Creophilus variegatus, MANNERHEIM

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 : 775-778

5. CREOPHILUS VARIEGATUS MANNERHEIM

( FIGS 1J, 2C, D, 4A , 23 , 24 )

Creophilus variegatus Mannerheim, 1830: 20 . Type locality: ‘Brasilia ad Rio Janeiro’; Mannerheim, 1831: 434; Nordmann, 1837: 21; Laporte de Castelnau, 1840: 173; Gemminger & Harold, 1868: 575; Fauvel, 1875: 54; Lynch Arribalzaga, 1884: 129; Lüderwaldt, 1911: 417; 419; Bernhauer & Schubert, 1914: 399; Scheerpeltz, 1933: 1413; Herman, 2001b: 3325.

Staphylinus variegatus ; Dejean, 1821: 21 (nomen nudum: Article 12.1); Erichson, 1839: 351; Blanchard, 1842: 76; Murray, 1870: 57. 77.

Emus variegatus ; Fauvel, 1877: xxvii.

Type material: Creophilus variegatus Mannerheim. Lectotype (here designated). ♀, ‘ ♀ / Silberm/ Brasilia/ [red] LECTOTYPE | Creophilus | variegatus| Mannh.| designated by| A. Newton 1978/ Mus. Zool. H: fors| Spec. typ. no. 15232| Creophilus | variegatus Mannh. / FMNH-INS 0000 016 813/ [red] LECTOTYPE | Creophilus | variegatus Mannerheim, 1830 | designated by| D. J. Clarke 2008’ (in MZHF). Newton’s lectotype designation has not been published (A. F. Newton, pers. comm.).

Other material examined: 333 specimens. See supporting information, Appendix S1.

Diagnosis: With characters of the maxillosus -group; antennae subincrassate; apex of antennomere 11 convex ( Fig. 23B ); pronotum strongly constricted basally with sharply delimited hind angles ( Fig. 23H ); elytral epipleura yellowish; with extensive pattern of white and golden brown vestiture on pronotum, elytra, and abdomen ( Fig. 1J); tergal chaetotaxic formula 6-6-6-4(6)-6-8.

Description: Measurements (N = 10♂, 10♀). Forebody length: ♂ 6.4–9.9 mm, ♀ 6.6–8.3 mm. See supporting Table S4 for comparison of ranges of male and female ratios. Head. Head slightly wider anteriorly in large males, subtrapezoidal (wider posteriorly) to suborbicular in females and smaller males; HW/HL = 1.25– 1.56; basal margins setose; dorsal punctation coarse, moderately dense; eyes large ( EYL /HL = 0.48–0.70), dorsolateral in large males, more lateral in females and smaller males, lateral margins of head visible in dorsal view only in large males, obscured by eye in smaller specimens, HL1/HL2 greater in females than males (♂ = 1.54–3.00, ♀ = 3.20–5.00); antennae characteristic ( Fig. 23B ), subincrassate; antennomeres 1–3 brownish-black, 4–11 variably yellowish-brown and black; antennomeres 9 and 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 Figures 2C, D and 23A , longer than head in large males, shorter in females ( ML /HL ♂ = 0.95– 1.48, ♀ = 0.89–0.96), T 3 larger than T 1 and T 2. Thorax and abdomen. Pronotum ( Fig. 23H ) slightly transverse ( PW / PL = 1.07–1.22), with sides subparallel to slightly convex anteriorly, distinctly constricted posteriorly, and hind angles sharply delimited; PL 1.23–1.50 ¥ ESL; vestiture of anterolateral declivities and sides sparse, whitish-grey and brown; vestiture of basal margin golden-brown, dense, and 2¥ as long as setae on anterior angles, with central setae radiating outwards; basolateral impressions broad and deeply set, with dense golden-brown setae; vestiture of scutellum whitish-grey anteriorly, brown posteriorly; disc of elytra brown-black, epipleura yellowish; vestiture of elytral surface dense, with patches of whitishgrey, brown, and golden setae forming complex pattern, posterior-most setae forming loose golden apical band ( Figs 1J, 23G shows white setae only); apical setae 2¥ length of discal setae; hind wings fully developed, clear yellowish-brown, without black spot in medial field between MP 3 and MP 4 veins; pterothorax and legs covered with whitish and golden vestiture; dorsal abdominal vestiture arranged into characteristic pattern of whitish-grey, golden, and brown maculation ( Fig. 1J); vestiture on all sternites mostly whitish, with irregular black patches laterally; abdominal tergite VII with well-developed palisade fringe. Male genitalia and secondary sexual characters. Sternite VII distinctly emarginate medioapically ( Fig. 23K ). Aedeagus as in Figure 23F ; median lobe apex long, pointed ( Fig. 23M ); paired apicolateral sclerites (as) fused to sides of median lobe. Paramere as in Figure 23C ; with apex entire; and with setae along entire length. Internal sac inverted as in Figure 23F , everted as in Figure 23E , paired basoventral spiculose regions absent; with additional sclerotized bar ( Fig. 23J ) connecting ventral sclerite (vs) with base of copulatory piece (cp); copulatory piece sclerites mostly fused along midline, without membranous sheath ( Fig. 4A ). Female internal genitalia. Apex of tergite X narrowly rounded ( Fig. 23L ). Internal female genitalia as in Figure 23I ; vaginal plate (vp) rounded posteriorly, without median sclerotized strip. Chaetotaxy. Basiantennal and basisternal macrosetae absent; elytral discal series with 4–6 macrosetae; mesocoxae with one macroseta at distal margin; tergal chaetotaxic formula 6-6-6–4(6)-6-8, inner laterals present or absent on tergite VI.

Comparison: Creophilus variegatus is easily distinguished from other species of the C. maxillosus -group by the body vestiture patterning ( Fig. 1J), antennae ( Fig. 23B ), and sharply delimited basal pronotal angles ( Fig. 23H ). Males are unusual within the genus in having the apex of sternite VII distinctly emarginate medially ( Fig. 23K ) and the parameral apex entire ( Fig. 23C ).

Distribution ( Fig. 24 , triangles): Endemic to South America: Argentina, Bolivia, Brazil, Chile (one record), Paraguay, Peru (one record), Uruguay.

Biology and ecology: Lüderwaldt (1911) records the species from carrion, and Lynch Arribalzaga (1884) additionally reports predation of calliphorid maggots. Habitat: unknown. Altitude: sea-level to 3000 m. Phenology: throughout the year. Other biology and life-history characteristics are unknown. Larvae and pupae are unknown.

Remarks: Dejean (1821) listed Staphylinus variegatus Dej. , attributing the name to himself, but did not provide a description. Mannerheim (1830) published the name Creophilus variegatus Dej. , attributing the name to Dejean, but also provided the first description so authorship is attributed to him. Mannerheim (1830) did not indicate the number of specimens examined.

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 23. Creophilus variegatus, details of morphology (setae of mandibles, antenna, pronotum, and abdominal structures omitted. A, mandibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown; anterior is to the left); C, paramere, dorsal; D, ventral sclerite of internal sac, ventral (lower) and apical (upper); 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, right elytron (only whitish setae shown); H, pronotum (small male); I, female internal genitalia, ventral; J, ventral sclerite of internal sac with attached sclerotized bar (bar), ventral; K, male abdominal sternites VII and VIII; L, female abdominal tergite X; M, median lobe apex, ventral. Abbreviations: T1, T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars: D and J = 0.1 mm; rest = 1 mm.

Figure 24. Distribution of all georeferenced Creophilus specimens from South America. C. variegatus (triangles), C. erythrocephalus (squares), C. maxillosus maxillosus (circles).