Creophilus, LEACH, 1819

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 : 755-757

publication ID	https://doi.org/ 10.1111/j.1096-3642.2011.00725.x
publication LSID	lsid:zoobank.org:pub:FBFE9195-BE04-4AFE-9417-6E38BCE6AB84
persistent identifier	https://treatment.plazi.org/id/039B414F-195C-FFD9-FEC4-FAF249A5F8C4
treatment provided by	Valdenar
scientific name	Creophilus
status

Treatment

GENUS CREOPHILUS LEACH View in CoL

Creophilus Leach, 1819: 172 View in CoL . Type species: Staphylinus maxillosus Linnaeus, 1758 View in CoL , fixed by original designation according to action of Commission in Opinion 546 ( ICZN, 1959). Blackwelder (1952: 111) listed synonymic homonyms and variant spellings for Creophilus View in CoL .

Saprophilus Streubel 1839: 136 View in CoL ; Blackwelder 1952: 344 (synonym of Staphylinus View in CoL ). Type species: Staphylinus maxillosus Linnaeus, 1758 View in CoL , fixed by monotypy.

Adult diagnosis: Within the Creophilus -complex, the following characters are exclusive to Creophilus : antennae clavate with antennomeres 7–10 much wider than long (e.g. Fig. 13F View Figure 13 ) and only 7–11 densely pubescent; left mandible with distinct ventral basolateral ridge ( Fig. 2C, vbr); protibiae with two spines on inner face ( Fig. 9E, arrows); sternacostal ridge broadly arcuate medially ( Fig. 2E, sr); mesothoracic peritremes enlarged medially, with posterior edge produced into point ( Fig. 2E, ptm); mesoventral intermesocoxal process truncate or broadly angulate, with mesocoxae widely separated ( Fig. 2F, mip); female with gonostyle articulated subapically on broadly rectangular or subcolumnar second gonocoxite ( Fig. 5D View Figure 5 , sty).

Description: Habitus: Generally large (10–28 mm) black or partly colourful rove beetles (e.g. Fig. 1A–J), with (five spp., e.g. Fig. 1F–J) or without (seven spp., e.g. Fig. 1A–E) variegated whitish-grey or yellowishbrown vestiture. Both sexes are strongly polymorphic in size, males generally larger, with larger heads and longer mandibles (both allometric). Head ( Fig. 2A, B). Head trapezoidal to subrectangular; strongly transverse; shorter than pronotum; narrower than pronotum in females, slightly wider than pronotum in males; mostly glabrous dorsally (disc). Frons and vertex coarsely but sparsely punctured; or impunctate; punctures when present less than half diameter of macrosetal puncture. Antennal fossae separated from anterior margin of head by sclerotized strip ( Fig. 2B, atf). Dorsum of neck sparsely punctate to impunctate, punctures when present as above. Configuration of head ridges as in Figure 2A, B: nuchal ridge generally incomplete dorsomedially, close to infraorbital ridge laterally; dorsal and ventral basal ridges obliterated laterally by anastomosing setiferous punctures; postmandibular ridge absent; gular sutures narrowly separate; basal gular impression not demarcated by anterior transverse ridge; infraorbital ridge present; ventral basal ridge on postgena separated from postoccipital suture, terminating at gular sutures. Antennae (e.g. Fig. 13F View Figure 13 ). Antennae 11-segmented; clavate to subincrassate; longer than head; insertions exposed dorsally. Antennomeres 1–6 shining, not pubescent; 7–11 densely pubescent; 1–3 elongate; 1 longer than 2 and 3 together; 2–3 subequal, dilated distally; 4 globular; 5–10 increasingly transverse (10 widest), somewhat dorsoventrally flattened; 11 narrower than preceding, dorsoventrally compressed apically forming distinct ridge (except C. rekohuensis ), and bearing two pairs of prominent setae. Mouthparts. Labrum ( Fig. 2B) short and transverse; broadly incised mesially; partly sclerotized, and with numerous macrosetae. Mandibles falciform; each with distinct mandibular blades ( Fig. 2D, mb); with dorsal basolateral ridge bordered by setiferous groove removed from edge ( Fig. 2D, dbl). Right mandible with two dorsal teeth ( Fig. 2D, T 1, T 2) and one ventral tooth ( Fig. 2D, T 3). Left mandible with two teeth in same plane ( Fig. 2D); with distinct ventral basolateral ridge bordered laterally (especially in large specimens) by distinct groove to receive right mandible ( Fig. 2C, vbr). Maxillae as in Figure 2H. Labium as in Figure 2I. Mentum short, transverse; with two pairs of anterolateral macrosetae. Ligula minutely incised apically. Thorax. Pronotum transverse; strongly transversely convex; glabrous (on disc), shining; without definite microsculpture on disc; with anterolateral declivities, sides, and base sparsely to rather densely setose. Basolateral pronotal margins faintly to distinctly emarginated. Pronotal hypomeron visible in lateral view; glabrous. Pronotal postcoxal process large and triangular; hyaline; fimbriate anteriorly. Superior pronotal marginal line separate from inferior line, never joining it anteriorly; deflexed beneath anterior pronotal angles, thus pronotum anteriorly visible in ventral view ( Fig. 2E). Inferior pronotal marginal line continuous with anterior pronotal margin ( Fig. 2E). Prosternum ( Fig. 2E) fused with hypomeron (pronotosternal suture absent). Basisternum broadly rounded; sparsely setose; acarinate; without distinct intercoxal depressions. Sternacostal ridge broadly arcuate medially. Furcasternum glabrous; almost vertical; with distinct fovea; posteriorly with broad groove connecting posterolateral fossae ( Fig. 2E, plf). Mesothoracic peritremes very large and acute; approximate mesially; with reinforced poriferous membrane behind ( Fig. 2E, ptm). Scutellum densely punctured; setose. Elytra. Elytra together subrectangular; longer than pronotum (when measured from humerus to apex); with epipleura faintly impressed anteriorly; with subapical surface depressed laterally. Apices obliquely truncate; forming obtuse angle with suture. Disc and epipleuron evenly setose, but without micropubescence; without microsculpture (but frequently rugosely sculptured). Humeri sparsely punctate and setose to impunctate and at least partly glabrous; often with distinctive shining humeral callus (e.g. Fig. 1A). Wings. Wings well developed and functional (except C. rekohuensis and C. huttoni ); with MP3, MP4, and CuA veins completely separate ( Fig. 33N View Figure 33 ). Mesoventrite acarinate; with short broadly transverse intermesocoxal process ( Fig. 2F, mip). Metaventrite broadly convex; with marginal carinae of mesocoxal acetabula obsolete posteromedially ( Fig. 2F, mac); with deep premetacoxal groove ( Fig. 2F, pmg). Legs. Tibiae multispinose, with numerous longer bristles interspersed (e.g. Figs 2K, 9E). Tarsi pentamerous; with bisetose empodia. Prothoracic legs robust. Protibiae ( Fig. 2K) moderately expanded distally; spinose only posteriorly; with two prominent spines on ventral face (opposing profemur when leg adducted; Fig. 9E). First four protarsomeres transversely bilobed in both sexes; clothed ventrally with tenent setae. Mesocoxae widely separated. Abdomen. Abdomen densely setose, with setae moderately appressed, and sparser or absent anteriorly and around spiracles. Basal tergal margins slightly sinuous laterally, straight medially ( Fig. 2L, btf). All segments with distinct basal transverse fold dorsally and ventrally, extending onto inner parasclerites ( Fig. 2L, btf). Tergites and sternites arcuate at middle; with shallow transverse lateral impressions. Tergite I setose laterally; without prototergal glands, but sometimes with shallow acetabula laterally. Tergite II as in Figure 2J; with short internal anterolateral flanges (af); with paired muricate fossae (f). Second abdominal spiracle large; situated in membrane outside of tergite. Apical margin of tergite VII with distinct white palisade fringe in all macropterous species (e.g. Fig. 1A; reduced or absent in flightless species). Paired abdominal defensive gland reservoirs present (for detail see Dajoz & Caussanel, 1969: fig. 10A). Male. Head as wide as or slightly wider than maximum pronotal width. Sternite VIII deeply subtriangularly emarginate (e.g. Fig. 23K View Figure 23 ). Apices of tergite X and sternite IX subtruncate to shallowly emarginate, and sternite IX with elongate basal portion ( Fig. 3Q). Aedeagus (e.g. Fig. 3A–C). Aedeagus bilaterally symmetrical; with paramere facing left in repose (in abdomen). Apex of median lobe with paired apicolateral sclerites (as); deeply incised laterally, proximal to lateral recurved teeth (lit; except in C. albertisi ). Basal bulb with large dorsal sclerite, and paired ventral sclerites separated medially by membrane. Paramere spatulate; narrower than median lobe; shaft widest apically or preapically; not extending beyond apex of median lobe; emarginate apically (except C. variegatus ); with several apicolateral setae; dorsoventrally movable at base (i.e. not fused to median lobe; except C. albertisi ). Internal sac with structures as labelled in Figure 3A–D; bilaterally symmetrical, everting symmetrically along axis of median lobe ( Fig. 3C, D). Membranous parts of internal sac, especially distally around copulatory piece, armed with minute tooth- or hair-like scales ( Fig. 4 View Figure 4 , sh). Female. Head as wide as or narrower than maximum width of pronotum. Sternite VIII entire. Apices of tergite X (except C. variegatus ) subtruncate to shallowly emarginate. Gonocoxites I and II ( Fig. 5D View Figure 5 , gc) separated by membrane. Distal gonocoxites broadly rectangular or subcolumnar; with gonostyle (sty) articulated subapically (i.e. articulation point not visible in ventral view). Internal female genitalia (e.g. Fig. 5A–C View Figure 5 ). Vaginal cavity with structures as labelled in Figure 5A–C View Figure 5 . Spermatheca unsclerotized, tubular to only very slightly expanded distally. Chaetotaxy ( Fig. 6 View Figure 6 ). All named macrosetae, when present, represented by single macrosetae, never clusters (excluding procoxal and posterior elytral ‘patches’: pp, pt), and are illustrated for a hypothetical member of the Creophilus complex in Figure 6 View Figure 6 . (Head) Anteromarginal (am) anterior to antennal insertion; temporal and occipital at posterior margin of head (tm, oc); ventroscapal (vs) same size as and indistinguishable from other setae, or else absent. (Prothorax) Pronotum at most with only antero- and basolateral (an, bp); without anterior pronotal group (ap). Procoxa with mesal patch (pp); protrochanter with at least one (tr); profemur with none. (Pterothorax) Elytra with at least one basohumeral (bh); elytral discal series comprising two to eight (ds); posterior elytral patch with at least three (pt); mesoventrite with broadly arcuate indistinct row (mr); mesocoxae ventrally, and hind legs, with none. Metaventrite with one posteromedially (pm). (Abdomen) Abdominal tergals placed at apical margins on tergites III–VI, nearer middle on tergites VII and VIII (ol, il, mm); parasclerites with none; outer laterals (ol) present on all tergites.

Distribution: Worldwide, but apparently absent from sub-Saharan Africa. Several species have become established outside their native ranges.

Biology and ecology: The biology and life history of C. maxillosus are well known (see references in Herman, 2001b). Comparable mouthpart morphology and specimen data suggest that all species (except possibly C. huttoni ) are primarily predators of dipteran larvae at carrion, although they may occasionally be found in other decaying substrates, or take other prey. Facultative cannibalism has also been observed ( Fichter, 1948). They are not normally necrophagous and prefer carrion in the early stages of decomposition ( Fichter, 1949a) when blowfly larvae are most plentiful. A preference for blowfly larvae has been demonstrated for C. m. villosus adults and larvae in feeding experiments ( Abbott, 1934; Fichter, 1948; Greene, 1996), and orientation to the carrion resource is achieved using long-range olfactory cues ( Abbott, 1934, 1936, 1938). One consequence of the carrion microhabitat association is that adults can be collected en masse using carrion-baited pitfall traps. Single squidbaited traps ( Newton & Peck, 1975) left for several weeks can collect hundreds of specimens. In contrast, larvae and pupae are evidently never collected using this method, but can be collected in the vicinity of carrion (pers. observ.). This suggests that larvae hatch from eggs laid directly on or near the carrion following arrival of adults from elsewhere. The likely requirement that the life cycle be completed in situ only at carrion or similar substrates might explain the absence of larvae from baited traps. Pre-adult development takes up to 35 days in C. maxillosus (details in Kramer, 1955), so newly emerged adults will probably disperse due to local conditions unsuitable for feeding and development (carrion dried out, low maggot abundance). Arrival and departure times for adults at carrion are highly variable, depending on environmental conditions affecting decomposition rates ( Archer, 2004).

Nomenclature and taxonomy: Samouelle (1819) published the new generic name Creophilus , for which he designated Staphylinus maxillosus Linnaeus, 1758 as type species. This designation was later formally established in Opinion 546 ( ICZN, 1959); Leach can be inferred as author of the genus by similar logic to that of Opinion 1722 ( ICZN, 1993). Since 1819, Creophilus has been considered either as a valid genus or a synonym or subgenus of Staphylinus (see Smetana & Davies, 2000). Nordmann (1837) described C. badiipennis , C. erithacus , and C. chloris , but these were removed to the xanthopygine genus Torobus Herman ( Herman, 2001a) . Staphylinus luctuosus Blanchard, 1842 was transferred to Creophilus by Bernhauer & Schubert (1914), but Newton (1996) transferred it to Xanthopygus Kraatz after examination (and designation) of the lectotype.

The nomenclature of C. maxillosus -group species is marked by proliferation of specific, infraspecific, and infrasubspecific synonyms, reflecting variability within some species. Recent works (e.g. Smetana & Davies, 2000; Smetana, 2004) treat C. m. maxillosus (Linnaeus) and C. m. villosus (Gravenhorst) as subspecies, and this classification is maintained here. Types of 12 of the 20 subjective C. maxillosus synonyms were studied, and most synonyms have been allocated to the appropriate subspecies. Several names were also determined to be unavailable. A Galapagos Islands species, frequently referred to as C. m. villosus , is here described as the new species C. galapagensis .

Nomenclatural issues in the C. erythrocephalus - group are fewer, involving confusion between C. erythrocephalus , C. lanio , and C. oculatus , or disagreement in their rank. However, both Fauvel (1875) and Steel (1949) clearly distinguished them at the specific level. For the New Zealand region, Herman (2001b) listed only one valid species ( C. oculatus ), but Emberson (1998) suggested that C. oculatus tends to be replaced by another species in the south of South Island. This distinctive beach-inhabiting species ( C. huttoni ), known also to Newton (1985) and Hammond (2000), was synonymized by Gourlay (1950) without comment. Moreover, Emberson (1998) also reported a species from the New Zealand Chatham Islands indicating that it was ‘very similar or identical to the one [ C. huttoni ] found in the southern South I. (SL) and Stewart I.’ Creophilus huttoni is here reinstated, and C. rekohuensis is described from the Chatham Islands.