Abstract Abstract
An “Orthrius-group” of genera is proposed, and defined to include Aphelochroa Quedenfeldt, 1885; Caridopus Schenkling, 1908; Dozocolletus Chevrolat, 1842; Gyponyx Gorham, 1883; Languropilus Pic, 1940; Orthrius Gorham, 1876; Pieleus Pic, 1940; Xenorthrius Gorham, 1892; plus three new genera Neorthrius gen. n., Nonalatus gen. n. and Pseudoastigmus gen. n. A phylogeny of the 11 constituent Orthrius-group genera (analysis of 22 morphological characters using Clerus Geoffroy as the out-group taxon was performed with TNT v1.1) is proposed. Four genera are synonymised: Burgeonus Pic, 1950, syn. n. (with Aphelochroa Quedenfeldt, 1885); Brinckodes Winkler, 1960, syn. n. and Quasibrinckodes Winkler, 1960, syn. n. (both with Dozocolletus Chevrolat, 1842); and Dedana Fairmaire, 1888, syn. n. (with Orthrius Gorham, 1876). The genera Falsoorthrius Pic, 1940 and Mimorthrius Pic, 1940 are transferred from Clerinae to the subfamily Tillinae.
Keywords: Cleridae, genus-group, synonymy, phylogeny
Introduction
The checkered beetles (Cleridae and Thanerocleridae) contain approximately 3600 described species, which are classified into seven subfamilies (Lawrence and Newton Jr. 1995) and involve 303 genera. By far, the Clerinae is the most specious subfamily with approximately 45% of the species of the family. Checkered beetles are largely tropical insects with an approximate faunal distribution as follows: 1030 species in the Afrotropics, 840 in the Neotropics, 690 in the Orientalis, 510 in the Australis and 490 in the Palaearctis (Gerstmeier 2000).
The higher classification of the Cleridae has undergone considerable categorical oscillations (Opitz 2002, 2010). Several landmark publications of (Crowson (1955, 1964, 1966, 1970) form the basis for a modern classification of Cleroidea, while some nomenclatural amendments were made by Lawrence and Newton Jr. (1995). More recently, significant contributions dealing with suprageneric taxa include the elevation of Thaneroclerinae (Kolibáč 1992, 2004) and Metaxina Broun, to family rank (Kolibáč 1992, 2004), the proposition of two subfamily classifications (Kolibáč 1997, Opitz 2010) plus revisions of the genera and species of Epiphloeinae (Opitz 1997, 2004, 2005, 2006, 2007, 2008a, 2008b, 2008c), the genera of Hydnocerinae (which included a tribal classification for that subfamily)(Kolibáč 1998) and the Australian Korynetinae (Kolibáč 2003). Nevertheless, some discontinuities are obvious and not all changes made at the subfamily-level are universally accepted among cleridologists. From a world viewpoint, much remains to be done with clarification of generic concepts and zoogeographic relationships at supraspecific levels (Opitz 2002). In our opinion, Opitz’s (2010) concept of 12 subfamilies seems to result in the best system.
The Clerinae is the largest of all subfamilies of the Cleridae and the most difficult in which to define generic limits (Chapin 1924). Furthermore, the paucity of clearly defined morphological gaps among these genera renders their generic delimitation very difficult. A paper dealing with genera related to Clerus Geoffroy (Gerstmeier 2002) represents an initial step in clarifying generic limits within Clerinae. After an extensive review of Indo-Australian clerid material, a generic concept of clerine genera such as Clerus Geoffroy, 1762, Omadius Laporte, 1836, and Stigmatium Gray, 1832 became apparent and resulted in a preliminary concept of “Clerus-series” (Gerstmeier 2002).
A recent revision of the genus Xenorthrius Gorham (Gerstmeier and Eberle 2010) represents besides Mawdsley’s (1994) revision of the genus Aphelochroa the second in a series of papers dealing with the genera of a so-called “Orthrius-group”. In the Xenorthrius revison 11 species were transferred from Orthrius to Xenorthrius, and 22 new species were described, so that the genus Xenorthrius now includes 50 species (from 20 species formerly listed in Corporaal 1950). The aim of the present paper is to define the characters for a generic group, to determine those genera constituting the Orthrius group and examine the relationships among those genera. The following genera have been taken into consideration: Aphelochroa Quedenfeldt, 1885, Caridopus Schenkling, 1908, Dozocolletus Chevrolat, 1842, Gyponyx Gorham, 1883, Languropilus Pic, 1940, Neorthrius gen. n., Orthrius Gorham, 1876, Pieleus Pic, 1940, Nonalatus gen. n., Pseudoastigmus gen. n., Xenorthrius Gorham, 1892, Falsoorthrius Pic, 1940 and Mimorthrius Pic, 1940 (during this study, the latter two genera were discovered to belong to the subfamily Tillinae).
Historic overview
Gorham (1876) described the genus Orthrius for Orthrius cylindricus and noticed the relationship to Thanasimus, and, on the basis of the tarsal structure, to Clerus. Seven years later, the same author (Gorham 1883) established the genus Gyponyx and mentioned its relationship to Thanasimus and Axina. Chevrolat (1842) described the species “oblongus”, drawing attention to its flightlessness and established the genus Dozocolletus, without a generic diagnosis; a diagnosis was given later by Lacordaire (1857). Quedenfeldt (1885) described the genus Aphelochroa (with Aphelochroa carneipennis as type species) comparing it with Opilo and Natalis. Later, Gorham (1892) established the new genus Xenorthrius for three new species (Xenorthrius balteatus, Xenorthrius mouhoti and Xenorthrius subfasciatus). For another two wingless species Schenkling (1908) erected the genus Caridopus and in the same publication, described the species Apteroclerus brevis from the Kilimanjaro, though with reservations about its generic placement. In two different publications (Pic (1940a, 1940b) respectively described the genera Languropilus and Pieleus, while in an earlier paper (Pic 1933), he had expressed his view that the flightless Astigmus pygidialis differs greatly from all other Astigmus species.
Material and methods
Abbreviations
A
CuA2
MNHN
MRAC
MSNG
MZLU
RGCM
r3, r4
RP2
SDEI
T
Cladistic analysis
23 characters with their respective states (Tab. 1) were analysed. Character polarity was determined by the outgroup method (Nixon and Carpenter 1993); no ancestral states were forced. The genus Clerus Geoffroy, 1762, was considered the outgroup taxon. The data matrix (Tab. 2) was analysed with the Willi Hennig Society edition of TNT 1.1 from September 2009 (Goloboff et al. 2003, 2008). To receive an exact solution, every possible tree was computed by using the “implicit enumeration” routine.
For characters with more than one state per genus, multiple character states were used; they appear enclosed by square brackets in the matrix. Characters that were ambiguous, or missing in the available specimen, appear as a question mark. All characters were chosen to be nonadditive and none were weighted. Implied weighting was also turned off. The species were sorted alphabetically within the input file.
Diagnosis
Species of the Orthrius-group are readily distinguished from other Clerinae by the presence of the following characters (in combination):
– Eyes distinct, more or less protruding laterally, coarsely facetted
– Eyes separated by more than one eye width
– Labrum bilobed to broadly V-shaped (Fig. 1)
Figures 1–10.
1–4 Labrum, labium, maxille and antenna of Orthrius sepulcralis 5 Pro-intercoxal process of Xenorthrius robustus 6 Pro-intercoxal process of Orthrius sepulcralis 7 Anterior mesosternal process of Xenorthrius robustus 8–9 Tarsus and claws of Orthrius sepulcralis 10 Claws of Xenorthrius robustus.
– Terminal segment of labial palpi securiform (Fig. 2)
– Terminal segment of maxillary palpi cylindrical (to digitiform) (Fig. 3)
– Antennal flagellum more or less filiform (Fig. 4)
– Antennomere 2 shorter than antennomere 3 (except Languropilus)
– Procoxal cavities broadly open posteriorly (Figs 5, 6)
– Pro-intercoxal process not (or only slightly) dilated distally (Figs 5, 6)
– Metendosternites without anterior tendons, furcal arms distinct, furcal laminae mostly distinct, furcal stalks mostly of normal length or very short in wingless genera (Figs 11–20)
Figures 11–20.
Metendosternites of 11 Aphelochroa sp. 12 Caridopus sp. 13 Dozocolletus discophorus 14 Gyponyx sp. 15 Languropilus fortipes 16 Neorthrius sp. 17 Nonalatus brevis 18 Orthrius sepulcralis 19 Pseudoastigmus pygidialis 20 Xenorthrius loricus. Scale bars 0.5mm.
– Elytra without sharply-defined basal margin
– Typical wing venation (if winged), with open wedge cell, r3, r4 and CuA2 (except Pieleus) present, RP2 absent (Figs 21–28)
Figures 21–28.
Wings of 21 Aphelochroa sp. 22 Caridopus sp. 23 Gyponyx sp. 24 Languropilus sp. 25 Neorthrius sp. 26 Orthrius sp. 27 Pieleus sp. 28 Xenorthrius sp. Scale bars 5mm.
– Pro- and meso-tarsi each with four pulvilli (number of metatarsal pulvilli variable) (Fig. 8)
– Hind tarsi: T2<T3 + T4 (Tarsomere 2 smaller than tarsomeres 3 and 4 together)
– Spiculae of spicular fork more or less dilated (Figs 29–37)
Figures 29–37.
Spicular forks of 29 Aphelochroa sp. 30 Caridopus monstruosus 31 Dozocolletus discophorus 32 Gyponyx sp. 33 Neorthrius sp. 34 Nonalatus brevis 35 Orthrius sepulcralis 36 Pseudoastigmus pygidialis 37 Xenorthrius simplex. Scale bars 0.5mm.
Key to genera
| 1 | Pronotum with six gibbosities on disc | Caridopus |
| – | Pronotum without six gibbosities on disc | 2 |
| 2 | Wingless species | 3 |
| – | Species with wings | 5 |
| 3 | Antennomeres of flagellum from A4 dilated, antennal club absent | 4 |
| – | Flagellum filiform, antennal club with 3 antennomeres | Dozocolletus |
| 4 | Tarsal pulvillar formula 4-4-2 | Nonalatus gen. n. |
| – | Tarsal pulvillar formula 4-4-3 | Pseudoastigmus gen. n. |
| 5 | Claws dentate (Fig. 10) | 6 |
| – | Claws simple (Fig. 9) | 7 |
| 6 | Tibial spur formula 1-1-2, elytral punctation not arranged into 10 striae | Pieleus |
| – | Tibial spur formula 1-2-2, elytral punctation arranged into 10 striae | Xenorthrius |
| 7 | Anterior mesosternal process present (Fig. 7), tarsal pulvillar formula 4-4-4 | Gyponyx |
| – | Anterior mesosternal process absent, tarsal pulvillar formula 4-4-3 | 8 |
| 8 | Tibial carinae absent | Languropilus |
| – | Tibial carinae present | 9 |
| 9 | Eyes weakly emarginate | 10 |
| – | Eyes conspicuously emarginate | Neorthrius gen. n. |
| 10 | Tibial spur formula 1-2-2 | Aphelochroa |
| – | Tibial spur formula 0-1-1 | Orthrius |
Description of genera
Aphelochroa
Quedenfeldt, 1885
http://species-id.net/wiki/Aphelochroa
Figures 38–46.
Aedeagi of 38 Aphelochroa sp. 39 Caridopus sp. 40 Dozocolletus discophorus 41 Gyponyx sp. 42 Neorthrius sp. 43 Nonalatus brevis 44 Orthrius sepulcralis 45 Pseudoastigmus pygidialis 46 Xenorthrius simplex. Scale bars 1mm.
Figures 47–57.
Terminal abdominal segments of 47 Aphelochroa sp. 48 Caridopus sp. 49 Dozocolletus discophorus 50 Gyponyx sp. 51 Languropilus fortipes 52 Neorthrius sp. 53 Nonalatus brevis 54 Orthrius sepulcralis 55 Pieleus irregularis 56 Pseudoastigmus pygidialis 57 Xenorthrius simplex. Scale bars 1mm.
Figures 58–63.
Habitus of 58 Aphelochroa sp. 59 Caridopus sp. 60 Dozocolletus discophorus 61 Gyponyx sp. 62 Languropilus fortipes 63 Neorthrius sp.
Burgeonus Pic, 1950 syn. n.; Pic 1950: 158.
Type species:
Aphelochroa carneipennis Quedenfeldt, 1885. Quedenfeldt 1885: 267; Kraatz 1899: 86; Schenkling 1902: 326; Schenkling 1903: 29, 57; Mawdsley 1994: 128; Mawdsley and Sithole 2010: 1.
Distribution:
Aethiopian region.
Material examined:
Aphelochroa sanguinea (Thomson, 1857), Kenya, Voi, Sagala Region, 12.1991, leg. K. Werner. Aphelochroa sanguinalis (Westwood, 1852), Congo, VIII.1959, Albertville. Aphelochroa fulva Kraatz, 1899, Kenya, Meru Distr., Materi (Mitunguu), mt. 800, R. Mourglia legit; and several other specimens of this genus (all RGCM). Burgeonus freynei Pic, 1950 (Holotype), Coll. Mus. Congo, Lulua: Luashi, XI-1938, F. Freyne; R. DET., X., 5621; desiré; Burgeonus freynei n sp [handwritten by Pic](MRAC).
Description
Head:
Eyes strongly protruding, only slightly emarginate at antennal insertion; interocular space more than one eye width; gular sutures converging, gular process broad; A1 large, stout, almost twice as long as A2, A2 shorter than A3, A3-A8 filiform, antennomeres becoming shorter, A9 dilated distally, A10 broader than long, A11 sub-ovate, apical third pinched, terminal three antennomeres forming a loose club.
Thorax:
Proepimeron short, not acute; anterior mesosternal process absent; proepimeron short; metendosternite with normal furcal stalk, short, normal furcal arms and very slightly emarginate stalk base (Fig. 11). Elytra long, subparallel, broadest behind middle, apices broadly rounded, elytral punctation not arranged into striae.
Legs:
Of normal size, stout; tarsal pulvillar formula 4-4-3, tibial spur formula 1-2-2; tibiae with longitudinal carinae; claws simple.
Abdomen:
Apical margin of male ventrite 6 distinctly emarginate (Fig. 47); tegmen slender, tapering to a curved acumination distally, phallic struts acuminate, not fused, phallobasic apodeme slightly dilated distally (Fig. 38).
Caridopus
Schenkling, 1908
http://species-id.net/wiki/Caridopus
Type species:
Caridopus monstruosus Schenkling, 1908. Schenkling 1908: 71.
Distribution:
Aethiopian region.
Material examined:
Caridopus monstruosus (Type), Kilimandj., Sjöstedt; Kibonoto, kulturz.; 30. April; Caridopus monstruosus, Typus! (NRM). Caridopus affinis Schenkling, 1908 (Type), Meru, Regenwald; Meru, Sjöstedt; Caridopus affinis Schklg., Typus! (NRM).
Description
Head:
Eyes strongly protruding, only slightly emarginate at antennal insertion; interocular space more than 1.5 eye widths; gular sutures converging, gular process broad; antennae long, A2 shorter than A3, A3-A8 filiform, antennomeres becoming shorter, A9 and especially A10 dilated distally, A10 shorter than A9, A11 sub-ovate, apical third pinched, without club.
Thorax:
Conspicuously longer than broad, with six gibbosities on disc; pro-intercoxal process narrow, linear; proepimeron short, acute to slightly rounded; anterior mesosternal process present; metendosternite with normal furcal stalk length, furcal arms acute distally, stalk base conspicuously emarginate (Fig. 12). Elytra compact (broadest behind middle), conspicuously constricted at base and strongly dilated apically in wingless species, apices broadly rounded, elytral punctation arranged into ten more or less regular striae; wingless or with hindwings.
Legs:
Long, very stout, femora conspicuously thickened; tarsal pulvillar formula 4-4-4, tibial spur formula 1-2-2; tibiae without longitudinal carinae; claws simple.
Abdomen:
Apical margin of male ventrite 6 deeply emarginate (Fig. 48); phallobasic struts fused, phallic struts very broad, phallobasic apodeme strongly dilated distally (Fig. 39).
Dozocolletus
Chevrolat, 1842
http://species-id.net/wiki/Dozocolletus
Brinckodes Winkler, 1960 syn. n.; Winkler 1960: 130.
Quasibrinckodes Winkler, 1960 syn. n.; Winkler 1960: 134.
Type species:
Dozocolletus oblongus Chevrolat, 1842. Chevrolat 1842: 278; Lacordaire 1857: 442; Schenkling 1903: 28, 38.
Distribution:
Aethiopian region (southern Africa).
Material examined:
Dozocolletus discophorus (Boheman, 1851)(Type), Caffraria, J. Wahlb, Type. Dozocolletus puberulus (Boheman, 1851)(Type), Caffraria, J. Wahlb, Type. Dozocolletus oblongus Chevrolat, 1842, Pretoria, 2.XII.1963, leg. A.L. Capener (all NRM). Brinckodes apterus Winkler, 1960 (Holotype and two Paratypes), S. Afr. Transvaal, 16 miles NE of Pretoria, Oct.-Nov. 1954, G. Rudebeck; Brinckodes apterus n.g., n.sp., Det. J.R. Winkler, 1959. Brinckodes apterus ab. ater Winkler, 1960 (Holotype), S. Afr. Transvaal, 16 miles NE of Pretoria, Oct.-Nov. 1954, G. Rudebeck; Brinckodes apterus n.g., n.sp., n.ab., Det. J.R. Winkler 1959. Quasibrinckodes pictus Winkler, 1960 (Holotype), 8200 ft.; S. Afr. Cape Prov., Drakensbergen, 8 miles ENE Rhodes, 10.III.51, No 223; Swedish South Africa Expedition, 1950–1951, Brinck - Rudebeck; Quasibrinckodes pictus n.g., n.sp., Det. J.R. Winkler 1959, Holotypus (all MZLU).
Description
Head:
Eyes protruding, very slightly emarginate at antennal insertion; interocular space two to three eye widths; gular sutures converging, gular process broad; antennae long, A1 large, stout, almost twice as long as A2, A2 shorter than A3, A3-A8 filiform, antennomeres becoming shorter, A9 short, transverse, A10 larger than A9, transverse, A11 approximately equal in length to A9+A10, sub-ovate, apical half pinched, terminal three antennomeres forming a distinct club.
Thorax:
Pronotum conspicuously constricted towards base, without transverse impression, proepimeron short to medium-sized, not acute; anterior mesosternal process present, broadly bent, proepimeron broad, short; metendosternite with very short furcal stalk, stalk base broad, with a deep emargination, furcal arms long, acute distally (Fig. 13). Elytra short, elytral base strongly constricted, broadest behind middle, apices rounded, elytral punctation arranged into ten striae; wingless.
Legs:
Relatively short, stout; femora conspicuously thickened (especially profemora); tarsal pulvillar formula 4-4-4, tibial spur formula 2-2-2; tibiae with longitudinal carinae; claws simple, stout.
Abdomen:
Apical margin of male ventrite 6 not emarginate (Fig. 49); tegmen relatively broad, phallobasic struts fused, phallic struts broad, dilated distally, phallobasic apodeme not dilated distally (Fig. 40).
Gyponyx
Gorham, 1883
http://species-id.net/wiki/Gyponyx
Type species:
Notoxus chinensis Fabricius, 1794. Gorham 1883: 604; Schenkling 1900: 14; Schenkling 1903: 29, 45; Schenkling 1907: 199.
Distribution:
Aethiopian region.
Material examined:
Gyponyx apicalis (Chevrolat, 1842), Südafrika, SE 3130AA, Umtanvuma 3.1.1989, leg. T. Beyers; Gyponyx signifer (Boheman, 1851), Tanzania, Nufindi Dist., Nafinga 1000m, 21.11.-4.12.1989, leg. R. Mourglia; and several further specimens of this genus (all RGCM).
Description
Head:
Eyes strongly protruding, broadly but not deeply emarginate at antennal insertion; interocular space more than 1.5 eye widths; gular sutures converging, gular process broad; A1 large, stout, almost two times longer than A2, A2 shorter than A3, A3-A6 filiform, A7-A10 slightly dilated distally, antennomeres becoming shorter, A11 sub-ovate, apical third pinched, without club.
Thorax:
Proepimeron medium-sized, more rounded than acute; anterior mesosternal process present; metendosternite with normal furcal stalk, short, normal furcal arms and very slightly emarginate stalk base (Fig. 14). Elytra long, subparallel, strongly dilated apically (broadest behind middle), apices broadly rounded, elytral punctation arranged into ten more or less regular striae.
Legs:
Of normal size; tarsal pulvillar formula 4-4-4, tibial spur formula 2-2-2; tibiae with longitudinal carinae; claws simple.
Abdomen:
Apical margin of male ventrite 6 very slightly emarginate (Fig. 50); tegmen broad, phallobasic struts fused, phallic struts and phallobasic apodeme broad, but not conspicuously dilated distally (Fig. 41).
Languropilus
Pic, 1940
http://species-id.net/wiki/Languropilus
Type species:
Languropilus fortipes Pic, 1940. Pic 1940a: 3.
Distribution:
Aethiopian region (East Africa).
Material examined:
Languropilus fortipes (females), Tanzania, Shinyanga Prov., Serengeti Sopa L., 19.XI.93, LF., Heiss (RGCM).
Description
Head:
With weakly protruding eyes, only very slightly emarginate at antennal insertion; interocular space about two times one eye width; gular sutures long, converging, gular process broad; antennae short, A1 large, stout, almost two times longer than A2, A2=A3 or A2>A3, A3-A8 filiform, antennomeres becoming shorter, A8 almost spherical, A9 and A10 transverse, A11 ovate, terminal three antennomeres forming a distinct club.
Thorax:
Proepimeron medium-sized, more rounded than acute; anterior mesosternal process absent; metendosternite with normal furcal stalk length, stalk slender, base almost straight, furcal arms of more or less normal length, acute distally (Fig. 15). Elytra long, broadest behind middle, apices broadly rounded, elytral punctation arranged into ten striae.
Legs:
Of normal size, stout; tarsal pulvillar formula 4-4-3, tibial spur formula 1-2-2; tibiae without longitudinal carinae; claws simple.
Neorthrius
Gerstmeier & Eberle gen. n.
urn:lsid:zoobank.org:act:37980052-1760-48A2-8AA8-F677612AA8AE
http://species-id.net/wiki/Neorthrius
Type species:
Neorthrius monticola Schenkling, 1906 Schenkling 1906: 267.
Distribution:
Indo-Australian region.
Material examined:
Neorthrius monticola (Holotype), Kina-Balu-Geb., 1500m, Coll. Waterstrad; Schenkling det (SDEI); and several unidentified specimens of this genus.
Description
Head:
Eyes strongly protruding, conspicuously emarginate at antennal insertion; interocular space at least more than one eye width; gular sutures converging, gular process broad, compact, only slightly emarginate at middle; antennae long, A1 about two times longer than A2, A2 shorter than A3, A3-A8 filiform, A9 and A10 slightly dilated distally, A3-A5 more or less equal in length, A6-A8 becoming shorter, A11 sub-ovate, apical half pinched, sometimes without club, sometimes terminal three antennomeres forming a loose club.
Thorax:
Proepimeron short to medium-sized, more rounded than acute; anterior mesosternal process absent; metendosternite with normal furcal stalk length, furcal arms normal, stalk base slightly emarginate (Fig. 16). Elytra long, subparallel, sometimes constricted apically, apices rounded separately, elytral punctation arranged into ten striae.
Legs:
Of normal size, sometimes with thickened femora; tarsal pulvillar formula 4-4-3, tibial spur formula 1-2-2; tibiae with longitudinal carinae; claws simple.
Abdomen:
Apical margin of male ventrite 6 sometimes deeply emarginate (Fig. 52); phallobasic struts not fused, phallic struts and phallobasic apodeme dilated distally (Fig. 42).
Nonalatus
Gerstmeier gen. n.
urn:lsid:zoobank.org:act:4331C030-7A93-4F7D-A6F1-D896965CF99D
http://species-id.net/wiki/Neorthrius
Figures 64–68.
Habitus of 64 Nonalatus brevis 65 Orthrius sepulcralis 66 Pieleus irregularis 67 Pseudoastigmus pygidialis 68 Xenorthrius mouhoti.
Type species:
Apteroclerus brevis Schenkling, 1908, comb. n. Schenkling 1908: 71.
Distribution:
Aethiopian region (Kilimanjaro).
Material examined:
Apteroclerus brevis (Type), Kilimandj., Sjöstedt; Kiboscho, 3’-4000m; 15. febr.; Bärgs-ängarne; Typus; Bergwiesen, Ericinella-Region, In den trockenen Blumenständen von Lobelia deekeni (NRM).
Description
Head:
Eyes protruding, emarginate at antennal insertion; interocular space two to three eye widths; gular sutures strongly diverging, gular process broad; antennae long, A2 shorter than A3, from A4 slightly dilated apically, A3-A7 becoming shorter, A9 and A10 more or less equal in length, A11 longer than A10, A11 sub-ovate, apical third pinched, without club.
Thorax:
Proepimeron short, not acute; anterior mesosternal process present, broadly bent; metendosternite with very short furcal stalk, stalk base deeply emarginate, furcal arms acute distally (Fig. 17). Elytra ovate, short, compact, strongly constricted at base and towards apex, broadest behind middle, apices broadly rounded, elytral punctation arranged into ten irregular striae; wingless.
Legs:
Relatively long, stout; tarsal pulvillar formula 4-4-2, tibial spur formula 1-2-2; tibiae without longitudinal carinae; claws simple, with a very small, acute basal denticle.
Abdomen:
Apical margin of male ventrite 6 distinctly emarginate (Fig. 53); tegmen very broad, parameres expanded laterally, tapering to an acumination distally, phallobasic struts not fused, phallic struts and phallobasic apodeme not dilated distally (Fig. 43).
Orthrius
Gorham, 1876
http://species-id.net/wiki/Orthrius
Figs 1–4, 6, 8–9 18 26 35 44 54 65
Dedana Fairmaire, 1888, syn. n.; Fairmaire 1888: 26; Schenkling 1903: 4, 23.
Type species:
Orthrius cylindricus Gorham, 1876. Gorham 1876: 74.
Distribution:
Indo-Australian region.
Material examined:
Orthrius cylindricus (Type), NSW; Orthrius Gorh., cylindricus G., Type; Museum Paris, Coll. Gorham, 1914 (MNHN); and several other specimens of this genus. Dedana rufodorsata Fairmaire, 1888 (Type), Fokien; Dedana rufodorsata Fairm.; ExMusaeo Arm. David, 1900 (MNHN).
Description
Head: Eyes strongly protruding, only slightly emarginate at antennal insertion; interocular space more than one eye width; gular sutures converging, gular process broad; antennae long, A2 shorter than A3, A2-A8 filiform, A10 broadest, A11 sub-ovate, apical half pinched, terminal three antennomeres forming a more or less conspicuous club.
Thorax: Proepimeron short to medium-sized, not acute; anterior mesosternal process absent; metendosternite with normal furcal stalk length, furcal arms normal, stalk base very slightly emarginate (Fig. 18). Elytra long, subparallel, sometimes dilated apically (broadest behind middle), apices rounded, elytral punctation not arranged into striae.
Legs: Long, especially profemora intermediately to strongly thickened; tarsal pulvillar formula 4-4-3, tibial spur formula 0-1-1; tibiae with longitudinal carinae; claws simple.
Abdomen: Apical margin of male ventrite 6 straight or slightly emarginate (Fig. 54); tegmen relatively broad, parameres expanded laterally, tapering to a curved acumination distally, phallobasic struts not fused, phallobasic apodeme dilated distally (Fig. 44).
Pieleus
Pic, 1940
http://species-id.net/wiki/Pieleus
Type species:
Pieleus irregularis Pic, 1940. Pic 1940b: 4.
Distribution:
China.
Material examined:
Pieleus irregularis (Type female), T’ienmu Shan, Musée Heude; 20.VII.36, O. Piel, coll.; Orthrius irregularis mihi [handwritten by Pic](MNHN).
Description
Head:
Eyes strongly protruding, conspicuously emarginate at antennal insertion; interocular space about 1.5 eye widths; gular sutures diverging, gular process broad; antennae short, A1 more than two times longer than A2, A2 shorter than A3, A2-A6 filiform, antennomeres becoming shorter, A7 shorter than A6, slightly dilated distally, A11 sub-ovate, apical third pinched, terminal three antennomeres forming a loose club.
Thorax:
Proepimeron medium-sized, slightly rounded; anterior mesosternal process present; metendosternite missing. Elytra compact, strongly dilated apically (broadest behind middle), apices broadly rounded, elytral punctation not arranged into striae; without CuA2 in hindwings (Fig. 27).
Legs:
Of normal size; tarsal pulvillar formula 4-4-4, tibial spur formula 1-1-2; tibiae without longitudinal carinae; claws with basal denticle.
Pseudoastigmus
Eberle gen. n.
urn:lsid:zoobank.org:act:56520F10-5440-4188-ACCE-CBA847869133
http://species-id.net/wiki/Pseudoastigmus
Type species:
Astigmus pygidialis Pic, 1933, comb. n. Pic 1933: 257.
Distribution:
Aethiopian region (Ruwenzori).
Material examined:
Astigmus pygidialis (Syntype), Musée du Congo, Ruwenzori (4200m), VII-1932, L. Burgeon; type; Stigmatium (Astigmus) pygidiale n sp [handwritten by Pic], and four additional syntypes (MRAC).
Description
Head:
Eyes strongly protruding, conspicuously emarginate at antennal insertion; interocular space about two eye widths; gular sutures subparallel to slightly diverging, gular process of medium width; antennae long, A2 shorter than A3, from A4 onwards slightly dilated distally, A11 sub-ovate, apical half pinched, without club.
Thorax:
Proepimeron very short, not acute; anterior mesosternal process present; metendosternite with very short furcal stalk length, furcal arms acute distally, stalk base conspicuously emarginate (Fig. 19). Elytra short, compact, dilated apically (broadest behind middle), apices broadly rounded, elytral punctation arranged into more or less regular ten striae; wingless.
Legs:
Long, stout; tarsal pulvillar formula 4-4-3, tibial spur formula 1-2-2; tibiae without longitudinal carinae; claws with basal denticle.
Abdomen:
Apical margin of male ventrite 6 deeply emarginate (Fig. 56); tegmen relatively broad, tapering to a curved acumination distally, phallobasic struts not fused, phallic struts and phallobasic apodeme not dilated distally (Fig. 45).
Xenorthrius
Gorham, 1892
http://species-id.net/wiki/Xenorthrius
Figs 5, 7, 10 20 28 37 46 57 68
Type species:
Xenorthrius mouhoti Gorham, 1892. Gorham 1892: 733, 1893: 575; Schenkling 1903: 46–47.
Distribution:
Indo-Australian and Palaearctic region.
Material examined:
Xenorthrius mouhoti, Lectotype (MSNG), Paralectotypes, and additional species (see Gerstmeier and Eberle 2010).
Description
Head:
Eyes strongly protruding, conspicuously emarginate at antennal insertion; interocular space larger than one eye width; gular sutures subparallel to divergent, gular process varying in width, from narrow to broad; antennal length interspecifically variable and sometimes sexually dimorphic (longer in males), A2 shorter than A3, A3-A8 more or less filiform, A10 broader than long, A11 sub-ovate, apical half pinched, mostly without club, sometimes terminal three antennomeres forming a loose club.
Thorax:
Proepimeron medium-sized, more or less acute; anterior mesosternal process present, with a subtriangular sulcus in the middle (Fig. 7); metendosternite with normal furcal stalk length, furcal arms broad, apically dilated, stalk base very slightly to deeply emarginate (Fig. 20). Elytra subparallel, sometimes broadest behind middle, apices rounded (most species), strongly dehiscent (Xenorthrius prolongatus and Xenorthrius furcalis), or dentate (Xenorthrius truncatus and Xenorthrius scordalus); elytral punctation arranged into ten striae.
Legs:
Mostly relatively short; tarsal pulvillar formula 4-4-4, tibial spur formula 1-2-2; tibiae with or without longitudinal carinae; claws with pronounced basal denticle (Fig. 10).
Abdomen:
Apical margin of male ventrite 6 more or less distinctly emarginate (Fig. 57); tegmen mostly elongate, cross-section subrectangular; phallobasic struts not fused, phallic struts acute, phallobasic apodeme not dilated distally (Fig. 46).
Discussion of cladistic results
The cladistic analysis resulted in a single most parsimonious tree with a length of 37 steps (Fig. 69). Common to all taxa of the Orthrius-group are four mesotarsal pulvilli (char. 0-0) and coarse ommatidial facets (char. 5-0) which distinguishes them from the Clerus-series.
Figure 69.
Cladistic tree of the genera of the Orthrius-group.
Table 1.
Characters and character states used in the cladistic analysis of the genera.
| Character 0 | Mesotarsal pulvilli: (0) 4; (1) 3 |
| Character 1 | Metatarsal pulvilli: (0) 4; (1) 3; (2) 2 |
| Character 2 | Protibial spurs: (0) 2; (1) 1; (2) 0 |
| Character 3 | Mesotibial spurs: (0) 2; (1) 1 |
| Character 4 | Metatibial spurs: (0) 2; (1) 1 |
| Character 5 | Ommatidial facets: (0) coarse; (1) fine |
| Character 6 | Flagellomeres: (0) filiform; (1) dilated |
| Character 7 | Eye’s emargination: (0) absent or weak; (1) conspicuous |
| Character 8 | Eye’s separation: (0) more than two eyes width; (1) between one and two eyes width |
| Character 9 | Gular sutures: (0) convergent to subparallel; (1) subparallel to divergent |
| Character 10 | Gular process: (0) broad; (1) narrow |
| Character 11 | Relation between A2 and A3: (0) A2 < A3; (1) A2 = A3 or A2 > A3 |
| Character 12 | Anterior mesosternal process: (0) present; (1) absent |
| Character 13 | Metendosternite, furcal stalk length: (0) normal; (1) very short |
| Character 14 | Metendosternite, furcal arms: (0) normal; (1) acute |
| Character 15 | Metendosternite, furcal stalk base: (0) normal; (1) deeply emarginate |
| Character 16 | Wings: (0) present; (1) absent |
| Character 17 | CuA2: (0) present; (1) absent |
| Character 18 | RP2: (0) present; (1) absent |
| Character 19 | Elytral punctation: (0) with 10 regular striae; (1) with 10 irregular striae; (2) with more than 10 irregular striae |
| Character 20 | Tibial carinae: (0) present; (1) absent |
| Character 21 | Claws: (0) simple; (1) with basal denticle |
| Character 22 | Phallobasic struts: (0) not fused; (1) fused |
Pseudoastigmus gen. n.and Nonalatusgen. n. appear together at the base of the tree. This pair is supported by the acute form of the furcal arms of the metendosternite (char. 14-1) as well as the complete reduction of the hind wings (char. 16-1).
The remaining taxa share the filiform flagellum (char. 6-0). The development of four pulvilli at the metatarsus (char. 1-0) is also synapomorphic at this point, but is reduced to three pulvilli for the cluster of Neorthrius gen. n., Languropilus, Orthrius and Aphelochroa (char. 1-1).
These four genera also share the loss of the anterior mesosternal process (char. 12-1). Like in Dozocolletus and Caridopus the emargination of the eyes is weak or absent (char. 7-0) in Languropilus, Orthrius and Aphelochroa. For this reason, Neorthrius adopts a basal position in this group. The monophyly of Orthrius and Aphelochroa is supported by their elytral punctation (char. 19-2). Orthrius differs from all other taxa in this revision in its tibial spur formula which is 0-1-1 (chars. 2-2, 3-1 and 4-1).
The aethiopian genera Gyponyx, Dozocolletus and Caridopus have in common, that the phallobasic struts are fused with the phallobasic apodeme (char. 22-1). The monophyly of Dozocolletus and Caridopus is well supported by the weak or absent emargination of the eyes (char. 7-0) and similarities of their metendosternites: the furcal arms are acute (char. 14-1) and the furcal stalk base (char. 15-1) is deeply emarginate.
A common ancestor can be assumed for the latter two clusters of genera.This is supported by two synapomorphies: the gular sutures are convergent to parallel (char. 9-0) and the claws are simple (char. 21-0). The presence of the tibial carinae (char. 20-0) also is apomorphic at this node but reduced in Caridopus and Languropilus. As Solervicens (2007) mentioned, it also may be considered a symplesiomorphy, because it is a common character of the Clerinae.
Table 2.
Character matrix of 23 adult morphological characters of Clerus (outgroup) and genera of the Orthrius
| Taxa | Characters | ||||||||||||||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 0 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | |
| Clerus | 1 | 2 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 |
| Aphelochroa | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 0 | 0 |
| Caridopus | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | [01] | ? | ? | 0 | 1 | 0 | 1 |
| Dozocolletus | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | ? | ? | 0 | 0 | 0 | 1 |
| Gyponyx | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 |
| Languropilus | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | ? |
| Neorthrius | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
| Nonalatus | 0 | 2 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | ? | ? | 1 | 1 | 1 | 0 |
| Orthrius | 0 | 1 | 2 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 0 | 0 |
| Pieleus | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | ? | ? | ? | 0 | 1 | 1 | 2 | 1 | 1 | ? |
| Pseudoastigmus | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | ? | 1 | ? | 1 | ? | ? | 0 | 1 | 1 | 0 |
| Xenorthrius | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | [01] | 0 | 0 | 0 | 0 | [01] | 0 | 0 | 1 | 0 | [01] | 1 | 0 |
Supplementary Material
Acknowledgements
Our sincere thanks go to Marianne Müller (the senior author's wife) who helped with the photos of specimens. For the kind loan of specimens we thank Thierry Deuve, Antoine Mantilleri and Azadeh Taghavian (MNHN), Marc de Meyer, MRAC, Roberto Poggi (MSNG), Roy Danielsson, MZLU and Lothar Zerche, SDEI. We would like to express our sincere thanks to Justin S. Bartlett (Brisbane, Australia) and Weston Opitz (Salina, USA) for important comments on earlier drafts of this manuscript. This research received support from the SYNTHESYS Project http://www.synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP6 “Structuring the European Research Area Programme”.
References
- Chevrolat A. (1842) Descriptions de quelques Térédiles de l’Afrique australe, du voyage de M. Drege. Revue Zoologique 5:276-278. [Google Scholar]
- Chapin EA. (1924) Classification of the Philippine components of the coleopterous family Cleridae. The Philippine Journal of Science 25 (2):159-286. [Google Scholar]
- Corporaal JB. (1950) Coleopterorum Catalogus, Supplementa pars 23, Cleridae. Uitgeverij Dr. W. Junk, ‘s-Gravenhage. [Google Scholar]
- Crowson RA. (1955) The natural classification of the families of Coleoptera. Nathaniel Lloyd, London, 187 pp. [Google Scholar]
- Crowson RA. (1964) A review of the classification of Cleroidea (Coleoptera) with descriptions of two new genera of Peltidae and of several new larval types. Transactions of the Royal Entomological Society of London 116 (12):275-327. [Google Scholar]
- Crowson RA. (1966) Further observations on Peltidae (Coleoptera: Cleroidea), with definitions of a new subfamily and of four new genera. Proceedings of the Royal Entomological Society of London (B) 35:119-127. 10.1111/j.1365-3113.1966.tb00523.x [DOI] [Google Scholar]
- Crowson RA. (1970) Further observations on Cleroidea (Coleoptera). Proceedings of the Royal Entomological Society of London (B) 39 (1–2):1-20. [Google Scholar]
- Fairmaire L. (1888) Coléoptères de l’intérieur de la Chine. Annales de la Société entomologiques Belgiques 32:7-76. [Google Scholar]
- Gerstmeier R. (2000) Aktueller Stand der Buntkäfer-Forschung (Coleoptera, Cleridae, Thanerocleridae). Entomologica Basiliensia 22:169-178. [Google Scholar]
- Gerstmeier R. (2002) Generic concept of clerid taxa related to Clerus Geoffroy, 1762 (Coleoptera: Cleridae: Clerinae). Entomological Problems 33 (2):99-111. [Google Scholar]
- Gerstmeier R. (2006) Key to Genera of Checkered Beetles of the Himalayas (Fam. Cleridae and Thanerocleridae). In: Hartmann M, Weipert J. (Eds) Biodiversität und Naturausstattung im Himalaya II. Verein der Freunde und Förderer des Naturkundemuseums Erfurt e.V., Erfurt, 467–470. [Google Scholar]
- Gerstmeier R, Eberle J. (2010) Revision of the Indo-Australian checkered beetle genus Xenorthrius Gorham, 1892 (Coleoptera: Cleridae, Clerinae). Zootaxa 2584:1-121. [Google Scholar]
- Goloboff P, Farris J, Nixon K. (2003) T.N.T.: Tree Analysis Using New Technology. Program and documentation, available from the authors and at www.zmuc.dk/public/phylogeny.
- Goloboff P, Farris J, Nixon K. (2008) TNT, a free program for phylogenetic analysis. Cladistics 24 (5), 774–786. 10.1111/j.1096-0031.2008.00217.x [DOI] [Google Scholar]
- Gorham HS. (1876) Notes on the coleopterous family Cleridae, with descriptions of new genera and species. Cistula Entomologica 2:57-106. [Google Scholar]
- Gorham HS. (1883) Descriptions of Malacodermata in the Civic Museum of Natural History at Genua. Annali del Museo Civico di Storia Naturale di Genova 18:591-606. [Google Scholar]
- Gorham HS. (1892) Viaggio di Leonardo Fea in Birmania e Vicine. Annali del Museo Civico di Storia Naturale di Genova, Ser. 2a 12 (32):718-746. [Google Scholar]
- Gorham HS. (1893) A list of the Coleoptera, of the family Cleridae, collected by Mr. Doherty in Burmah and Northern India, with descriptions of new species; and of some species from Borneo, Perak, &c., from the collection of Alexander Fry, Esq. Proceedings of the Zoological Society of London 39:566-581. [Google Scholar]
- Kolibáč J. (1992) Revision of Thanerocleridae n.stat. (Coleoptera, Cleroidea). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 65:303-340. [Google Scholar]
- Kolibáč J. (1997) Classification of the subfamilies of Cleridae (Coleoptera: Cleroidea). Acta Musei Moraviae, Scientiae naturale 81(1/2): 307–361. [Google Scholar]
- Kolibáč J. (1998) Classification of the subfamily Hydnocerinae Spinola, 1844 (Coleoptera: Cleridae). Acta Musei Moraviae, Scientiae biologicae 83:127-210. [Google Scholar]
- Kolibáč J. (2003) A review of Australian genera of Korynetinae (Coleoptera, Cleridae). Entomologica Basiliensia 25:41-97. [Google Scholar]
- Kolibáč J. (2004) Metaxinidae fam.nov., a new family of Cleroidea (Coleoptera). - Entomologica Basiliensia 26:239-268. [Google Scholar]
- Kraatz G. (1899) Verzeichnis der von Hrn. Conradt in Westafrica (Kamerun, Togo) gesammelten Cleriden. Deutsche entomologische Zeitschrift 1899 (1):81-107. [Google Scholar]
- Lacordaire JT. (1857) Histoire Naturelle des Insectes. Genera des Coleopteres IV. Libraire encyclopédique de Roret, Paris, 579 pp. [Google Scholar]
- Lawrence JF, Newton Jr. AF. (1995) Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). In: Pakulak J, Slipinski SA. (Eds) Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson, Muzeum i Instytut Zoologii PAN, Warszawa, Volume two, 779–1006. [Google Scholar]
- Mawdsley JR. (1994) A revision of the genus Aphelochroa Quedenfeldt (Coleoptera: Cleridae: Clerinae). Journal of African Zoology 108:121-132. [Google Scholar]
- Mawdsley JR, Sithole H. (2010) Observation on the biology of the South African checkered beetle Aphelochroa sanguinalis (Westwood) (Coleoptera: Cleridae). Insecta Mundi 0111:1–6. http://centerforsystematicentomology.org/insectamundi/0111MawdsleyandSithole.pdf [Google Scholar]
- Nixon KC, Carpenter JM. (1993) On outgroups. Cladistics 9:413-426. 10.1111/j.1096-0031.1993.tb00234.x [DOI] [PubMed] [Google Scholar]
- Opitz W. (1997) Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae). Part I. The genera of Epiphloeinae. Insecta Mundi 11 (1):51-96. [Google Scholar]
- Opitz W. (2002) 73. Cleridae Latreille 1804. In: Arnett Jr. RH. et al. (Eds) American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, 267–280. [Google Scholar]
- Opitz W. (2004) Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae). Part II. The genera Chaetophloeus Opitz and Plocamocera Spinola. Bulletin of the American Museum of Natural History 280:1-82. [DOI] [Google Scholar]
- Opitz W. (2005) Classification, natural history, and evolution of the genus Aphelocerus Kirsch (Coleoptera: Cleridae: Clerinae). Bulletin of the American Museum of Natural History 293:1-128. [DOI] [Google Scholar]
- Opitz W. (2006) Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae). Part III. The genera Parvochaetus, n.gen., Amboakis, n.gen., and Ellipotoma Spinola. Insecta Mundi 20 (3–4):97-164. [Google Scholar]
- Opitz W. (2007) Classification, natural history, and evolution of Epiphloeinae (Coleoptera, Cleridae). Part IV. The genera Pyticeroides Kuwert, 1894 and Diapromeces, Opitz, 1997. Entomologica Basiliensia et Collectionis Frey 29:77-166. [Google Scholar]
- Opitz W. (2008a) Classification, natural history, and evolution of Epiphloeinae (Coleoptera, Cleridae). Part VI. The genera Epiphlaeus Spinola and Opitzius Barr. Annales Zoologici (Warszawa) 58 (1):1-34. [Google Scholar]
- Opitz W. (2008b) Classification, natural history, and evolution of Epiphloeinae (Coleoptera: Cleridae). Part VII. The genera Hapsidopteris Opitz, Iontoclerus Opitz, Katamyurus Opitz, Megatrachys Opitz, Opitzia Nemésio, Pennasolis Opitz, new genus, Pericales Opitz, new genus, Pteroferus Opitz, new genus, and Turbophloeus Opitz, new genus. Zootaxa 1754:1-40. [Google Scholar]
- Opitz W. (2008c) Classification, natural history, and evolution of Epiphloeinae (Coleoptera: Cleridae). Part V. Decorosa Opitz, a new genus of checkered beetles from Hispaniola with description of its four new species. American Museum Novitates 3628:1-19. 10.1206/599.1 [DOI] [Google Scholar]
- Opitz W. (2010) Classification, natural history, phylogeny, and subfamily composition of the Cleridae and generic content of the subfamilies. Entomologica Basiliensia et Collections Frey 32: 31–128. [Google Scholar]
- Pic M. (1933) Clérides du Congo Belge et du Mozambique. Revue de Zoologie et de Botanique Africaines 23 (3–4):255-258. [Google Scholar]
- Pic M. (1940a) Diagnoses de Coléoptères exotiques. L’Echange, Rev.Linn. 56 (479): 2–4. [Google Scholar]
- Pic M. (1940b) [......]. Opuscula martialis 1:1-16. [Google Scholar]
- Pic M. (1950) Nouveaux Clérides, originaires surtout du Congo belge (Col.). Revue de Zoologie et de Botanique Africaines 43 (1–2):152-163. [Google Scholar]
- Quedenfeldt G. (1885) Vier neue Cleriden aus dem tropischen Westafrika. Berliner Entomologische Zeitschrift 29 (2):267-271. 10.1002/mmnd.18850290211 [DOI] [Google Scholar]
- Schenkling S. (1900) Neue Cleriden des Hamburger Museums. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 17:13-20. [Google Scholar]
- Schenkling S. (1902) Clérides nouveaux du Muséum d’Histoire naturelle de Paris. Bulletin du Muséum d’Histoire Naturelle 8:317-333. [Google Scholar]
- Schenkling S. (1903) Coleoptera. Malacodermata. Fam. Cleridae. In: Wytsman P. (Ed) Genera Insectorum, Fasc. 13, P. Wytsman, Bruxelles, 124pp. [Google Scholar]
- Schenkling S. (1906) Die Cleriden des Deutschen Entomologischen National-Museums, nebst Beschreibungen neuer Arten. Deutsche entomologische Zeitschrift 1906:241-320. [Google Scholar]
- Schenkling S. (1907) New species of Cleridae (Coleoptera) in the Entomological Collection of the South African Museum. Annals of the South African Museum 5 (4):197-202. [Google Scholar]
- Schenkling S. (1908) 7. Coleoptera. 7. Cleridae, Erotylidae und Endomychidae. In: Sjöstedt Y Wissenschaftliche Ergebnisse der Schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906, 7 (7):69-78. [Google Scholar]
- Solervicens JA. (2007) Cladistic analysis of the species of Calendyma Lacordaire (1857) and Epiclines Chevrolat (1838) (Coleoptera: Cleridae: Clerinae). Zootaxa 1497:57-68. [Google Scholar]
- Winkler JR. (1960) Coleoptera: Cleridae. In: Hanström B, Brinck P, Rudebeck G (Eds) South African Animal Life 7:128-162. [Google Scholar]
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