Oedodactylus Fairmaire and Germain

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 : 46-62

Oedodactylus Fairmaire and Germain Figures 4, 77–89

Oedodactylus Fairmaire and Germain, 1861: 441 . Type species: Oedodactylus fuscobrunneus Fairmaire and Germain, 1861: 441 , fixed by R. Lucas (1920: 452) by subsequent designation when he cited only one of the two originally included species.

— Fauvel, 1867: 2, 16 (key; characters). — Fauvel, 1868: 7, 21 (key; characters). — Gemminger and Harold, 1868: 629 (catalog; world). — Sharp, 1876: 337 (notes; Brazilian species). — Duvivier, 1883: 176 (catalog). — Sharp, 1886: 630 (Mexican species). — Bernhauer and Schubert, 1912: 198 (catalog; world). — Bruch, 1915: 493 (catalog; Argentina). — Bruch, 1928: 440 (catalog; Argentina). — Scheerpeltz, 1933: 1213 (catalog; world). — Blackwelder, 1944: 130 (checklist; Latin America). — Blackwelder, 1952: 270 (type species). — Coiffait and Saiz, 1968: 369 (characters; Chilean species; cited as Aedodactylus). — Navarrete- Heredia et al., 2002: 293 (characters; notes; list of Mexican species).

DIAGNOSIS: Oedodactylus can be separated from all other Procirrina except Pseudoprocirrus by the inflated basal three protarsomeres (fig. 89); the basal four are swollen in other genera (fig. 92). Both Oedodactylus and Pseudoprocirrus have a paratergite on each

80. Prothorax, ventral. 81. Maxillary palpus. 82. Sternum IV, base. 83. Labium. 84. Labrum.

side of abdominal segment III. Oedodactylus lacks an interantennal, transverse frontoclypeal ridge, the gular sutures are most approximate just proximad of the middle (fig. 78), tergum and sternum VIII have a transverse, basal ridge, and terga IX and X are fused (fig. 87). Pseudoprocirrus has a transverse, interantennal, frontoclypeal ridge, the gular sutures are most approximate medially, tergum and sternum VIII lack a transverse, basal ridge, and terga IX and X are separate (fig. 103). Oedodactylus is a New World tropical genus and Pseudoprocirrus is African. Some species of Oedichirus have paratergites on abdominal segment III, but in Oedichirus the tergum and sternum of segment VII are fused basally whereas in Oedodactylus they are separated. The pres- ence of setae on the edge of the posterior margin of the elytra will separate Oedodactylus from Procirrus , Oedichirus , Palaminus , and Paraprocirrus .

DESCRIPTION: Head not pedunculate, wider than long (fig. 77); frontoclypeus without subapical, interantennal, transverse ridge; postocular lateral margin broadly rounded to basal angles or to neck; postocular lateral margin moderately long; basal angle present and strongly rounded (fig. 77) or absent (fig. 4); basal margin of head truncate or indistinguishable from lateral margin, with or without marginal ridge, and with median portion feebly to moderately emarginate anterior to neck. Neck across nuchal constriction one half to two thirds of greatest postocular width of head; nuchal groove shallow to moderately deep; nuchal ridge present (fig. 77). Eye length longer (fig. 77) to slightly shorter than postocular length of head. Dorsal surface with moderately dense to dense umbilicate punctation. Ventral surface without postocular groove (cf. fig. 2). Gular sutures (fig. 78) separated and present basally; sutures most approximate just proximad of middle. Gula without pubescence. Antennomere 11 about as long as or slightly longer than 10; apex without spinelike pencil of setae. Mandibles with apically bifid denticle (fig. 79); prostheca digitiform apically and with cluster of cuticular processes medially (fig. 79). Maxillary palpus (fig. 81) with palpomere 4 longer than second or third, securiform, broad apically, slightly longer than wide, and compressed. Labium (fig. 83) with glossae separated, apically rounded, long, broad, and with a few cuticular processes on medial margin. Hypopharynx without spinelike setae or lobes on anterior margin; laterodorsal margin with dense cluster of setae; median groove present. Labrum without denticles on anterior margin, but with broad lobe adjacent to emargination (fig. 84). Epipharynx with dense, transverse cluster of cuticular processes across anterior portion and with five setae on lateral portion near anterior margin; median groove present; epipharynx slightly visible anterolaterally in dorsal view.

Prothorax rounded trapezoidal, as long as (fig. 77) to longer than wide (fig. 4); widest near anterior third and with lateral margins rounded and gradually convergent anteriad and posteriad from widest point. Pronotum with umbilicate punctation; punctation dense to moderately dense, uniform, and present on midline in most species, but absent in one. Pronotal marginal ridge present and entire. Notosternal suture absent. Hypomeron with fine microsculpturing and with a few punctures in some species. Postprocoxal lobe long and with a few to many punctures; transverse ridge present or absent; apices widely separated from each other. Probasisternum with median carina present (fig. 80) or absent; surface punctate. Mesospiracular peritreme (fig. 80) with anteromedial margin fused to furcasternum. Elytra longer than pronotum; humeral angles present; posterior edge with row of setae; subapical region without long, thicker, posteromediad directed seta near lateroapical corner. Scutellum with a few setae. Mesosternum with short median carina extending from mesosternal process. Mesocoxal acetabulum with marginal carina laterally and posteriorly. Intersternal suture present and well developed.

Profemur with rounded ridge on proximal half of anteroventral surface. Protibia with multiple (11 in a Venezuelan species) combs extending from near base to tibial apex; tibia with feeble depression on ctenidial surface; apical portion not constricted or enlarged. Protarsomeres 1–3 inflated (fig. 89); base of tarsomere 1 not surrounded by cupulate protibial apex; tarsomere 4 not expanded beneath tarsomere 5, apical margin entire, not bilobed, and with setae ventrally, but without setose pad; tarsomere 4 inserted laterad of middle of apex of asymmetrical third segment; tarsomere 5 sparsely pubescent ventrally. Mesotarsomere 1 longer than 2. Metatarsomere 1 longer than 2–4 combined.

Abdominal segments without imbricate macrosculpturing (cf. fig. 28). Segment III with one paratergite; paratergal carina absent; sternum and tergum III separated by paratergite. Segment VII with tergum and sternum separated. Segments IV to VII without oval ‘‘windows’’ in intersegmental membrane. Sternum I present, lightly sclerotized, and medially divided. Sterna IV (fig. 82) through VII with small, median, membrane covered, possibly glandular, slit on anterior margin. Tergum IX with transverse base narrow and fused middorsally (fig. 87); lateroapical process of IX (fig. 85, 87) long, slender, gradually curved ventrally, and extending well beyond apex of tergum X; lateroapical process fused basally to remain- der of segment. Tergum X with base fused to base of tergum IX (fig. 87); apical margin attenuate, long, slender, and narrowly round- ed apically.

Female genital sclerites separated medially; proximal gonocoxite long, broad, apically acute, and separated from short, setate, apically rounded distal gonocoxite (fig. 86).

Aedeagus (fig. 88) slightly asymmetrical; parameres absent; basal piece absent.

DISTRIBUTION AND HABITAT: Oedodactylus is a Neotropical genus collected at scattered localities from Mexico (Morelos and Veracruz States) south to Argentina (Buenos Aires Province) and Chile (Santiago District). I have seen specimens from Mexico, Costa Rica, Venezuela, Guyana, Brazil, Paraguay, Argentina, and Chile, but few specimens from any one place and few in total.

Almost nothing has been reported about the habitat of the species and the species are rarely collected. Oedodactylus fuscobrunneus was reported to have been collected from under stones in humid places ( Fairmaire and Germain, 1861: 442) and, according to the locality labels, a few specimens of an unnamed Venezuelan species (near anceps ) were collected at lights.

DISCUSSION: Oedodactylus is the only genus in the subtribe with terga IX and X fused (fig. 87). This fusion is unusual in the subfamily.

The similarity of some species of Oedodactylus and those of Pseudoprocirrus is remarkable, but features of abdominal segments III, IX, and X distinguish them. I dissected only the apical segments of the abdomen of a female of Pseudoprocirrus arrowi and only three specimens of two species of Oedodactylus . Nonetheless, these few dissections reveal tergum IX of P. arrowi to be fused midbasally, the lateroapical processes to be attached to IX despite an incision (fig. 104), and tergum X to be fully separated from IX (fig. 103). Fagel (1971: 49) wrote that Pseudoprocirrus abyssinicus possesses a single paramere; he had no males of P. arrowi . I have insufficient material of the genus to corroborate Fagel’s observation and was unable to study his dissection of P. abyssinicus .

The largest, most robust species in the genus is O. fuscobrunneus (fig. 4), but the species also differs in other ways. The cephalic and pronotal punctation is not as coarse as for other species, the postocular lateral margin of the head is gradually curved to the neck, the basal angles of the head are absent, the pronotum is long, and the anterior angles are broadly rounded; the neck is about two thirds as wide as the postocular width of the head, but for other species the neck is about half the width. The basal angles of the head and the anterior angles of the pronotum are acute in most species, but rounded in O. fuscobrunneus . Oedodactylus fauveli is similar to O. fuscobrunneus , but the anterior pronotal angles of the former are moderately angulate and the pronotal midline is impunctate.

Revision of the genus is needed, in part to assess the variation described above. Only four species are currently in the genus, but among the few specimens I examined there appear to be additional ones.

SPECIES INCLUDED AND

MATERIAL EXAMINED

anceps Sharp – H ( BMNH)........... Brazil aper Sharp – Syn ( BMNH).......... Mexico errans Sharp , transferred to Stylokyrtus

fauveli Sharp – Syn (BMNH, FMNH)... Mexico fuscobrunneus Fairmaire and Germain – L (IRSN), Pl? (BMNH), sp (FMNH)... Chile, Argentina (Note: It is unclear whether the specimen designated as lectotype is part of the original series. Coiffait and Saiz [1968: 370] designated the lectotype, cited it from Rancagua, and wrote that Fauvel [1868: 22] added the locality to the original locality of ‘‘Santiago.’’ These two cities are in different provinces now, but I don’t know whether, when the species was collected, both were in the province of Santiago. This discrepancy needs reconciliation.)

UNDETERMINED SPECIMENS: Argentina, Brazil, Costa Rica, Paraguay, Guyana, Venezuela.

DISSECTIONS: Disarticulation: Undetermined species near O. anceps (1 female, Venezuela) and abdominal dissection (1 male, Venezuela); Abdominal dissection Oedodactylus near fauveli (1 male, Paraguay).

Palaminus Erichson Figures 5, 10–13, 19–21, 26–29, 40–43, 90–102

Palaminus Erichson, 1839: 29 . Type species: Palaminus pilosus Erichson, 1840: 682 , fixed by Duponchel (1841: 57) by subsequent designation.

— Erichson, 1840: 681 (characters; first includ- ed species: pilosus , piceus , variabilis , testaceus ). — Lacordaire, 1854: 104 (characters; notes; list of species). — Kraatz, 1857: 666, 668 (notes; key). — LeConte, 1861: 66 (key). — LeConte, 1863: 25 (list of species; North America). — Gemminger and Harold, 1868: 629 (catalog; world). — Crotch, 1873: 33 (checklist; North America). — Sharp, 1876: 340 (notes; Brazilian species). — Fauvel, 1878b: 225 (characters; notes; New Guinea species). — Duvivier, 1883: 175 (catalog). — Lynch Arribálzaga, 1884: 327 (characters; notes; Argentina). — Sharp, 1886: 631 (notes; Mexico and Central America). — Fauvel, 1891: 95 (key; Venezuela). — Blackburn, 1891: 75 (notes). — Casey, 1910: 192, 197 (characters; notes; key North America). — Blatchley, 1910: 440, 441 (characters). — Bernhauer and Schubert, 1912: 198 (catalog; world). — Bruch, 1915: 493 (catalog; Argentina). — Leng, 1920: 100 (catalog; U.S.A.). — Cameron, 1921: 352, 370, 402 (characters; key and catalog; Singapore). — Lea, 1923: 12 (notes). — Cameron, 1925: 33, 106 (catalog; British India). — Bruch, 1928: 440 (catalog; Argentina). — Notman, 1929: 2 (key; world). — Cameron, 1931: 1, 20 (characters; key; India). — Scheerpeltz, 1933: 1213 (catalog; world). — Bierig, 1935: 31 (natural history notes). — Glick, 1939: 31 (high aerial capture). — Blackwelder, 1943: 229, 388 (characters; notes; type species). — Bierig, 1943: 154 (notes). — Blackwelder, 1944: 130 (checklist; Latin America). — Blackwelder, 1952: 187 (type species). — Adachi, 1955: 14 (characters; key). — Arnett, 1963: 243, 267 (characters; notes). — Fagel, 1971: 11, 52 (characters; type species; Africa). — Seevers, 1971: 84 (amber fossil; late Oligocene to early Miocene; Mexico). — Shibata, 1973: 42 (catalog; Taiwan). — Moore and Legner, 1974: 555 (characters). — Blackwelder and Arnett, 1974: 48 (checklist; North America; West Indies). — Bordoni, 1975: 418 (characters). — Moore and Legner, 1975: 151 (catalog; U.S.A.). — Shibata, 1977: 20 (catalog; Japan).

98. Mandible, right.

— Moore and Legner, 1979: 122 (characters; notes). — Spahr, 1981: 96 (references to specimens from amber and copal). — Frank, 1982: 27 (parasites). — Hammond, 1984: 203 (checklist; Borneo). — Frank, 1986: 367 (checklist; Florida). — Lecoq, 1986: 81 (characters; discussion). — Biswas and Biswas, 1995: 266 (key to species of West Bengal). — Downie and Arnett, 1996: 424 (characters; key to species of Northeastern North America). — Lundgren, 1998: 48 (list of species from Florida; records for other States listed for each species). — Newton et al., 2000: 389 (characters in key; notes; U.S.A. and Canada). — Navarrete-Heredia et al., 2002: 294 (characters in key to genera for Mexico; general notes; list of species; Mexico). — Smetana, 2004: 624 (Palaearctic catalog).

Parapalaminus Bierig, 1943: 155 . Type species: Palaminus symphylus Bierig, 1943: 155 , fixed by original designation. New Synonym.

— Blackwelder, 1952: 292 (type species; subgenus of Palaminus ). — Blackwelder and Arnett, 1974: 48 (synonym of Palaminus ). — Navarrete- Heredia et al., 2002: 294 (subgenus of Palaminus ).

DIAGNOSIS: Palaminus is one of the most recognizable genera in the subfamily; it is distinguished from all other paederines (and other staphylinids) by the imbricate abdominal macrosculpturing (figs. 26–28). Unlike other genera of the Procirrina , the elytron of Palaminus has a long, thick, medially directed seta near the apicolateral margin with a similar laterally directed seta near the apicomedial margin (figs. 11, 96); the protibia is gradually expanded apically from the base to the apex, which is about the same diameter as the base of tarsomere 1 (fig. 41), which appears to nest into the tibial apex. Like most genera of the subtribe abdominal segment III of Palaminus lacks paratergites (fig. 26). Only Oedodactylus , Pseudoprocirrus , and some species of Oedichirus have paratergites on segment III. Tergum and sternum VII are fused basally and separated apically in Oedichirus (figs. 33, 34), entirely fused in Palaminus , but separated in the other genera. Abdominal segments III–VII of Palaminus (fig. 28) and Oedichirus have a pair of ‘‘cells’’ or ‘‘windows’’ in the intersegmental membrane at the anterior edge of the segment on both the dorsal and ventral surfaces; these structures are lacking in the other genera. The absence of a row of setae on the edge of the posterior margin of the elytra (fig. 96) will separate Palaminus from Neoprocirrus , Oedodactylus , Pseudoprocirrus , and Stylokyrtus .

DESCRIPTION: Head not pedunculate, wid- er than long (fig. 90); frontoclypeus with or without subapical, interantennal, transverse ridge; postocular lateral margins short and straight (fig. 90); basal angle strongly angulate (fig. 90) to strongly rounded (fig. 5); basal margin of head gradually curved to neck (fig. 5) or to basal angle with marginal ridge extending from basal angle to neck (figs. 10, 90), and with median portion slightly to moderately emarginate anterior to neck. Neck across nuchal constriction about two fifths to half as wide as greatest postocular width of head; nuchal groove well developed; nuchal ridge moderately to poorly developed or absent. Eye length shorter to longer than postocular length of head. Dorsal surface with umbilicate punctation (fig. 10), some species with barely discernible punctation. Ventral surface without postocular groove. Gular sutures present (fig. 91) and distinct to barely visible (fig. 12) to absent; sutures moderately widely separated and most approximate near or proximad of middle to narrowly separated and most approximate posteriorly. Gula without pubescence. Antennomere 11 shorter than 9 and 10 combined, constricted sub- apically and tapered to narrowly rounded apex, and without apical, spinelike pencil of setae. Mandibles with apically acute denticle (fig. 98), not bifid; prostheca evident as cluster of cuticular processes at base (fig. 98). Maxillary palpus (fig. 95) with palpomere 4 transverse, securiform, and greatest length longer than third palpomere, and compressed. Labium (fig. 94) with glossae separated, apically rounded and tapered and with cuticular processes on medial margin. Hypopharynx without lobes or cluster of spinelike setae on anterior margin; lateral region with dense cluster of cuticular processes. Labrum (fig. 97) with one pair of submedial denticles on anterior margin. Epipharynx with seta near anterior margin laterad of middle and with small cluster of setae near lateral base; lateral region with long, cuticular processes; median groove present; epipharynx not visible along anterior margin of labrum in dorsal view.

Prothorax (fig. 90) curved trapezoidal, wider than long to slightly longer than wide; widest near anterior margin or anterior third and with lateral margins curved and gradually convergent posteriad. Pronotum with moderately dense umbilicate punctation; punctation weak in some species; punctation absent from midline basally. Pronotal marginal ridge absent or present. Notosternal suture present distad of procoxa (fig. 18). Hypomeron polished and impunctate. Postprocoxal lobe (figs. 19, 93) short to poorly developed, impunctate, and with curved ridge near ventral edge; transverse ridge absent; apices widely separated from one another. Probasisternum (fig. 93) short, without median carina, and without punctation. Mesospiracular peritreme (fig. 19) with anteromedial margin fused to furcasternum; suture absent. Elytra (fig. 5) longer to shorter than pronotum; humeral angles present or absent; posterior edge without row of setae; subapical region with one prominent, long, thicker, posteromediad-directed seta near lateroapical corner and another posterolaterad one near medioapcial corner (figs. 11, 96). Scutellum with scattered setae. Mesosternum with short, median carina extending from mesosternal process. Mesocoxal acetabulum with marginal carina laterally and partly posteriorly. Intersternal suture present medially but feebly developed.

Profemur with ridge and slit (fig. 40) on ventral edge extending from near middle toward base. Protibia with multiple (8 and 9 in species from U.S.A. and Peru respectively) combs (fig. 41); surface without (fig. 41) or with feeble depression on ctenidial surface; tibia widest apically, gradually expanded from base to apex. Protarsomeres 1– 4 inflated (fig. 92); base of tarsomere 1 approximately same diameter as apex of expanded tibia (fig. 41); tarsomere 4 not expanded beneath tarsomere 5 and apical margin entire, not bilobed; tarsomere 5 unmodified and inserted laterad of middle of asymmetrical tarsomere 4; tarsomere 5 densely pubescent ventrally. Mesotarsomere and metatarsomere 1 as long as to longer than combination of tarsomeres 2 and 3 through 2–5.

Abdominal segments III to VI with imbricate macrosculpturing (figs. 26–28), VII without or with feeble imbricate macrosculpturing, and VIII and IX without it. Segment III without paratergites; paratergal carina present, short, and at base ventrad of spiracle (figs. 26, 27); tergum and sternum III fused. Segment VII with tergum and sternum completely fused and without notch at midlateral edge of posterior margin. Segments IV to VII with ‘‘windows’’ or ‘‘cells’’ covered with white, opaque membrane in intersegmental membrane at anterior edge of tergum and sternum (fig. 28). Sternum I absent. Sterna IV–VII without glandular lobe or slit on anterior margin. Tergum IX (figs. 29, 99) fused basally; emargination deep, three or more times deeper than length of tergum, and wide; lateroapical process (fig. 99) long, slender, straight to slightly curved ventrally, and extending well beyond tergum X; lateroapical process separated basally from remainder of segment (figs. 99, 100). Tergum X of male and female with apical margin rounded; base separated from tergum IX.

Female genital sclerites fused to form broad, distal gonocoxal plate partially fused to basally divided, proximal gonocoxal plate and vulva embedded in proximal region (fig. 101) in some species and in others to form one median gonocoxal plate.

Aedeagus (fig. 102) asymmetrical; parameres long, complex, and separated from median lobe for most of length; basal piece absent.

DISTRIBUTION AND HABITAT: The first four species were described by Erichson (1840: 682, 683) the year after he made the name Palaminus available. Three of the species, P. piceus , P. pilosus , and P. variabilis , were from Colombia; the latter was also reported from the West Indies and the fourth, P. testaceus , from the U.S.A. The most recently described species was from the Comoros ( Lecoq, 1996: 138). In contrast to the sharply skewed Old versus New World distribution of Oedichirus , Palaminus is more evenly distributed with 141 New World and 165 Old World species. Madagascar, with 55 species, has more than any other country and the next closest is Brazil with 38. The genus is mostly a tropical and subtropical group with few species in temperate and semiarid regions.

In the New World the genus reaches north to the southern edge of Ontario, Canada ( Campbell and Davies, 1991: 113), on the north shore of Lake Erie at 42.28 ° N (Anthony Davies, personal commun.) and extends south through eastern, southeastern, and southwestern United States and West Indies and Mexico to Argentina. For the United States, I have seen material for most of the eastern states to Arkansas, Missouri, Kansas, and Oklahoma, and species are recorded in Texas, New Mexico, and Arizona. The only far-western species was described from the southern Arizona / California border along the Colorado River. No species are reported in the northern Plains, central and northern Rockies, Great Basin, or western coastal states from southern California (except near Arizona) to British Colombia. In the West Indies I have seen specimens or species have been reported in the literature for the Bahamas, Greater Antilles ( Cuba, Jamaica, Dominican Republic, Puerto Rico), and Lesser Antilles ( Dominica, Trinidad). The genus is known in most countries from Mexico to Argentina where it reaches the north and central provinces of Jujuy (23 ° S), Salta (22 ° S), Tucumán, Misiones, San Luis (32 ° S), and Buenos Aires (33 ° S). The genus may be absent from most of Canada, the western half of the United States (except southern California, Arizona, New Mexico, and Texas), southern Argentina, and Chile.

Most of the 165 Old World species are described from Africa and Madagascar and nearby islands (117 spp.). In Asia species are known from India, Sri Lanka, Myanmar, Singapore, Malaysian Borneo, Indonesia, Philippines, China, and Japan. Further south Palaminus is known from New Guinea and Australia, and in the Pacific from New Caledonia, Vanuatu, Fiji, Samoa, and Guam.

In Africa no species are reported north of the Sahel. The genus is unknown in Europe, Turkey, and the Levant west through Pakistan, central Asia, Russia, and New Zealand. All of the continental Asian species are in India, Bhutan, Myanmar, and China. It is found in Singapore and insular China in Hong Kong and Taiwan; the only record for mainland China is in Zhejiang. The species reported ( Rougemont, 2001: 43) from Hong Kong ( P. parvus ) was described from Singapore. The species from Zhejiang ( P. truncatus ; Zheng, 2001: 324) was described from Coonoor in Tamil Nadu, southern India. The three other Chinese species are from Taiwan and two of those are the only representatives in Japan. The northernmost Old World locality is at almost 35 ° N (Kobe, Japan; P. japonicus ) and the most southerly is an unspecified locality in Victoria, Australia ( P. australiae ). In India seven of the 10 species are reported from the south in Tamil Nadu and one of those is also from West Bengal. Only two species are reported in the north, one in Sikkim ( P. rufulus ) and one, the more northern species, at about 30 ° N in Arunachal Pradesh ( P. morosus ). Palaminus indicus Kraatz is reported from the Indian states of Tamil Nadu, West Bengal, and Sikkim ( Biswas, 2003: 259) and may also be in Myanmar. The type locality of the species is ‘‘ India orientali’’ ( Kraatz, 1859: 153). Many of the species reported from that site by Kraatz were from what is now Myanmar (see Herman, 2001: 7, and Frisch, 2005: 77, for discussion and references).

In the New World the genus is distributed between approximately 42 ° N and 33 ° S and in the Old World about 35 ° N and perhaps to 38 ° S. At the northern and southern edges few species are known. The regions from which it appears to be unknown may be too arid, too cool, or both.

Collections have been made in lowland and tropical forests and in montane regions to elevations as high as 2500 meters ( P. kaboboensis ; Fagel, 1971: 63). Species are found on shrubs, trees, and overhanging vegetation, and may be collected by sweeping or by using a beating sheet, or in forest leaf litter, decaying litter, litter near streams, and ground debris. I have collected and/or have seen specimens of both flightless and flying specimens from leaf litter and from beating trees and bushes. It is unclear whether the winged species found on the ground live there or have fallen there from the vegetation, but I have seen many specimens collected from both habitats. Fagel (1971: 55), in his revision of the African species, thought that individuals collected from ground litter probably fell from vegetation. Bierig (1935: 32) wrote that flightless species were solitary vagabonds that lived in humus (‘‘formas ápteras terrícolas… pueblan en terrenos húmedos el humus de boscajes, son seres solitarios y vagamundos’’). Species have been collected in the upper tree canopy in Panama and probably occur there in most tropical regions.

SYNONYMY: Considering that Palaminus includes more than 300 species, there is surprisingly little external variation. This uniformity of external features is reflected in the lack of subgenera. Bierig (1943: 155) described one subgenus, Parapalaminus , for nine species ( P. barbiellinii , P. bifidus , P. bruchianus , P. coriaceus , P. diffinis , P. peralutaceus , P. pusillus , P. scitulus , and P. symphylus ) with alutaceous microsculpturing on the cephalic dorsum and pronotum. Bierig presented no other characters to support recognition of a subgenus. Another species, P. erichsoni , also has alutaceous sculpturing, but whereas the species cited by Bierig are small and yellowish brown, this species is larger and reddish brown with yellowish-brown elytral maculations. At least four species ( P. bifidus , P. grandicollis , P. parvipennis , P. scitulus ) have this microsculpturing only on the pronotum. The head (dorsal surface) and pronotum of the great majority of species of Palaminus , including its type species, are polished and lack microsculpturing. A few others have feeble cephalic and pronotal microsculpturing, so it appears that this sculpturing defines no group and Parapalaminus should be synony- mized. It is doubtful that the stated sculpturing defines a monophyletic group. Furthermore, by separating a subgenus, Parapalaminus , a paraphyletic group, with 296 species, is created because there are no unique characters common to all the remaining species of Palaminus that are not also shared by those of Parapalaminus . If description or recognition of a subgenus results in the remaining species becoming a paraphyletic group then the subgenus should not be described or, if it already exists, should be synonymized, so as to maintain a genus defined by synapomorphic features. Parapalaminus is hereby synonymized with Palaminus .

DISCUSSION: Palaminus species are small, slender, elegant beetles with polished integument. Externally the species are similar, differing in color and size, but with beautiful, imbricate macrosculpturing (triangular or diamond shaped) (figs. 26–28) on the cylindrical abdomen. Most species are honeycolored yellowish brown, but others are black or reddish brown and some have maculations. The subterminal abdominal sterna of the males may exhibit elaborate secondary sexual characteristic, but the most striking variation occurs in the aedeagus which has rich, complex structure in many species (fig. 102) and is one of the principal, most reliable means of defining species of Palaminus with use of the males (see for example the illustrations in Fagel, 1971: 52–126). Preliminary studies suggest that the female genital sclerites also offer a rich unexploited resource that may be useful for species identification of females and for phylogenetic hypotheses. As yet no infrageneric classification exists for Palaminus , so it is unclear that aedeagal characters or female genitalic sclerites will determine groups. Because of the dearth of external characters, I expect that many specimens identified without reference to the aedeagus are misidentified and that some are synonyms or complexes of cryptic species.

The number of named species for the New and Old World differs only slightly. Howev- er, during the 117-year interval between when the first four species were described ( Erichson, 1840) and 1956, when the most recent six were added ( Wendeler, 1956a, 1956b), all of the 142 species known for Palaminus in the New World were described. It was not until 1859 that the first two Old World species were described ( Kraatz, 1859: 153) and by 1952 only 60 species had been described for the entire hemisphere, so by 1956 more than two thirds of the named species were in the New World. During the 26 years from 1971– 1996, Fagel (1971), Jarrige (1974, 1978), and Lecoq (1984, 1986, 1991, 1996) described 102 more species from Africa and Madagascar. Five others were described from Bhutan ( Coiffait, 1978a: 114), Myanmar ( Scheerpeltz, 1965: 103, 172), and Indonesia ( Scheerpeltz, 1957: 260–262). Considering the external uniformity of the species and the fact that the most diagnostic features are associated with the aedeagus, the male terminal abdominal segments, and the female genital sclerites, there are surely many species awaiting discovery. By contrast, during the nearly 170 years the genus has been known, only one name has been listed as a synonym, one as a synonym and valid name in the same publication, two were published as varieties, one as a subspecies, and five names are homonyms and were replaced. Since in the New World and in the eastern part of the Old World the aedeagus was not used to distinguish species, it is probable that some synonymy will be discovered in those regions. Since in Africa and Madagascar the aedeagus and the terminal abdominal segments were employed extensively, it is less likely that heretofore unnoted synonyms will be found.

Palaminus requires revisionary study in nearly all parts of its range. The Nearctic species are being revised by Larry Watrous at this writing. Among some of the important and more or less recent taxonomic studies on Palaminus are a review of and key to the species of India ( Cameron, 1931: 20), a revision and illustrations of the species for Africa ( Fagel, 1971: 52) and description and illustrations for Madagascar ( Lecoq, 1984, 1986, 1991, 1996; Jarrige, 1974, 1978). No keys were published for Africa or Madagascar and identification relies on comparison of specimens with descriptions and aedeagal illustrations. Downie and Arnett (1996: 424) provide a key and diagnoses for the species of northeastern North America, but without aedeagal characters or illustrations. Notman (1929) published a key to the 144 species then known; the number of species has more than doubled since. All of the characters Notman uses are external (color, form, punctation), he does not indicate whether the key was based on characters from the literature or from actual specimens, and no use was made of the aedeagus or secondary sexual characteristics of the male abdomen, so the accuracy of identification using the key is limited. Many other articles present a list of species for a region or include isolated descriptions of new ones.

By beating bushes, trees, lianas, and so on, one can relatively easily collect Palaminus and large numbers of individuals can be quickly accumulated. Many of the identified Palaminus I examined were represented by few specimens, but many collections often have large accumulations of undetermined specimens. Obviously, any given species can be uncommon at a site and for that reason might be rare in museum collections, but the paucity of specimens of Palaminus may also partly reflect the lack of revisionary study that develops the tools needed to identify these masses of similar specimens.

Most species of Palaminus can fly. A few species ( P. apterus , P. larvalis , P. rosariensis , plus other unidentified species from various regions) have shorter elytra that lack humeral angles and may be incapable of flight.

A specimen of an unidentified species of Palaminus was reported in Oligocene/Miocene amber from Mexico ( Seevers, 1971: 84) and at least nine specimens of approximately the same age in Dominican amber are known in collections (Herman, personal obs.). The most diagnostic characters for species are the aedeagus and the female genital sclerites. The males of some living species have especially elaborate modifications of the abdominal sterna, but most do not, so identity often requires dissections. The extant species of the genus are far too poorly known for most parts of the world, but especially for the New World, to defend naming any of these fossils species. If and when the New World species are revised, describing and naming the fossil might be warranted.

Over a period of five years flights were made over Tallulah, Louisiana, in a biplane fitted with an aerial plankton net in an effort to gather information on dispersal and migration of insects, spiders, and mites. Flights were made during the day and night, at various elevations. During five years there were over 1300 flights. Over 4400 beetles were collected of which 19 were Palaminus spp. ; of these 19 species, 16 were collected at night at about 500 feet altitude ( Glick, 1939: 31), and which only shows that the species are more apt to fly at night when it is more humid.

SPECIES INCLUDED AND

MATERIAL EXAMINED

aequicollis Bernhauer – L ( MRAC), Pl, sp ( FMNH)

................................. D.R. Congo africanus Fagel – H ( MRAC)........ Guinea allocerus Fauvel – L ( IRSN)...... Madagascar alluaudi Jarrige – H ( IRSN)........ Mauritius aluticollis Bernhauer – H ( FMNH)...... Brazil amplipennis Fall – Lit. Att. [ U.S.A.] anacoreta Bierig – H, sp ( FMNH)...... Cuba anceps Sharp – H ( BMNH)........... Brazil andapanus Lecoq – Lit. Att. [ Madagascar] andohahelo Lecoq – Lit. Att. [ Madagascar] andrewesi Cameron – Syn ( BMNH), sp ( FMNH)

.............................. India angolensis Fagel – sp ( BMNH)

(Note: No locality with specimen; known from

Angola.) anjavidilavanus Jarrige – Lit. Att. [ Madagascar] ankazobensis Lecoq – Lit. Att. [ Madagascar]

proximus Lecoq anosyanus Lecoq – Lit. Att. [ Madagascar] apicalis Sharp – H ( BMNH).......... Brazil apicatus Bernhauer – Syn ( FMNH).... Colombia apicipennis Sharp – Syn ( BMNH), Syn, sp ( FMNH)

.......................... Guatemala, Mexico apterus Bernhauer – H ( FMNH)... Guadeloupe armatus Fagel – H ( MRAC)..... D.R. Congo assimilis Lecoq – Lit. Att. [ Madagascar] asymmetricus Bernhauer – Syn ( FMNH).....

.......................... Costa Rica atriventris Bernhauer – Syn ( FMNH)...... Fiji

lateralis Cameron – P ( BMNH)........ Fiji australiae Fauvel – H ( BMNH), sp ( FMNH, IRSN)

..................... Australia, Loyalty Islands

hebridensis Cameron – H ( BMNH)........

........................ New Hebrides bafianus Fagel – Lit. Att. [ Cameroon] balzani Bernhauer – Syn ( FMNH)..... Bolivia barbiellinii Bernhauer – H, sp ( FMNH)..... Brazil barombiensis Fagel – Lit. Att. [ Nigeria] bermudezi Bierig – H ( FMNH)........ Cuba betschi Jarrige – Lit. Att. [ Madagascar] bicolor Cameron – H ( BMNH)............

.................... Papua New Guinea bifidus Notman – H, P ( AMNH), P ( FMNH)....

................................. Puerto Rico biguttatus Fauvel – Syn ( IRSN), sp ( FMNH).....

................................... Venezuela biimpressus Bernhauer – Syn ( FMNH).......

.......................... Costa Rica bipustulatus Bernhauer – Syn ( FMNH).......

........................... Colombia birmanus Cameron – H ( BMNH)...... Burma bivittatus Cameron – Syn ( BMNH), sp ( FMNH)

......................... New Guinea

obliteratus Cameron – Syn ( BMNH).......

......................... New Guinea bivittipennis Lea – Lit. Att. [New Guinea] blanci Jarrige – Lit. Att. [ Madagascar] bolivianus Bernhauer – Syn, sp ( FMNH).....

........................ Bolivia, Brazil bonariensis Lynch Arribálzaga – Syn? ( IRSN), sp

( FMNH)................... Argentina

(Note: Lynch Arribálzaga [1884: 330] wrote ‘‘El

ejemplar que poseo lo coleccioné en Chaca-

buco,…’’ which suggests that he examined only

one specimen. Since he collected the specimen it

is probable that it is deposited with his collection

in Buenos Aires and it seems unlikely that type

material is in the IRSN.) borneenis Cameron – H ( BMNH), sp ( FMNH)...

.................................... Malaysia brachelytratus Lecoq – Lit. Att. [ Madagascar] brachypterus Lecoq – Lit. Att. [ Madagascar] breviceps Sharp – H ( BMNH), sp ( FMNH)...

.............................. Brazil brevicollis R. Sahlberg – sp ( FMNH).... Brazil brevipennis Bernhauer –H,sp( FMNH) ... Argentina bruchi Bernhauer – H ( FMNH)..... Argentina bruchianus Bierig – Syn ( FMNH)....... Cuba brunneus Fauvel – H ( IRSN)...... Madagascar bryanti Cameron – Syn ( BMNH).... Singapore buehleri Scheerpeltz – Syn ( NHMW)... Indonesia capitalis Lecoq – Lit. Att. [ Madagascar] carayoni Lecoq – Lit. Att. [ Madagascar] carinatus Lecoq – Lit. Att. [ Madagascar] ceylanensis Kraatz – Syn ( SDEI), sp ( BMNH)...

............................... Sri Lanka cheesmani Cameron – H ( BMNH)..............

.......................... Papua New Guinea chiriquensis Sharp – H ( BMNH), sp ( FMNH)....

.................................... Panama circumflexus Fauvel – L ( IRSN), sp ( FMNH)....

................................. Madagascar cognatus Sharp – Syn ( BMNH, FMNH)......

............................ Panama collaris Bernhauer – Syn ( FMNH)... Costa Rica columbinus Bernhauer – Syn ( FMNH).......

........................... Colombia compressiventris Bernhauer – Syn ( FMNH)....

.............................. Mexico concolor Sharp – Syn ( BMNH), sp ( FMNH)...

.................... Guatemala, Mexico confusus Fagel – H ( BMNH), P ( MRAC).....

................... D.R. Congo, Angola congoensis Cameron – Syn ( BMNH, MRAC)....

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

(Note: Fagel [1971: 125] considers species to be

incertae sedis for lack of a male.) consimilis Lecoq – Lit. Att. [ Madagascar] continentalis Bernhauer – Syn, sp ( FMNH)....

......................... South Africa contortus LeConte – Lit. Att. [ U.S.A.] coriaceus Cameron – Syn ( BMNH), sp ( FMNH)...

................................ Haiti, Jamaica crassus Sharp – H ( BMNH), sp ( FMNH).....

.............................. Brazil crenatus Sharp – H ( BMNH)...... Guatemala crenulatus Lecoq – Lit. Att. [ Madagascar] cribratus LeConte – Lit. Att. [ U.S.A.] debilis Sharp – H ( BMNH), sp ( FMNH).....

.................... Mexico, Nicaragua decorus Lecoq – Lit. Att. [ Madagascar] decussatus Wollaston – Syn ( BMNH)... Madeira deformis Sharp – Syn ( BMNH, FMNH)......

.................... Guatemala, Mexico depressus Lecoq – Lit. Att. [ Madagascar] densicollis Bernhauer – H ( FMNH).... Mexico descarpentriesi Jarrige – Lit. Att. [ Madagascar] dieganus Jarrige – Lit. Att. [ Madagascar] difficilis Cameron – P ( BMNH).......... Fiji diffinis Sharp – H ( BMNH), sp ( FMNH).....

................. Nicaragua, Costa Rica discretus Sharp – H ( BMNH)......... Brazil distans Sharp – H ( BMNH)........... Brazil dubius Notman – H ( AMNH)..... Guadeloupe elegans Wendeler – H, P ( MNHB)...... Brazil erichsoni Bernhauer – H ( FMNH)..... Bolivia evansi Bernhauer – Syn ( FMNH)......... Fiji exiguus Lecoq – Lit. Att. [ Madagascar] fageli Jarrige – H ( IRSN)........ Madagascar falsus Fagel – H ( MRAC)....... D.R. Congo fauveli Jarrige – H ( IRSN)....... Madagascar ferrugineus R. Sahlberg – sp ( FMNH)... Brazil fijiensis Cameron – P ( BMNH).......... Fiji filicornis Fagel – H ( MRAC)..... D.R. Congo filum Sharp – Syn ( BMNH), Syn, sp ( FMNH).

........... Costa Rica, Mexico, Nicaragua fiziensis Fagel – H ( MRAC)..... D.R. Congo flavescens Fagel – Lit. Att. [ Guinea] flavipennis LeConte – Lit. Att. [ U.S.A.] flavocinctus Lecoq – Lit. Att. [ Madagascar] flavoguttatus Cameron – Syn ( BMNH).......

.......................... Philippines flavus Fagel – H ( MRAC)....... D.R. Congo formosae Cameron – P ( BMNH)...... Taiwan foutadjallonensis Fagel – P ( MRAC).... Guinea fragilis Sharp – H ( BMNH)........... Brazil fraternus Casey – Lit. Att. [ U.S.A.] freyianus Fagel – Lit. Att. [ Guinea] fuscatus Lecoq – Lit. Att. [ Madagascar] fuscipes Sharp – H ( BMNH), sp ( FMNH)....

.............................. Brazil fusciventris Sharp – H ( BMNH), sp ( FMNH)..

................. Nicaragua, Costa Rica fuscus Lecoq – Lit. Att. [ Madagascar] gabonicus Fagel – Lit. Att. [ Gabon] garambanus Fagel – H ( MRAC)..... D.R. Congo germanus Cameron – P ( BMNH)..... Malaysia gracilipes Sharp – Syn ( BMNH), Syn, sp ( FMNH)

............. Panama, Costa Rica, Mexico gracilis Sharp – H ( BMNH), sp ( FMNH).....

................. Brazil, Colombia, Peru grandicollis Notman – H, P ( AMNH), P ( FMNH)

......................... Puerto Rico grandis Lecoq – Lit. Att. [ Madagascar] griveaudi Jarrige – Lit. Att. [ Madagascar] guineensis Fagel – Lit. Att. [ Guinea, Ivory Coast] heraldicus Fauvel – Syn ( IRSN), sp ( FMNH).....

......................... Venezuela, Colombia hudsonicus Casey – Lit. Att. [ U.S.A.] hylaeus Bierig – Syn, sp ( FMNH)...... Cuba illotus Lecoq – sp ( BMNH)....... Madagascar implicatus Lecoq – Lit. Att. [ Madagascar] incertus Fagel – H ( MRAC)..... D.R. Congo inconspicuus Lecoq – Lit. Att. [ Madagascar] indicus Kraatz – Syn ( SDEI), sp ( FMNH)....

........................ India, ‘‘ India

orientali, Indonesia, Philippines, Singapore, Vietnam infuscatus Bernhauer – Syn, sp ( FMNH).....

............................ Mexico insignis Sharp – H ( BMNH), sp ( FMNH).....

................... Panama, Costa Rica insularis Cameron – Syn ( BMNH), sp ( FMNH)

.................. Jamaica, Guadeloupe invidus Casey – sp ( FMNH)......... Mexico isthmianus Casey – Lit. Att. [ Panama] japonicus Cameron – H ( BMNH)....... Japan jarrigei Lecoq – Lit. Att. [ Madagascar] jocquei Lecoq – Lit. Att. [ Comoros] kaboboensis Fagel – H ( MRAC)..... D.R. Congo

ruwenzoricus Fagel – H ( MRAC)... D.R. Congo kamerunensis Bernhauer –Pl( FMNH) ... Cameroon kasaiensis Fagel – H ( MRAC).... D.R. Congo katanganus Fagel – H ( MRAC)... D.R. Congo kivuensis Fagel – H ( MRAC)..... D.R. Congo kokodanus Cameron – H ( BMNH)..........

.................... Papua New Guinea kundelungensis Fagel – H ( MRAC)... D.R. Congo kwangensis Fagel – H ( MRAC)... D.R. Congo lacertinus Sharp – Syn ( BMNH)... Guatemala lancifer Fauvel – Syn ( IRSN)....... Venezuela larvalis LeConte – sp ( FMNH)............

...... United States (North Carolina, Texas). lateripennis Bernhauer – Syn ( FMNH).......

.......................... Costa Rica laticollis Wendeler – H ( MNHB)....... Brazil latior Bernhauer – Syn ( FMNH)... Costa Rica lengi Notman – H ( AMNH)...... Puerto Rico liberiensis Fagel – Lit. Att. [ Liberia] ligulifer Lecoq – Lit. Att. [ Madagascar] limbifer Fauvel – H ( BMNH), sp ( IRSN).....

......................... New Guinea

(Note: The collection of the IRSN has a

specimen with a ‘‘type’’ label, but according to

the original description [ Fauvel, 1879: 82] the

species was described from one specimen col-

lected by Wallace and that specimen is in the

BMNH.) lividus LeConte – Lit. Att. [ U.S.A.] lobiventris Bernhauer – Syn ( FMNH)........

.......................... Costa Rica longelytratus Lecoq – Lit. Att. [ Madagascar] longicollis Sharp – H ( BMNH)...... Panama longicornis Sharp – H ( BMNH), sp ( FMNH)..

.............................. Brazil luteus Casey – Lit. Att. [ Canada, U.S.A.] luberensis Fagel – H ( MRAC).... D.R. Congo lumiventris Herman – Lit. Att. [ Taiwan]

spiniventris Bernhauer

(Note: Holotype missing from FMNH collec-

tion.) maculatus Bernhauer – Syn ( FMNH)... Australia madecassa Fauvel – L ( IRSN), sp ( FMNH)...

......................... Madagascar magniceps Bernhauer – H, sp ( FMNH)......

........................... Argentina magnipennis Bernhauer – Syn ( FMNH)......

.......................... Costa Rica malaisei Scheerpeltz – Lit. Att. [Type not found in

Vienna – May 2006] malandanus Bernhauer – Syn ( BMNH, FMNH)

........................... Australia masoalaensis Lecoq – Lit. Att. [ Madagascar]

difficilis Lecoq mexicanus Sharp – Syn ( BMNH), sp ( FMNH).

............................ Mexico milloti Lecoq – Lit. Att. [ Comoros] minor Bernhauer – Syn ( FMNH)...... Mexico minutissimus Bernhauer – Syn ( FMNH)... Guam minusculus Lecoq – Lit. Att. [ Madagascar] minutulus Lecoq – Lit. Att. [ Madagascar] minutus Fagel – Lit. Att. [D.R. Congo] mohelianus Lecoq – Lit. Att. [ Comoros] motoensis Fagel – H ( MRAC).... D.R. Congo modestus Sharp – H ( BMNH)......... Brazil molestus Jarrige – Lit. Att. [ Madagascar] montanus Cameron – H ( BMNH)... Mauritius monticola Cameron – Syn ( BMNH), sp ( FMNH)

........................... Sri Lanka morosus Cameron – Syn ( BMNH)...... India multifidus Lecoq – Lit. Att. [ Madagascar] nevermanni Bernhauer – Syn ( FMNH).......

.......................... Costa Rica niger Sharp – Syn ( BMNH), sp ( FMNH).....

........................ Bolivia, Brazil nigropiceus Cameron – Syn ( BMNH), sp ( FMNH)

............................... India nigrosuturalis Bernhauer – Syn ( FMNH).....

........................... Argentina nilgiriensis Cameron – Syn ( BMNH).... India nimbaensis Fagel – Lit. Att. [ Guinea] nitidulus Fagel – H ( MRAC)..... D.R. Congo normalis LeConte – Lit. Att. [ U.S.A.] nossibianus Fauvel – L ( IRSN), sp ( FMNH)...

.................. Madagascar, Tanzania novaeguineae Fauvel – sp ( FMNH).... Indonesia occidentalis Lecoq – Lit. Att. [ Madagascar] oculatus Fagel – H ( MRAC)..... D.R. Congo ogloblini Bernhauer – H ( FMNH)... Argentina omissus Lecoq – Lit. Att. [ Madagascar] opaculus Bernhauer – H ( FMNH)..... Bolivia ophtalmicus Lecoq – sp ( BMNH)... Madagascar pallidus R. Sahlberg – sp ( FMNH)...... Brazil pallipes LeConte – sp ( FMNH)............

................. United States (Florida) palmatus Sharp – H ( BMNH)....... Panama papuanus Cameron – H ( BMNH)...........

.................... Papua New Guinea parcus Sharp – H ( BMNH)........... Brazil pauliani Lecoq – Lit. Att. [ Madagascar] parviceps Sharp – H ( BMNH)........ Mexico parvipennis Notman – H, P ( AMNH), P ( FMNH)

......................... Puerto Rico parvulus Sharp – Syn ( BMNH), sp ( FMNH)...

............................ Mexico parvus Cameron – Syn ( BMNH).... Singapore pauloensis Bernhauer – H, sp ( FMNH)... Brazil pellax Sharp – H ( BMNH)........... Brazil pennifer Fauvel – L ( IRSN), sp ( FMNH).....

..... India, Indonesia, Philippines, Seychelles peralutaceus Bierig – Syn ( FMNH)...... Cuba perforatus Wendeler – H ( MNHB)...... Brazil pertusus Lecoq – Lit. Att. [ Madagascar] peyrierasi Lecoq – Lit. Att. [ Madagascar] philippinus Bernhauer – Syn, sp ( FMNH).....

.................. Philippines, Singapore piceus Erichson – H ( MNHB)...... Colombia pictus Bernhauer – Syn ( FMNH, IRSN)......

............................. Bolivia pilosus Erichson – Syn ( MNHB).... Colombia pilum Bierig – H, P ( FMNH).......... Cuba plagiatus Fauvel – Syn ( IRSN)..... Venezuela problematicus Fagel – H ( MRAC)... D.R. Congo procerus Notman – H ( AMNH).... Puerto Rico productus Schubert – Syn ( MNHB)... Tanzania prolongatus Jarrige – P ( IRSN).... Madagascar propinquus Lecoq – Lit. Att. [ Madagascar] proximus Cameron – H, P ( BMNH).........

........................ New Hebrides pubescens Lecoq – Lit. Att. [ Madagascar] pullus Lecoq – Lit. Att. [ Madagascar] pulvereus Lecoq – Lit. Att. [ Madagascar] pumilus LeConte – Lit. Att. [ U.S.A.] puncticeps Sharp – H ( BMNH), sp ( FMNH)...

.............................. Brazil puncticollis Fagel – H ( MRAC)... D.R. Congo pusillus Notman – H ( AMNH).... Puerto Rico quadriguttatus Fauvel – Syn ( IRSN).........

........................... Venezuela quadripustulatus Bernhauer – H, sp ( FMNH)...

...................... Bolivia, Colombia rejectus Lecoq – Lit. Att. [ Madagascar] robustus Sharp – H ( BMNH).......... Brazil rosariensis Bierig – Syn ( FMNH)....... Cuba rotundicollis Wendeler – H ( MNHB).... Brazil rubripennis Sharp – Syn ( BMNH, FMNH)....

............................ Panama rufotestaceus Lecoq – Lit. Att. [ Madagascar] rufulus Coiffait – Lit. Att. [ Nepal, India, Bhutan] rufus Cameron – Syn ( BMNH), sp ( FMNH)...

.............................. India rugicollis Fauvel – Syn ( IRSN)..... Venezuela sambavanus Lecoq – Lit. Att. [ Madagascar] samoensis Cameron – P ( BMNH)..... Samoa saopaoloanus Wendeler – H ( MNHB).... Brazil schirmi Wendeler – H ( MNHB)........ Brazil scitulus Notman – H, P ( AMNH).. Puerto Rico sellatus Sharp – Syn ( BMNH)......... Brazil sharpi Bernhauer – H ( FMNH)......... Peru signifer Casey – Lit. Att. [ U.S.A.] silvestris Jarrige – Lit. Att. [ Madagascar] similis Fagel – H ( MRAC)...... D.R. Congo simoni Fauvel – Syn ( IRSN, BMNH)........

...................... Sri Lanka, India simplex Sharp – Syn ( BMNH), sp ( FMNH)...

.............................. Brazil sinuatus Sharp – H ( BMNH).......... Brazil sobrinus Sharp – Syn ( BMNH)........ Brazil sogai Jarrige – Lit. Att. [ Madagascar] solitus Lecoq – Lit. Att. [ Madagascar] sordidus Sharp – Syn ( BMNH)....... Mexico spiniventris Bernhauer – Syn ( FMNH)... Brazil stipes Sharp – H ( BMNH)............ Brazil strigosus Bierig – Syn ( FMNH).... Costa Rica sumbaensis Scheerpeltz – Syn ( NHMW)......

........................... Indonesia sutteri Scheerpeltz – Syn ( NHMW).. Indonesia suturalis Fauvel – L ( IRSN)...... Madagascar swezeyi Bernhauer – Syn ( FMNH)...... Guam symphylus Bierig – H, sp ( FMNH)... Costa Rica tanalensis Jarrige – H ( IRSN)..... Madagascar tegulatus Casey – Lit. Att. [ Nicaragua] tener Bernhauer – H ( FMNH)......... Brazil tenuis Lecoq – Lit. Att. [ Madagascar] testaceus Erichson – sp ( FMNH)...........

......... U.S.A. (Florida, Georgia, Illinois,

Kansas, New Jersey, North Carolina, Tennessee) thiemei Bernhauer – Syn, sp ( FMNH).......

........................... Colombia timidus Casey – Lit. Att. [ U.S.A.] tortilis Casey – Lit. Att. [ Nicaragua] transmarinus Fauvel – Syn ( IRSN), sp ( FMNH)

........................... Tanzania trapezicollis Fauvel – H ( IRSN)..... Colombia

(Note: No type label is attached to this

specimen, but Fauvel [1901: 76] wrote that the

species was based on one specimen.) trapezimorphus Herman

trapezicollis Bernhauer – H ( FMNH)... Brazil trifidus Lecoq – Lit. Att. [ Madagascar] trinotatus Fauvel – Syn ( IRSN)..... Venezuela

(Note: The species was described in a key and no

locality was cited. In the IRSN collection the

type is labeled from ‘‘Colonia Tovar’’.) trivittipennis Lea – Lit. Att. [ Fiji] truncatus Fauvel – Syn ( IRSN), sp ( FMNH)...

.............................. India tsaratananus Lecoq – Lit. Att. [ Madagascar] tucumanensis Bernhauer – Syn ( FMNH)......

........................... Argentina uncinatus Lecoq – Lit. Att. [ Madagascar] unicus Lecoq – Lit. Att. [ Madagascar] uniformis Cameron – H ( BMNH)..........

.................... Papua New Guinea validus Sharp – Syn ( BMNH)....... Panama variabilis Erichson – Syn ( MNHB), sp ( FMNH)

.............. Brazil, Colombia, Grenada variiventris Bernhauer – Syn ( FMNH).. Bolivia varipennis Cameron – Syn ( BMNH).........

......................... New Guinea veronicae Lecoq – Lit. Att. [ Madagascar]

maculatus Lecoq villiersi Lecoq – Lit. Att. [ Madagascar] vitiensis Fauvel – Syn ( IRSN)........... Fiji vittatus Sharp – Syn ( BMNH), sp ( FMNH)...

.................... Costa Rica, Mexico

humeralis Sharp – H ( BMNH)...... Mexico vittiger Bernhauer – Syn ( FMNH)..........

........................ Bolivia, Peru volans Lecoq – Lit. Att. [ Madagascar] volitans Lecoq – Lit. Att. [ Madagascar] yangambiensis Fagel – H ( MRAC), P ( FMNH).

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

UNDETERMINED SPECIMENS: Africa: Liberia, Rhodesia.

Asia: Borneo, India, Indonesia, Japan, Malaysia, Taiwan.

Australian and Pacific Region: Australia, Caroline Islands, Marianna Islands, New Caledonia, Papua New Guinea, Philippines.

South America: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Paraguay, Peru, Surinam, Venezuela.

North America: Bahamas, Belize, Costa Rica, Cuba, Dominica, Dominican Republic,

El Salvador, Guatemala, Honduras, Jamaica, Mexico, Nicaragua, Panama, Puerto Rico, Trinidad, United States (Alabama, Arizona, Arkansas, Connecticut, Florida, Georgia, Illinois, Indiana, Kansas, Kentucky, Louisiana, Maryland, Michigan, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Virginia).

DISSECTIONS: Disarticulation: Palaminus sp. (1 male, 1 female, Peru) ; (1 male, 1 female, U.S.A.) .