Oedichirus Erichson

Herman, Lee, 2010, Generic Revision Of The Procirrina (Coleoptera: Staphylinidae: Paederinae: Pinophilini), Bulletin of the American Museum of Natural History 2010 (347), pp. 1-78 : 36-46

Oedichirus Erichson Figures 3, 14–18, 22–39, 44–51, 66–76

Oedichirus Erichson, 1839: 29 . Type species: Oedichirus paederinus Erichson, 1840: 685 , fixed by Erichson, 1840: 685, by subsequent monotypy.

— Erichson, 1840: 684 (characters; first includ- ed species: paederinus ). — Schaum, 1852: 28 (list; Europe). — Lacordaire, 1854: 104 (characters; notes; list of species). — Kraatz, 1857: 666, 668 (notes; key). — Jacquelin du Val, 1857: 49 (characters). — Redtenbacher, 1857: 217 (characters). — Jacquelin du Val, 1859: 73 (catalog; Europe). — Schaum, 1859: 29 (catalog; Europe). — Gemminger and Harold, 1868: 629 (catalog; world). — Redtenbacher, 1874: 235 (characters). — Fauvel, 1875a: 219 (catalog; France). — Fauvel, 1875b: xvii (catalog; France). — Sharp, 1876: 338 (notes; synonyms; Brazil). — Duvivier, 1883: 176 (catalog). — Fauvel, 1889: 253 (notes). — Heyden et al., 1891: 110 (list; Europe and Caucasus Region). — Heyden et al., 1906: 152 (list; Europe and Caucasus Region). — Bernhauer and Schubert, 1912: 201 (catalog; world). — Cameron, 1925: 34, 106 (catalog; British India). — Winkler, 1925: 358 (catalog; Palaearctic Region). — Porta, 1926: 67 (characters). — Cameron,

69. Labium. 70. Prothorax, ventral. 71. Maxillary palpus. 72. Labrum.

1931: 1, 25 (characters; key to Indian species). — Scheerpeltz, 1933: 1217 (catalog; world). — Blackwelder, 1944: 131 (checklist; Latin America). — Blackwelder, 1952: 269 (type species). — Fagel, 1955: 194 (characters). — Adachi, 1955: 14 (characters; key). — Kocher, 1958: 117 (checklist; Morocco). — Fagel, 1963: 342 (notes). — Fagel, 1971: 11, 126, 196, 369 (characters; notes; key to species; type species). — Tikhomirova, 1973: 175 (checklist; USSR). — Bordoni, 1975: 419 (characters). — Shibata, 1977: 21 (catalog; Japan). — Coiffait, 1978b: 327 (characters; key to western Palaearctic species; distribution). — Hammond, 1984: 203 (checklist; Borneo). — Outerelo and Gamarra, 1985: 22 (characters). — Lecoq, 1986: 7

(characters; key to Madagascar species; discussion). — Ciceroni and Zanetti, 1995: 20 (checklist; Italy). — Navarrete-Heredia et al., 2002: 293 (characters in key to genera for Mexico; general notes; unnamed species in Mexico). — Janák, 2003: 253 (list; Madagascar). — Smetana, 2004: 623 (Palaearctic catalog).

Elytrobaeus Sahlberg, 1847: 801 . Type species: Elytrobaeus geniculatus Sahlberg, 1847: 802 , fixed by monotypy.

— Gemminger and Harold, 1868: 629 (catalog). — Fauvel, 1875a: 219 (synonym of Oedichirus ). — Fauvel, 1875b: xvii (synonym of Oedichirus ). — Sharp, 1876: 338 (synonym of Oedichirus ). — Bernhauer and Schubert, 1912: 201 (synonym of Oedichirus ). — Cameron, 1931: 25 (synonym of Oedichirus ). — Blackwelder, 1952: 147 (type species). — Smetana, 2004: 623 (synonym of Oedichirus ).

Oedichiranus Reitter, 1906: 263 . Type species: Oedichirus dimidiatus Reitter, 1906: 263 , fixed by monotypy. New Synonym.

— Bernhauer and Schubert, 1912: 201 (subgenus of Oedichirus ). — Blackwelder, 1952: 269 (subgenus of Oedichirus ; type species). — Smetana, 2004: 623 (subgenus of Oedichirus ).

DIAGNOSIS: Oedichirus can be separated from all other Procirrina by the spiniform pencil of setae on antennomere 11 (figs. 31, 32) and from all other genera except Palaminus by the ‘‘windows’’ (fig. 24) at the edge of the anterior margin of terga and sterna III to VII. The abdomen of Oedichirus is punctate (fig. 24) and lacks imbricate macrosculpturing and the lateroapical edge of the elytra lacks a long, stout seta (fig. 14). The abdomen of Palaminus lacks punctation and has imbricate sculpturing (fig. 28) and the elytra have a long, stout seta on the lateroapical corner (fig. 11, 96). Unlike other genera of the Procirrina , both Oedichirus and Palaminus have a dense cluster of setae on the ventral surface of protarsomere 5 (figs. 45, 47) and tergum and sternum VII are fused, basally in Oedichirus (figs. 33, 34) and completely in Palaminus . The absence of setae on the edge of the posterior margin of the elytra will separate Oedichirus from Neoprocirrus , Oedodactylus , Pseudoprocirrus , and Stylokyrtus .

DESCRIPTION: Head (fig. 66) not pedunculate, wider than long; frontoclypeus with subapical, interantennal, transverse ridge (fig. 15); subapical ridge curved or sinuate and complete, extending from one antennal fossa to other, or incomplete, extending from each antenna fossa and absent medially; postocular lateral margin rounded to neck or to poorly to well-developed basal angle of head (figs. 15, 66); postocular lateral margin (fig. 15) short to moderately long; basal margin of head distinct to indistinguishable from postocular lateral margin; base of head with (fig. 15) or without ridge extending laterally from neck to near eye. Neck across nuchal constriction half to two thirds as wide as greatest postocular width of head; nuchal groove moderately well developed (fig. 15); nuchal ridge present and well developed (figs. 15, 66). Eye length greater than postocular length of head. Dorsal surface with umbilicate punctation (fig. 15). Ventral surface (fig. 16) without postocular groove. Gular sutures absent or present, moderately to feebly developed, and separated (fig. 68); sutures most approximate near middle. Gular pubescence absent. Antennomere 11 subequal to 10; apex with spiniform pencil of setae (figs. 31, 32). Mandibles (fig. 67) with apically acute denticle; prostheca (fig. 67) evident as cluster of cuticular processes at base. Maxillary palpus (fig. 71) with palpomere 4 longer than second or third, securiform, and moderately compressed. Labium (fig. 69) with glossae broad, widely separated, and apically truncate. Hypopharynx with long submedial seta on anterior margin, but without lobes or cluster of spinelike setae; lateromedial surface covered densely with cuticular processes; sublateral surface with row of coarse setae. Labrum (fig. 72) with one to three pairs of denticles on anterior margin. Epipharynx without setae; median region with transverse, dense cluster of cuticular processes; median groove present; epipharynx not visible along anterior margin in dorsal view.

Prothorax (fig. 66) trapezoidiform, longer than wide in most species, rarely wider than long; widest near anterior third and with lateral margins strongly convergent anteriorly and gradually convergent posteriorly. Pronotum (fig. 18) with umbilicate punctation; punctation sparse to moderately dense, and absent from midline; punctation confused, arranged in straight or curved rows, or more or less evenly distributed. Pronotal marginal ridge present (fig. 18) and well developed, poorly developed, incomplete, or absent. Notosternal suture present (figs. 17, 70); marginal ridge, when present, and suture separated. Hypomeron polished, with or without punctation, and without microsculpturing. Postprocoxal lobe moderately long; punctation present and asetate (fig. 17); transverse ridge absent; apices (fig. 70) moderately widely separated from each other. Probasisternum (fig. 17) with median carina absent in most species, some with short carina; surface with punctures. Mesospiracular peritreme (fig. 17) with anteromedial margin fused to furcasternum. Elytra shorter to longer than pronotum; humeral angles absent (fig. 14) or present; posterior edge (fig. 14) without row of setae; subapical region without long, thicker, medioposteriorly directed seta near lateral corner (fig. 14). Scutellum without pubescence. Mesosternum without median carina. Mesocoxal acetabulum with marginal carina laterally; carina poorly developed to absent posteriorly and position marked by row of punctures. Intersternal suture present, incomplete, and weakly developed.

Profemur with ridge and slit near middle of ventroanterior surface (fig. 36). Protibia with multiple (14 in an Australian species) combs extending from proximad of middle to near apex; tibia without or with feeble depression on ctenidial surface; apical portion slender and slightly tapered to nearly parallel. Protarsomeres (fig. 76) 1–4 inflated and ventral surface without setose pad (figs. 45, 46); base of tarsomere 1 not surrounded by cupulate protibial apex; tarsomere 4 not expanded beneath tarsomere 5 and apical margin entire or weakly emargin- ate (figs. 47, 76), not bilobed; tarsomere 5 unmodified and inserted laterad of middle of asymmetrical tarsomere 4; tarsomere 5 with dense pubescence ventrally (figs. 45, 47). Mesotarsomere and metatarsomere 1 (fig. 50) longer than combination of tarsomeres 2 and 3, 2–4, or 2–5.

Abdominal segments without imbricate macrosculpturing (fig. 24). Segment III with or without one paratergite, if paratergite absent then paratergal carina present basally (figs. 22, 23); tergum and sternum III fused or separated by paratergite. Segment VII with tergum and sternum fused basally and with incision leading to small notch at midlateral edge of posterior margin (figs. 33, 34). Segments III to VII with membranous ‘‘windows’’ or ‘‘cells’’ covered with white, opaque membrane in intersegmental membrane at anterior edge of sterna and terga (fig. 24). Sternum I absent. Sterna IV to VII without glandular lobe on anterior margin. Tergum IX (fig. 73) of male and female fused medially; emargination moderately deep, slightly shorter than length of tergum to more than twice length of tergum; lateroapical process (fig. 25) long, slender, moderately strongly curved ventrally, and extending beyond apex of tergum X; lateroapical process fused basally to remainder of segment, but with possible remnant of separation in some species (fig. 25). Tergum X (fig. 73) of male and female with apical margin broadly rounded; base separated from tergum IX.

Gonocoxites fused into large, broad medi- an gonocoxal plate (fig. 74); vulva proximad of (fig. 74) or embedded in gonocoxal plate.

Aedeagus (fig. 75) asymmetrical; parameres present and appressed to median lobe for entire length of paramere, or appressed basally and free distally; basal piece absent.

DISTRIBUTION AND HABITAT: Most species of Oedichirus are found in tropical and subtropical regions, a few are known from more temperate regions. In the Old World, Oedichirus occurs from southern Europe to South Africa, east to Madagascar and across southern Asia through India, southeastern Asia, to Indonesia, the Philippines, Papua New Guinea, and Australia, China, and Japan. One central Asian species is known ( Tajikistan), one is reported from Tasmania, and two from New Caledonia. New World species are known from Mexico and the Caribbean Islands south to Brazil, but have not been found in all the intervening countries.

Collections of Oedichirus have been made from leaf litter on the forest floor and moist litter near streams, swamps, or springs. Some species have been collected by sweeping lowland forest vegetation at night. Species are found in lowland forests, many live in montane forests, and occur to as high as 3500 m. Of the 175 species for which elevational data are available, 150 are found at elevations between 1000 and 2999 meters. At least 10 species have been collected at or above 3000 meters elevation; they include: O. celisianus , O. ericeticola , O. flavifrons , O. luberensis kenyacus , O. meruensis , O. microphthalmus , O. omoanus , O. orophilus , O. ruwenzoricus , and O. uviraensis (see Fagel, 1971). Two specimens of Oedichirus simoni were collected from pine/oak litter in Turkey ( Assing, 2004: 683; 2006: 227). Many species are known from few specimens and often only one specimen is collected at a site.

Since so many species of Oedichirus are flightless it would seem unlikely to find species in trees or bushes and in fact many specimens were taken from ground litter (see Fagel, 1971). However, Borys Malkin, who for several decades travelled to many remote sites and villages of indigenous peoples in Central and South America after the Second World War, collected 10 specimens, all flightless, of three species of Oedichirus in Brazil by sweeping vegetation.

SYNONYMY: Oedichiranus was originally described as and has remained a subgenus. The characters for Oedichiranus listed by Reitter (1906: 264) to distinguish the subgenus from Oedichirus include long elytra, welldeveloped humeral angles, and a deep depression adjacent to the suture. These features are present in species with fully developed elytra and those species presumably can fly. The depth of the sutural depression and the height of the sutural ridge vary. Examples of species with a shallow to moderately developed sutural depression and sutural ridge include O. arrowi , O. dollmani , O. idae , O. lewisius , O. segmentarius , O. strandi , and O. zumpti ; included among examples of species with a strongly developed depression and sutural ridge are O. latipennis , O. longipennis , O. oneili , O. pendleburyi , O. reitteri , and O. rhodesianus . Most species with poorly developed humeral angles lack the sutural depression, or it is poorly developed and the sutural ridge is indistinct or weakly developed. Other than species diagnostic features, no characters unrelated to the possession of wings can be found that distinguish O. reitteri from other species of Oedichirus , so Oedichiranus is herewith synonymized with Oedichirus .

Elytrobaeus was described for one Brazilian species by Sahlberg (1847: 801, 802). Fauvel (1875a: 219; 1875b: xvii) and Sharp (1876: 338) synonymized the name with Oedichirus ; Fauvel did so without comment. Sharp pointed out that the characters purporting to separate it from Oedichirus appeared to be ‘‘very indefinite.’’ Sharp further stated that Sahlberg placed emphasis ‘‘on the last joint of the antennae terminating in a spine.’’ To understand this spine Sharp studied specimens of Oedichirus paederinus and found that the terminal antennomere of two females was truncate whereas in the male it was terminated by a slender, long spine or seta. He also observed that the apex of the antennomere appears to be membranous and that the spine can be retracted and so the antennomere is apparently truncate. Sharp’s observations are basically correct. Although the ‘‘spine’’ is actually a compact, spiniform pencil of stout, flat setae (fig. 31), it is found in both sexes of all species of Oedichirus and, in dried specimens, is retracted into the apex of the antennomere when the dried apex of the antennomere collapses. The length of the pencil may vary. This spiniform pencil is unique to Oedichirus and supports the monophyly of the genus and the synonymy of Elytrobaeus with Oedichirus .

DISCUSSION: Nearly 98% of the 303 described species of Oedichirus occur in the Old World. As a result of the intense work of Fagel (1971), Lecoq (1986), and Janák (2003), over 80% of the species are found in subsaharan Africa (147 species) and Madagascar (107 species). The remaining Old World species are in southern Europe ( Spain, Italy, and Greece) and North Africa (six species), the western edge of southwestern Asia (two species), India (four species), Sri Lanka (three species), Indochina and Indonesia (five species), China (two species), Japan (four species), Papua New Guinea (eight species), Australia (11 species), and New Caledonia (two species). Only one species is in central Asia ( Tajikistan) and two in China ( Taiwan, Zhejiang). The four species reported in southern Europe also occur in North Africa. It is probable that the paucity of species outside Africa is an artifact of collecting and that with additional collecting more, perhaps many more, species will be found in all areas except temperate ones. New Caledonia has only two described species, but there are at least six others. Only eight species are known for all of the New World, but there are at least 12 others among a limited number of unidentified specimens.

Certainly more species are to be found for all the tropical and subtropical regions. However, with the exception of Europe, Africa, and Madagascar, the species from the remainder of the world need revision. Although the New World species are being revised (Herman, in prep.), few specimens from few localities are available, so many more species await discovery. Species continue to be described from Madagascar and undoubtedly more will be found in Africa.

Whether more species will be found at the periphery of the distribution of the genus is an open question, so these edges are discussed below. The Oedichirus fauna for most of the world is poorly understood and most identifications beyond the type series are suspect and need verification. The preponderance of flightless species in the genus makes likely a large number of species with restricted distributions.

Four North African species are in southern Europe in Italy ( O. oedypus and O. paederinus ), Spain ( O. paederinus and O. unicolor ), and Greece ( O. terminatus ). Two other species, O. pardoi Outerelo and Gamarra from Morocco, and O. simoni Eppelsheim from Israel, Lebanon, and Turkey; additional collecting may extend the range of these species. The species in Europe are the northern edge of the range of North African species.

The northernmost central Asian record for the genus is in Tajikistan ( O. reitteri ); the species also occurs in Afghanistan, Turkey, and Cyprus.

Five species are reported from Japan ( O. chapmani , O. idae , O. kiuchii , O. kuroshio , and O. lewisius ). Only O. lewisius is recorded from Korea. For Japan the species are reported from the southernmost islands (Iriomote and Ishigaki; Hayashi, 1989) north to Kyushu, Shikoku, and southern Honshu and various islands between. Hayashi (1989) identified specimens from southern islands of Japan as O. chapmani Cameron , a species originally reported from Vietnam. Nothing in the discussion or redescription of O. chapmani suggests that Hayashi studied the type of the species, or had available reliably identified material. Furthermore, Cameron’s description of O. chapmani , which was based on one specimen, is terse, including no illustrations, no description of the aedeagus or male external secondary sexual characteristics, nor even a statement as to the sex of the individual. For the preceding reasons and because of the great distance between Japan and Vietnam, records of O. chapmani from Japan need verification.

For China only O. flammeus (Zhejiang) and O. kuroshio (Taiwan) are known; the latter species is also in Japan and the former is the only record of the genus on mainland China. Given that most species of the genus occur in tropical and subtropical regions, additional species might be found in such areas of China.

Eleven species are reported from Australia and, as one would expect, all are along the periphery of the continent. Based only on the type localities, six species are from the tropical and subtropical regions of the north in Queensland ( O. cribricollis , O. cribriventer , O. grandis , O. intricatus , and O. paederoides ) and Western Australia ( O. terminalis ). Four are from the more temperate south in South Australia ( O. andersoni ), New South Wales ( O. pictipes ), Victoria ( O. cribripennis ), and Victoria and Tasmania ( O. tricolor ). The type locality of the eleventh species ( O. rubricollis ) is simply Australia.

Five Australian species ( O. cribricollis , O. cribripennis , O. cribriventer , O. pictipes , and O. terminalis ) are known only from the type locality. The remaining species are known from additional sites. The Australian species have not been revised and since perhaps 90% of the species of the genus are flightless and tend to have restricted distributions, the same is likely to be true of the Australian species. Some species reported from multiple, widely separated localities may be misidentified. For example, the syntypes of O. tricolor Lea are reported from both Victoria and Tasmania, but the species is stated to be apterous ( Lea, 1904), so the syntypes may be different species. Oedichirus andersoni Blackburn is listed from both Western Australia (Swan River, which is near Perth) ( Lea, 1904: 63) and South Australia (Port Lincoln) ( Blackburn, 1888: 10); these localities are about 1800 km apart, so if O. andersoni is flightless then these specimens may be different species. Oedichirus paederoides is said to be from southern Queensland (Gayndah, NNW of Brisbane) ( MacLeay, 1873: 147) and from New South Wales (Clarence River); the species is said to be winged, so these records, which are separated by about 480 km, may be correct. The Australian species are in great need of revision.

All the New World species are restricted to the Neotropical Region, but only eight have been described, one from Costa Rica, the others from Brazil. The poverty of species is the result of the lack of collecting. In a forthcoming article 10 new species will be described from Brazil, Bolivia, Peru, Ecuador, Mexico, and Dominican Republic (Herman, in prep.), but all are represented by few specimens and for most species only one locality is known. Collections of more specimens from more localities will result in discovery of many more new Neotropical species.

As is found in species of a number of staphylinid genera, some Oedichirus mimic a color pattern seen in many species of Paederus in which the head, elytra, and abdominal apex are black or metallic dark blue and the remaining regions are orange to reddish orange. Among some of the Oedichirus species with this color pattern are: O. arrowi , O. bicolor , O. cameronianus , O. desaegerianus , O. eppelsheimianus , O. fortepunctatus , O. garambanus , O. idae , O. katanganus , O. kivuensis , O. lamotteanus , O. latipennis , O. leleupianus , O. lewisius , O. longipennis , O. nimbaensis , O. obscuripes , O. oneili , O. paederinus , O. problematicus , O. reitteri , O. rhodesianus , O. segmentarius , O. strandi , O. terminatus , O. uelensis , O. villiersi , and O. zumpti . All but four of these species are from Africa; two are from Japan, one is from India, one is from Central Asia; one of the African species is also found in southern Europe. All but one of these species, O. paederinus , have fully developed elytra and may be capable of flight. All of the species fall into ‘‘Section III’’ (species lacking the pronotal marginal ridge) of the three groupings proposed by Fagel (1971: 129–130). There are certainly other species with this color pattern, but I recorded this feature inconsistently and have not seen all the species. According to Coiffait (1978b: 330, 332) two species from the Mediterranean Region, O. oedypus and O. simoni , have this color pattern. I have seen no Australian species so colored, but the head and apical abdominal segments of O. paederoides are reported to be black and the basal three abdominal segments reddish. The color of the elytra was not stated ( MacLeay, 1873: 147), but the name is suggestive and the species may fly. As yet no New World species are known that mimic the Paederus color pattern, although O. neotropicus is close.

Few keys to species have been published. Fagel (1971) published a key to the sub- Saharan African species. Lecoq (1986) published another for the species known to then for Madagascar, but many others have been described since. Coiffait (1978b: 329–330) published one for the six species of the Western Palaearctic Region and Cameron (1931: 25) for eight Indian species.

SPECIES INCLUDED AND

MATERIAL EXAMINED

abdominalis Boheman – sp ( FMNH)........

......................... South Africa abyssinicus Fagel – Lit. Att. [ Ethiopia] aethiopopygus Fagel – H ( MRAC)... D.R. Congo alatus Nietner – sp ( BMNH)....... Sri Lanka ampamoho Janák – Lit. Att. [ Madagascar]

analis Lecoq – Lit. Att. [ Madagascar]

andapanus Lecoq – Lit. Att. [ Madagascar] andersoni Blackburn – Syn (BMNH)... Australia andringitra Janák – Lit. Att. [ Madagascar] andringitranus Jarrige – Lit. Att. [ Madagascar] angavokeliensis Lecoq – Lit. Att. [ Madagascar] angolensis Fagel – H (MRAC)........ Angola anosibensis Lecoq – Lit. Att. [ Madagascar] anosyanus Lecoq – Lit. Att. [ Madagascar] antitra Janák – Lit. Att. [ Madagascar] anularis Lecoq – Lit. Att. [ Madagascar] appendiculatus Lecoq – Lit. Att. [ Madagascar] arrowi Bernhauer – Syn (FMNH)..... Zambia bacillus Fagel – H (MRAC)...... D.R. Congo balarombe Janák – Lit. Att. [ Madagascar] balazuci Lecoq – Lit. Att. [ Madagascar] baloghi Last – Lit. Att. [ Papua New Guinea] bambusicola Fagel – H (MRAC)....... Kenya bara Janák – Lit. Att. [ Madagascar] barbertonensis Fagel – P (MRAC)... South Africa basilewskyanus Fagel – H (MRAC)... Rwanda beltermanni Bernhauer – H, sp (FMNH).....

.......................... Cameroon bertiae Lecoq – Lit. Att. [ Madagascar] betschi Jarrige – Lit. Att. [ Madagascar] betsileo Janák – Lit. Att. [ Madagascar] bicolor Fagel – H (MRAC)...... D.R. Congo bifidus Lecoq – Lit. Att. [ Madagascar] biguttatus Fauvel – Lit. Att. [ Mozambique] birmanus Fauvel – Syn (IRSN)..... Myanmar biroi Last – Lit. Att. [ Papua New Guinea] blukwaensis Fagel – H (MRAC).... D.R. Congo boehmi Bernhauer – H, sp (FMNH)..... Brazil bonibona Janák – Lit. Att. [ Madagascar] bonsae Jarrige – Lit. Att. [ Madagascar] brachelytratus Lecoq – Lit. Att. [ Madagascar] brunneicolor Fagel – H (MRAC).... D.R. Congo brunneus Wendeler – Syn (MNHB)..... Brazil bulirschi Janák – Lit. Att. [ Madagascar] burgeoni Bernhauer – Syn (FMNH, MRAC)...

......................... D.R. Congo cameronianus Fagel – H (MRAC)... D.R. Congo camerounensis Fagel – Lit. Att. [ Cameroon] capensis Fagel – P (MRAC)..... South Africa capicola Fagel – P (MRAC)..... South Africa carayoni Lecoq – Lit. Att. [ Madagascar] carolinorum Janák – Lit. Att. [ Madagascar] celisianus Fagel – H (MRAC).... D.R. Congo chapmani Cameron – H (BMNH).... Vietnam clementi Lecoq – Lit. Att. [ Madagascar] congoensis Bernhauer – Syn (FMNH)........

......................... D.R. Congo conspicuus Fagel – H (MRAC)... D.R. Congo crebrepunctatus Fagel – H (MRAC).........

......................... D.R. Congo cribricollis Lea – Lit. Att. [ Australia] cribripennis Lea – Lit. Att. [ Australia]

geniculatus Lea cribriventer Lea – Lit. Att. [ Australia] curticornis Fagel – Lit. Att. [D.R. Congo] curtipennis Fagel – H (MRAC)... D.R. Congo curtulus Lecoq – Lit. Att. [ Madagascar] densoides Fagel – Lit. Att. [ South Africa] densus Bernhauer – H (FMNH)... South Africa denticulatus Lecoq – Lit. Att. [ Madagascar] depressipennis Lecoq – Lit. Att. [ Madagascar] desaegerianus Fagel – H (MRAC)... D.R. Congo descarpentriesi Jarrige – Lit. Att. [ Madagascar] didyanus Janák – Lit. Att. [ Madagascar] dimidiatus Eppelsheim – Lit. Att. [ India]

elegans Cameron dollmani Bernhauer – Syn (FMNH).... Zambia dubius Jarrige – Lit. Att. [ Madagascar] duflosi Lecoq – Lit. Att. [ Madagascar] dunayi Janák – Lit. Att. [ Madagascar] elgonensis Fagel – H (MRAC)......... Kenya ensifer Lecoq – Lit. Att. [ Madagascar] epiphyticola Janák – Lit. Att. [ Madagascar] eppelsheimianus Fagel – H (MRAC)...... Mali ericeticola Fagel – H (MRAC).... D.R. Congo excellens Cameron – sp (BMNH).... Indonesia fageli Lecoq – Lit. Att. [ Madagascar] femoralis Lecoq – Lit. Att. [ Madagascar] filicornis Fagel – H (MRAC)..... D.R. Congo fitorahana Janák – Lit. Att. [ Madagascar] flammeus Koch – Lit. Att. [ China] flavifrons Fagel – H (MRAC).... D.R. Congo flavipes Lecoq – Lit. Att. [ Madagascar] foveicollis Quedenfeldt – sp (FMNH)........

......................... Madagascar fortepunctatus Fagel – H (MRAC)... D.R. Congo franzi Lecoq – sp (BMNH)....... Madagascar furcatus Lecoq – Lit. Att. [ Madagascar] garambanus Fagel – H (MRAC).... D.R. Congo geniculatus (R. Sahlberg) – sp (FMNH)... Brazil gladiatus Lecoq – sp (BMNH)..... Madagascar gracilis Fagel – H (MRAC)...... D.R. Congo grandis Bernhauer – Syn (FMNH).... Australia graskopensis Fagel – P (MRAC)... South Africa griveaudi Lecoq – Lit. Att. [ Madagascar] hammondi Lecoq – H (BMNH).... Madagascar hanglipbosensis Fagel – Lit. Att. [ South Africa] haribe Janák – Lit. Att. [ Madagascar] hewitti Bernhauer – H (FMNH)... South Africa histrio Lecoq – Lit. Att. [ Madagascar] humansdorpensis Fagel – P (MRAC)........

......................... South Africa humicola Fagel – Lit. Att. [ South Africa] idae Sharp – Syn (BMNH), sp (FMNH)... Japan ingogoensis Fagel – Lit. Att. [ South Africa] insolitus Fagel – H (MRAC)..... D.R. Congo insularis Lecoq – Lit. Att. [ Madagascar] intermixtus Fagel – H (MRAC)... D.R. Congo intricatus Fauvel – H (IRSN)........ Australia itombwensis Fagel – H (MRAC).... D.R. Congo ivohibensis Jarrige – Lit. Att. [ Madagascar] janae Janák – Lit. Att. [ Madagascar] jarrigei Lecoq – Lit. Att. [ Madagascar] jenisi Janák – Lit. Att. [ Madagascar] jocquei Lecoq – Lit. Att. [ Comoros] kaboboensis Fagel – H (MRAC).... D.R. Congo kahololoensis Fagel – H (MRAC)... D.R. Congo kahuziensis Fagel – H (MRAC)... D.R. Congo

incertus Fagel [ssp] – H (MRAC).. D.R. Congo kalamba Janák – Lit. Att. [ Madagascar] kalehensis Fagel – H (MRAC).... D.R. Congo katanganus Fagel – H (MRAC)... D.R. Congo katbergensis Fagel – H (BMNH), P (MRAC)..

......................... South Africa keanus Fagel – Lit. Att. [D.R. Congo] kidundaensis Fagel – H (MRAC)..... Tanzania kilimanjarensis Fagel – H (MRAC)... Tanzania kimbiensis Fagel – H (MRAC).... D.R. Congo kirunguensis Fagel – H (MRAC).. D.R. Congo kiuchii Sawada – Lit. Att. [ Japan] kivuensis Fagel – H (MRAC)..... D.R. Congo kolbei Fauvel – L, Pl (IRSN), sp (BMNH)....

......................... Madagascar kolwezienus Fagel – Lit. Att. [D.R. Congo] kuroshio Hayashi – Lit. Att. [ Japan, Taiwan] kyandolirensis Fagel – H (MRAC)... D.R. Congo lamotteanus Fagel – sp (MRAC)...... Guinea laticeps Lecoq – Lit. Att. [ Madagascar] latipennis Bernhauer – Syn (FMNH)... Zambia lecoqi Janák – Lit. Att. [ Madagascar] leleupianus Fagel – H (MRAC)... D.R. Congo levasseuri Lecoq – Lit. Att. [ Madagascar] lewisius Sharp – H (BMNH), sp (FMNH)....

.............................. Japan loksai Last – Lit. Att. [ Papua New Guinea] longicornis Lecoq – Lit. Att. [ Madagascar] longipennis Kraatz – L (SDEI), sp (FMNH)...

.............. ‘‘ India orientali’’, Indonesia longipilis Fagel – H (MRAC)..... D.R. Congo luberensis Fagel – H (MRAC), sp (BMNH)...

......................... D.R. Congo

kalongensis Fagel [ssp] – H (MRAC).......

......................... D.R. Congo

obscurellus Fagel [ssp] – H (MRAC).......

......................... D.R. Congo

nigrinus Fagel [ssp] – Lit. Att. [D.R. Congo]

kenyacus Fagel [ssp] – H (MRAC).... Kenya

imitator Fagel [ssp] – H (MRAC)..... Kenya lucidiceps Fagel – P (MRAC).... South Africa lugubris Fagel – H (MRAC)..... D.R. Congo luikoensis Fagel – H (MRAC).... D.R. Congo luvubuensis Fagel – H (MRAC)... D.R. Congo lwiroensis Fagel – H (MRAC).... D.R. Congo madagascariensis Bernhauer – H (FMNH)....

......................... Madagascar madegassus Fagel – Lit. Att. [ Madagascar] mafana Janák – Lit. Att. [ Madagascar] magnus Last – Lit. Att. [ Papua New Guinea] mahasoa Janák – Lit. Att. [ Madagascar] malalaka Janák – Lit. Att. [ Madagascar] mandibularis Lecoq – Lit. Att. [ Madagascar] mangabensis Lecoq – Lit. Att. [ Madagascar] manarivo Janák – Lit. Att. [ Madagascar] mariepskopensis Fagel – P (MRAC).........

......................... South Africa mavo Janák – Lit. Att. [ Madagascar] melanurus Eppelsheim – Lit. Att. [ Cape Verde

Islands, Guinea, Ghana, Ethiopia, Tanzania,

Namibia] meruensis Fagel – H (MRAC)....... Tanzania microcephalus Fagel – P (MRAC)... South Africa microphthalmus Fagel – Lit. Att. [ Kenya] minimus Bernhauer –Syn (FMNH)... South Africa minor Cameron – Syn (BMNH)..... Sri Lanka minutus Fagel – P (MRAC)...... South Africa miskoi Janák – Lit. Att. [ Madagascar] mokotoensis Fagel – H (MRAC)... D.R. Congo moloensis Fagel – H (MRAC)......... Kenya montanus Last – Lit. Att. [ Papua New Guinea] monticola Fagel – H (MRAC).... D.R. Congo

surdus Fagel [ssp] – H (MRAC). D.R. Congo montishoyoensis Fagel – H (MRAC).........

......................... D.R. Congo moraveci Janák – Lit. Att. [ Madagascar] muscorum Jarrige – Lit. Att. [ Madagascar] mwengensis Fagel – H (MRAC)... D.R. Congo natalensis Fagel – Lit. Att. [ South Africa] neotropicus Blackwelder

pictipes Bierig – Syn (FMNH)... Costa Rica newtonianus Fagel – H (MRAC)... South Africa niger Cameron – Syn (BMNH), Syn, sp (FMNH)

.............................. India nigriceps Fagel – H (MRAC)..... D.R. Congo nigrolineatus Lecoq – Lit. Att. [ Madagascar] nimbaensis Fagel – sp (MRAC).... Ivory Coast nitidiventris Fagel – Lit. Att. [ Kenya] nodieri Lecoq – Lit. Att. [ Madagascar] nordicus Lecoq – sp (BMNH)..... Madagascar nosykombanus Lecoq – Lit. Att. [ Madagascar] novaeguineae Wendeler – Lit. Att. [ Papua New

Guinea] novus Jarrige – Lit. Att. [ Madagascar] noyesi Lecoq – H (BMNH)....... Madagascar nyakageraensis Fagel – H (MRAC)... D.R. Congo nyalengwensis Fagel – H (MRAC)... D.R. Congo obscuripes Fagel – H (MRAC).... D.R. Congo obscurus Fagel – P (MRAC)..... South Africa obsoletus Lecoq – Lit. Att. [ Madagascar] occipitopunctatus Lecoq – Lit. Att. [ Madagascar] oedypus Rottenberg – Syn (SDEI)....... Italy

walkeri Fauvel ohausi Wendeler – Syn (MNHB)....... Brazil oldeaniensis Fagel – H (MRAC)...... Tanzania omoanus Fagel – P (MRAC).......... Kenya oneili Péringuey – sp (FMNH)..... Zimbabwe? opacus Lecoq – Lit. Att. [ Madagascar] optatus Sharp – Syn (BMNH), sp (FMNH)...

.............................. Brazil orophilus Fagel – H (MRAC).... D.R. Congo

proximus Fagel [ssp] – H (MRAC)........

......................... D.R. Congo oundaensis Last – Lit. Att. [ New Caledonia] paederinus Erichson – sp (FMNH)..........

...... Algeria, Italy, Morocco, Spain, Tunisia

quedenfeldtii Schaufuss paederoides W.J. MacLeay – Lit. Att. [ Australia] papanganus Lecoq – Lit. Att. [ Madagascar] papuanus Cameron – Syn (BMNH).........

.................... Papua New Guinea pardoi Outerelo and Gamarra – Lit. Att. [ Morocco] parviceps Fagel – Lit. Att. [ South Africa, Zanzibar] parvus Lecoq – Lit. Att. [ Madagascar] pauliani Lecoq – Lit. Att. [ Madagascar] pearceanus Fagel – H (MRAC)..... Zimbabwe pendleburyi Cameron – Syn (BMNH), sp (FMNH)

.................... Malaysia, Singapore penicillatus Jarrige – Lit. Att. [ Madagascar] pictipes Oke – Lit. Att. [ Australia] pietersburgensis Fagel – P (MRAC).........

......................... South Africa problematicus Fagel – H (MRAC)..........

......................... D.R. Congo profundepunctatus Fagel – H (MRAC).......

......................... D.R. Congo pubescens Lecoq – Lit. Att. [ Madagascar] puguensis Bernhauer – H, sp (FMNH).......

........................... Tanzania pumilus Lecoq – Lit. Att. [ Madagascar] puncticollis Fagel – H (MRAC)... D.R. Congo pyricollis Lea – Lit. Att. [ Papua New Guinea] radama Janák – Lit. Att. [ Madagascar] ranavalona Janák – Lit. Att. [ Madagascar] ranomafanus Janák – P (MNHB)... Madagascar reitteri Bernhauer – Syn, sp (FMNH)........

....................... Iraq, Tajikistan

dimidiatus Reitter reticulatus Fagel – H (MRAC).... D.R. Congo rhodesianus Bernhauer – Syn, sp (FMNH), sp

(MRAC)........... Zambia, D.R. Congo

elegantulus Fagel [ssp] – H (MRAC).......

......................... D.R. Congo ruandaensis Fagel – H (MRAC)...... Rwanda rubricollis Fauvel – H (BMNH)...... Australia ruficeps Kraatz – L (SDEI)... ‘‘ India orientali’’ rufitarsis Fauvel – H (IRSN)...... Madagascar rufotestaceus Bernhauer – H (FMNH)... Sri Lanka

(Note: Holotype missing head and prothorax.) rufus Fauvel – Lit. Att. [ Zanzibar] rugegensis Fagel – H (MRAC)....... Rwanda ruteri Lecoq – Lit. Att. [ Madagascar] ruwenzoricus Fagel – H (MRAC)... D.R. Congo sambavanus Lecoq – Lit. Att. [ Madagascar] sanctamariae Lecoq – Lit. Att. [ Madagascar] satyrus Lecoq – Lit. Att. [ Madagascar] schultheissi Fauvel – Syn (IRSN),sp (SDEI)...

........................... Indonesia sedilloti Fauvel – Syn (IRSN)... New Caledonia segmentarius Bernhauer – Pl, sp (FMNH), L

(MRAC)................. D.R. Congo

(Note: Fagel [1971: 417] cited the specimen he

examined from Ituri as ‘‘ type,’’ but it is a

lectotype designation under ICZN, 1999: Article

74.5.) senegalensis (Laporte) – Lit. Att. [ Senegal, Cape

Verde Islands] serrulatus Lecoq – Lit. Att. [ Madagascar] silvestris Lecoq – Lit. Att. [ Madagascar] similis Fagel – H (MRAC)...... D.R. Congo simillimus Fagel – Lit. Att. [ South Africa] simoni Eppelsheim – Lit. Att. [ Israel, Lebanon,

Turkey] simplex Lecoq – Lit. Att. [ Madagascar] sogai Jarrige – Lit. Att. [ Madagascar] sparsepunctatus Fagel – H (MRAC).........

......................... D.R. Congo sparsipennis Bernhauer – Syn (FMNH)... Brazil sparsutus Fagel – H (MRAC)....... Rwanda spectabilis Fagel – H (MRAC).... D.R. Congo speculifrons Bernhauer – Syn, sp (FMNH)....

.............................. Brazil splendidus Lecoq – Lit. Att. [ Madagascar] stilicinus Gerstaecker – Lit. Att. [ Ethiopia,

Tanzania, Zanzibar]

ignicollis Fauvel strandi Bernhauer – H (FMNH)..... Tanzania strictus Fagel – H (MRAC)...... D.R. Congo subdensus Fagel – Lit. Att. [ South Africa] summicola Fagel – H (MRAC)...... Tanzania tenuis Fagel – P (MRAC)....... South Africa terminalis Lea – sp (BMNH)........ Australia terminatus Erichson – Syn (MNHB), sp

(BMNH)........... Nigeria, Sierra Leone

rubronotatus Pic testaceus Fagel – Lit. Att. [ South Africa] tigrinus Lecoq – Lit. Att. [ Madagascar] transvaalensis Fagel – P (MRAC).... South Africa triangulipennis Fagel – H (MRAC)... D.R. Congo tricolor Lea – sp (FMNH, BMNH, SDEI).......

................................... Australia tronqueti Lecoq – Lit. Att. [ Madagascar] tshiaberimuensis Fagel – H (MRAC)........

......................... D.R. Congo tshuruyagaensis Fagel – H (MRAC)... Rwanda turneri Bernhauer – Syn (FMNH)... South Africa uelensis Fagel – H (MRAC)..... D.R. Congo uhligi Janák – Lit. Att. [ Madagascar] uluguruensis Fagel – H (MRAC)..... Tanzania uncinatus Lecoq – Lit. Att. [ Madagascar] unicolor Aubé – sp (FMNH, SDEI)..... Spain unicus Fagel – H (MRAC)......... Tanzania uniformis Fagel – H (MRAC).... D.R. Congo usambarae Bernhauer – H (FMNH), sp (MRAC)

........................... Tanzania uviraensis Fagel – H (MRAC).... D.R. Congo vadoni Lecoq – Lit. Att. [ Madagascar] variabilis Fagel – H (MRAC).... D.R. Congo variegatus Fagel – sp (BMNH, MRAC).......

.......................... South Africa variipennis Fagel – H (MRAC)... D.R. Congo varius Lecoq – Lit. Att. [ Madagascar] vaovao Janák – Lit. Att. [ Madagascar] ventralis Fauvel – Lit. Att. [ Madagascar] villiersi Cameron – Pl (BMNH), sp (MRAC)...

......................... Senegal, Chad vohitrosa Janák – Lit. Att. [ Madagascar] witteanus Fagel – H (MRAC).... D.R. Congo woodbushensis Fagel – P (MRAC)... South Africa yangambiensis Fagel – H (MRAC)... D.R. Congo zanzibaricus Fagel – Lit. Att. [ Zanzibar] zumpti Bernhauer – Syn (FMNH)... Cameroon

UNDETERMINED SPECIMENS: Old World: Australia, Borneo, China, Indonesia, Madagascar, Malaysia, New Caledonia, Philippines, Zambia, Zimbabwe, Singapore, South Africa, Zaire.

New World: Brazil, Costa Rica, Ecuador, Mexico, Peru.

DISSECTIONS: Complete dissection: Oedichirus nr. pictipes (1 male, 1 female); Oedichirus geniculatus (1 male, 1 female).