Creophilus maxillosus, (LINNAEUS)

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 : 760-761

1. CREOPHILUS MAXILLOSUS (LINNAEUS)

( FIGS 1F, 2E–G, 3C, D, I, P, 4B, C , 5A , 9C–F, L, M, 10B , 13 , 14 , 15,19B, 24)

Staphylinus maxillosus Linnaeus, 1758: 421 . Type locality: ‘Europa’; Herman, 2001b: 3317 (as C. maxillosus ); Navarrete-Heredia et al., 2002: 322 (as C. maxillosus ). Type specimens: see C. maxillosus maxillosus .

Staphylinus anonymus Sulzer, 1761: 17 , fig. 49. Type locality: ‘Europa’; Erichson, 1839: 348 (synonym of S. maxillosus s.l.); Herman, 2001b: 3321 (synonym of C. maxillosus s.l.); Smetana, 2004: 671 (synonym of C. m. maxillosus ).

Staphylinus tertius Schaeffer, 1766 : plate 20, fig. 1, 1. Type locality: ‘Ratisbon’ (not seen); Erichson, 1839: 348 (synonym of S. maxillosus s.l.); Herman, 2001b: 3321 (synonym of C. maxillosus s.l.; nomen nudum, Article 11.4).

Staphylinus balteatus DeGeer, 1774: 18 . Type locality not cited; Fabricius, 1781: 334 (synonym of S. maxillosus s.l.); Herman, 2001b: 3321 (synonym of C. maxillosus s.l.); Smetana, 2004: 671 (synonym of C. m. maxillosus ).

Staphylinus fasciatus Füessly, 1775: 21 . Type locality: Switzerland; Gravenhorst, 1802: 3 (synonym of S. maxillosus s.l.); Herman, 2001b: 3322 (synonym of C. maxillosus s.l.); Smetana, 2004: 672 (synonym of C. m. maxillosus ).

Staphylinus nebulosus Geoffroy, 1785: 165 . Type locality: Paris; Olivier, 1795: (42): 10 (synonym of S. maxillosus s.l.); Herman, 2001b: 3322 (synonym of C. maxillosus s.l.); Smetana, 2004: 672 (synonym of C. m. maxillosus ).

Staphylinus bicinctus Mannerheim, 1843: 229 . Type locality: ‘insula Sitkha’; Solsky, 1870: 261 (synonym of C. arcticus ); Horn, 1879: 200 (synonym of C. villosus ); Ganglbauer, 1895: 415 (synonym of C. maxillosus s.l.); Herman, 2001b: 3323 (synonym of C. maxillosus s.l.; preoccupied by Staphylinus bicinctus Rossi, 1792 ); Navarrete-Heredia et al., 2002: 322 (synonym of C. maxillosus s.l.); Smetana, 2004: 671 (synonym of C. m. maxillosus ); primary junior homonym of Staphylinus bicinctus Rossi, 1792 .

Staphylinus orientalis Motschulsky, 1858b: 67 . Type locality: ‘Iles Kouriles’; Ganglbauer, 1895: 415 (synonym of C. maxillosus s.l.); Herman, 2001b: 3324 (synonym of C. maxillosus s.l.); Smetana, 2004: 672 (synonym of C. m. maxillosus ).

Creophilus fulvago Motschulsky, 1860: 120 . Type locality: ‘ Mongolie et Chine boréale’; Ganglbauer, 1895: 415 (synonym of C. maxillosus s.l.); Herman, 2001b: 3324 (synonym of C. maxillosus s.l.); Smetana, 2004: 672 (synonym of C. m. maxillosus ).

Creophilus maxillosus var. pulchellus Meier, 1899: 99 . Type locality: ‘Hamburg’; Blackwelder, 1943: 448 (synonym of C. maxillosus s.l.); Herman, 2001b: 3324 (synonym of C. maxillosus s.l.); Smetana, 2004: 672 (synonym of C. m. maxillosus ).

Diagnosis: With characters of the maxillosus -group; integument, including elytra, uniformly black; antennae moderately clavate; apex of antennomere 11 emarginate ( Fig. 13F ); elytra with white vestiture arranged into well-demarcated (but variable) central transverse fascia ( Figs 1F, 13B ); tergal chaetotaxic formula 6-6-6-2(4)-4(6)-8(10+).

Description: To previous detailed descriptions ( Blackwelder, 1936; Dajoz & Caussanel, 1969) I add measurements and body ratios (listed under each subspecies), chaetotaxic characters, and full description of male and female genital characters. Head. Head slightly wider anteriorly in large males, subtrapezoidal and wider posteriorly to suborbicular in females and smaller males; basal margins densely setose; dorsal punctation moderately dense; eyes large, dorsolateral, lateral margins of head visible in dorsal view (not obscured by eye), HL1/HL2 greater in females than males; antennae as in Figure 13F , moderately clavate; antennomeres 1–6 brownish-black, 7–11 greyish-black; 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 13A, T 1 and T 3 subequal in length, in large males T1 much larger than T2 and T3. Thorax and abdomen. Pronotum ( Figs 2G, 13E ) moderately to distinctly narrowed posteriorly; basolateral margins very shallowly emarginate, hind angles indistinct; with dense and long peripheral setae and coarse long black or whitish-grey vestiture on anterolateral declivities; scutellum with indistinct anterior band of whitish-grey vestiture; elytra uniformly black, densely setose, with characteristic welldelimited fascia of whitish-grey setae at middle; wings fully developed, clear yellowish-brown to black, without black spot in medial field between MP3 and MP4 veins; ventral thoracic vestiture entirely whitish-grey or black ( Fig. 14 ); anterior and middle legs with black vestiture, hind coxae, trochanters, and femora with white vestiture; abdominal vestiture arranged into characteristic pattern of whitish-grey or golden brown maculation dorsally, and mostly or entirely white on sternites and parasclerites IV–VI, sternite III, and sternites VII–IX; extensive whitishgrey vestiture interrupted dorsally by a large medial and two smaller anterolateral black patches, and ventrally by more or less circular black patches laterally; abdominal tergite VII with well-developed palisade fringe. Male genitalia. Aedeagus as in Figures 3C, 13D ; paired apicolateral sclerites articulated to internal edge of median orifice. Paramere as in Figure 13C . Internal sac inverted as in Figure 13D , everted as in Figure 3C, D; with short ventral process (vp), ventromedian spiculose strip slightly longer than the box-like ventral sclerite ( Figs 3I, 13H , vs). Female internal genitalia. Internal genitalia as in Figures 5A , 13G ; vaginal plate produced into two broad lobes posteriorly, and with median sclerotized strip (ss); vaginal fold (vf) completely membranous; spermatheca (sp) elongate sacculate, with distinct duct connecting to bursa copulatrix (bcx). Chaetotaxy. Elytral discal series with 4–8 macrosetae, frequently with additional macroseta medially disposed from series; tergal chaetotaxic formula = 6-6-6-2(4)-4(6)- 8(10+), outer laterals present or absent and inner laterals absent on tergite VI, inner laterals present or absent on tergite VII; tergite VIII often with inconsistent additional macrosetae.

Variation: Coloration and the patterning of body vestiture are the only notable geographically variable characters, and with the exception of North American and Palearctic forms (discussed under each subspecies) appears to be continuous. These forms exhibit more or less identical male and female genitalia, with only subtle differences in the copulatory piece/ appendix that may be significant for reproductive isolation but which require more detailed study. Much of the synonymic diversity is attributable to variations in vestiture coloration and to allometric characters of the head and thorax.

Comparison: Creophilus maxillosus is distinctive in habitus ( Fig. 1F) and most similar to C. incanus . It is reliably distinguished from this species by the concave apex of the apical antennomeres, and from all species by the definite whitish-grey elytral fascia, and dorsal pattern of body vestiture.

Distribution: Widespread throughout the northern hemisphere; apparently absent from austral regions (except as adventive), South East Asia, and the East Indies. See under each subspecies for details.

Biology and ecology: Creophilus maxillosus is probably the best-studied rove beetle species. Herman (2001b) lists many papers on its biology, morphology, and life history. The following papers provide details of: general biology and life history ( Abbott, 1938; Fichter, 1949b; Kramer, 1955; Greene, 1996); morphology and anatomy ( Talbot, 1928; Blackwelder, 1936; Dajoz & Caussanel, 1969); diet ( Fichter, 1948); chemical defence ( Jefson et al., 1983); olfaction ( Abbott, 1936); and immature morphology ( Voris, 1939; Paulian, 1941). Kirkpatrick & Olson (2002) note increases in predatory activity at night. Moon & Kajii (1997) observed fully grown larvae feeding on fly maggots beneath rabbit carcasses, where they are presumably able to evade the cannibalistic tendencies of adults ( Abbott, 1937, 1938) by burrowing into the substrate (as observed by Fichter, 1948).

Dynamics of arrival and departure at carrion have been recorded frequently in the forensic entomology literature (e.g. Early & Goff, 1986; Goff, 1991; Anderson & VanLaerhoven, 1996; Moon & Kajii, 1997; Richards & Goff, 1997; Sánchez Piñero, 1997; Avila & Goff, 1998). Summarizing these papers, adults arrive early at a carcass during the ‘bloat’ and ‘decay’ stages ( Payne, 1965; length of stages varies depending on environmental conditions), remaining there up to 13 days post-mortem. Creophilus maxillosus prefers open disturbed habitats over dense forest, and is typically found in synanthropic situations. Habitat: seashore, grassland, prairie, mixed forest, savannah, woodland, swamp, alpine meadow, etc. Altitude: sea level to 3640 m. Phenology: throughout the year, tending to be most common from late spring through autumn in northern USA.

Remarks: Measurement and ratio data for this species are provided for each subspecies, below. I could not determine details and/or depositions for type material of the following synonyms: Staphylinus anonymus Sulzer , Staphylinus tertius Schaeffer , Staphylinus balteatus De Geer , Staphylinus fasciatus Füessly , Staphylinus nebulosus Geoffroy , Staphylinus bicinctus Mannerheim (syntypes in ZMHB?), and C. maxillosus var. pulchellus Meier (holotype by monotypy). Syntypes (?) of S. orientalis and C. fulvago are presumably in ZMUM but according to A. Gussakov (ZMUM) these specimens are in poor condition and their identity cannot be confirmed. I list these nine names here because I could not confirm their status as synonyms of the nominate subspecies, which for most is nevertheless suggested by their type localities and synonymic history. All other synonyms and type material are listed under the subspecies to which they apply. Previous authors (e.g. Horn, 1879; Hatch, 1957) have treated C. bicinctus as a synonym of C. m. villosus or C. arcticus (see Hatch, 1938: 148). Motschulsky (1858b, 1860) notes the similarity of S. orientalis to C. m. maxillosus , noting that it differs only in (trivial) dimensions of the head and thorax. According to the original descriptions, C. fulvago Motschulsky and C. m. var. pulchellus Meir may represent forms similar to the C. ciliaris holotype, characterized by extensive golden or reddish-brown vestiture over most of the body. Most authors prior to 1938 treated C. villosus ( Gravenhorst, 1802) as either a separate species or a variety of C. maxillosus ( Herman, 2001b) . Hatch (1938) was the first to treat C. villosus as a subspecies of C. maxillosus , although Blackwelder (1943) later synonymized it with C. maxillosus , which was followed by Herman (2001b). The subspecific division of C. maxillosus has been readopted in some recent works ( Smetana & Davies, 2000; Smetana, 2004), including this one, because although superficial, the differences between them are more or less discrete, and may be supported by some internal sac differences that require further study.

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 5. Female genitalia, details of morphology (all but E, ventral view). A, Creophilus m. villosus, internal genitalia; B, C. erythrocephalus, internal genitalia; C, C. huttoni, internal genitalia; D, C. oculatus, gonocoxites and styli (setae omitted from left side); E, Thinopinus pictus, abdominal segments VIII–X, dorsal (vestiture omitted); F, T. pictus, internal genitalia; G, Staphylinus caesareus, internal genitalia; H, Platydracus maculosus, internal genitalia; I, Hadrotes crassus, right gonocoxites and stylus (vestiture omitted) – arrow indicates membranous articulation; J, T. pictus, right gonocoxite. Abbreviations: bcx, bursa copulatrix; fl, flagellum (note that the complete structure is not shown); gc, gonocoxite; gcm, gonocoxal macroseta; gc1, gonocoxite I, gc2, gonocoxite II; mg, median groove; pls, paired lateral sclerites of vaginal plate; sc, sclerite of bursa copulatrix; sd, spermathecal duct; sp, spermatheca; ss, sclerotized strip of vaginal plate; sty, gonostyle; vf, vaginal fold; vp, vaginal plate. Scale bars = 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 13. Creophilus maxillosus, details of morphology (setae of mandibles, antenna, and pronotum omitted). C. m. maxillosus (A–G); C. m. villosus (H). A, mandibles, dorsal (bases omitted); B, right elytron (only whitish setae shown) C, paramere, dorsal; D, aedeagus with details of inverted internal sac, right lateral (parameral setae omitted); E, pronotum (small male); F, right antenna, dorsal (only apical setae shown); G, female internal genitalia, ventral H, 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: H = 0.1 mm; rest = 1 mm.

Figure 14. Ventral vestiture patterning of A, C. maxillosus maxillosus and B, C. m. villosus. Scale bar = 10 mm.