elevation

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 16. Creophilus incanus, details of morphology (setae of mandibles, antenna, and pronotum omitted). A, mandibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown; anterior is to the left); C, median lobe apex, ventral; D, paramere, dorsal; E, apical part of median lobe with fully everted internal sac, right lateral; F, aedeagus with details of inverted internal sac, right lateral (parameral setae omitted); G, pronotum (small male); H, right elytron (only whitish setae shown); I, female internal genitalia, ventral (surface of vf with minute teeth as shown for vp of C. maxillosus villosus in Fig. 9M); J, ventral sclerite of internal sac, ventral (lower) and apical (upper). Abbreviations: T1, T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars = 1 mm.

Figure 17. Distribution of all georeferenced specimens of C. incanus in Madagascar and Mayotte (circles). This species is also known from Réunion and Mauritius islands.