Procirrus Latreille

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 : 28-34

Procirrus Latreille Figures 7, 52–64

Procirrus Latreille, 1829: 436 . Type species: Procirrus lefebvrei Latreille, 1829: 436 , fixed by monotypy.

— Laporte, 1835: 123 (characters). — Erichson, 1840: 685 (characters). — Laporte, 1840: 184 (characters). — Schaum, 1852: 28 (list of species; Europe). — Lacordaire, 1854: 108 (characters; notes; list of species). — Kraatz, 1857: 667, 668 (notes; key). — Jacquelin du Val, 1857: 50 (characters). — Redtenbacher, 1857: 217 (characters). — Jacquelin du Val, 1859: 74 (catalog; Europe). — Schaum, 1859: 29 (catalog; European species). — Gemminger and Harold, 1868: 630 (catalog). — Redtenbacher, 1874: 236 (characters). — Fauvel, 1875a: 219 (catalog). — Fauvel, 1875b: xvii (catalog). — Duvivier, 1883: 176 (catalog). — Heyden et al., 1891: 110 (list of species of Europe and Caucasus Region). — Heyden et al., 1906: 152 (list of species of Europe and Caucasus Region). — Bernhauer and Schubert, 1912: 197 (catalog). — Lea, 1923: 8 (characters). — Cameron, 1925: 33 (catalog; species of British India). — Porta, 1926: 67 (characters). — Winkler, 1925: 358 (catalog; Palaearctic Region). — Cameron, 1931: 1, 18 (characters). — Scheerpeltz, 1933: 1212 (catalog). — Blackwelder, 1952: 322 (type species). — Adachi, 1955: 13, 14 (characters; key). — Kocher, 1958: 117 (checklist of species; Morocco). — Fagel, 1971: 11, 22 (characters; key to African species; type species). — Tikhomirova, 1973: 175 (checklist of species of USSR). — Bordoni, 1975: 419 (characters). — Shibata, 1977: 19 (catalog; Japanese species). — Coiffait, 1978b: 323 (characters; key to western Palaearctic species; distribution). — Hammond, 1984: 204 (checklist; Borneo). — Outerelo and Gamarra, 1985: 21 (characters). — Ciceroni and Zanetti, 1995: 19 (list of species of Italy). — Smetana, 2004: 624 (Palaearctic catalog).

Procirrus Gray, 1832: 294 (species included: lefebvrei , cited as lefeburi ). Type species: Procirrus lefebvrei Gray, 1832: 294 , fixed by monotypy. Objective synonym of Procirrus .

— Blackwelder, 1952: 322 (synonym of Procirrus ). — Smetana, 2004: 624 (synonym of Procirrus ).

Microphius Chevrolat, 1846: 201 . Type species: Procirrus lefebvrei Latreille , fixed by monotypy. Objective synonym of Procirrus .

— Blackwelder, 1952: 322 (type species; synonym of Procirrus ).

Procirrinus Koch, 1934: 79. Type species: Procirrus saulcyi Fauvel, 1873: 291 , fixed by monotypy. New synonym.

— Blackwelder, 1952: 322 (type species). — Smetana, 2004: 624 (subgenus of Procirrus ).

DIAGNOSIS: Procirrus is separated from other Procirrina by the four inflated protarsomeres (fig. 59), pedunculate base of the head (figs. 7, 52), quadridentate labrum (fig. 55), elongate pronotum (figs. 7, 52), and absence of a pronotal marginal ridge. Abdominal segment III has a paratergal carina laterally (as in fig. 23), the tergum and sternum are fused, and the segment is cylindrical. Tergum and sternum VII are separated. The quadridentate labrum will separate Procirrus from Paraprocirrus . The absence of a ventral cephalic groove (cf. fig. 2) that extends diagonally from the margin of the eye to the neck distinguishes Procirrus from Neoprocirrus . The absence of setae on the edge of the posterior margin of the elytra will separate Procirrus from Neoprocirrus , Oedodactylus , Pseudoprocirrus , and Stylokyrtus .

DESCRIPTION: Head (figs. 7, 52) pedunculate, elongate, longer than wide, tapered from posterior margin of eyes to neck; frontoclypeus with subapical, interantennal, transverse ridge; postocular lateral (fig. 52) margin strongly rounded to neck; postocular margin long; basal angle absent (fig. 52); basal margin of head indistinguishable from lateral margin, without marginal ridge, and feebly distinct from occiput. Neck across nuchal constriction one fourth to three tenths as wide as greatest postocular width of head; nuchal groove feeble, indistinct; nuchal ridge absent. Eye length less than postocular length of head (fig. 52). Dorsal surface with dense umbilicate punctation. Ventral surface without postocular groove (fig. 53). Gular sutures (fig. 53) separated anteriorly and confluent posteriorly, some species with sutures separated nearly to neck; sutures most approximate posteriorly. Gula with minute to moderately long pubescence. Antennomere 11 slightly longer than to slightly shorter than 9 and 10 combined; apex without apical spinelike pencil of setae. Mandibles with apically bifid denticle (fig. 56); prostheca evident as cluster of cuticular processes at base (fig. 56). Maxillary palpus (fig. 57) with palpomere 4 shorter than second and subequal to third, symmetrically to asymmetrically fusiform, moderately compressed. Labium (fig. 58) with glossae large, widely separated, and with narrowly rounded apex. Hypopharynx without lobes or cluster of spinelike setae on anterior margin; lateral region with dense cluster of cuticular processes. Labrum (fig. 55) with two pairs of denticles on anterior margin. Epipharynx with two small setae near anterior margin laterad of middle; surface with dense cluster of cuticular processes on curved ridge adjacent to median groove; median groove present; epipharynx not visible along anterior labral margin in dorsal view.

Prothorax (fig. 52) trapezoidal, longer than wide; widest near anterior fourth and lateral margins broadly rounded to slightly sinuate and gradually convergent posteriorly. Pronotum with dense umbilicate punctation; punctation uniform and with or without impunctate, narrow, midlongitudinal ridge on posterior third or half. Pronotal marginal ridge poorly developed or absent. Notosternal suture poorly developed. Hypomeron densely punctate. Postprocoxal lobe long and punctate; transverse ridge present proximad of apex; apex of each lobe nearly touching one another medially (fig. 54). Probasisternum without median carina (fig. 54); surface with coarse, dense punctation. Mesospiracular peritreme (fig. 54) with anteromedial margin fused to furcasternum. Elytra slightly longer to shorter than pronotum; humeral angles present (fig. 7) or absent; posterior edge without row of setae; subapical region (cf. fig. 96) without long, prominent, medioposteriorly directed seta near lateroapical corner. Scutellum pubes- cent. Mesosternum without median carina. Mesocoxal acetabulum with marginal carina laterally and posteriorly. Intersternal suture feebly developed.

Profemur with carina on distal half of anteroventral surface. Protibia with many (12 in a specimen of P. lefebvrei ) combs extending from near base to apex; tibia with feeble, indistinct depression on ctenidial surface; apical portion neither constricted nor enlarged. Protarsomeres (fig. 59) 1–4 inflated; base of tarsomere 1 not surrounded by cupulate protibial apex; apical margin of tarsomere 4 not expanded beneath tarsomere 5 and not bilobed; tarsomere 5 unmodified and inserted laterad of middle of asymmetrical tarsomere 4; tarsomere 5 with moderately dense pubescence ventrally. Mesotarsomere 1 about as long as or longer than remaining tarsomeres. Metatarsomere 1 longer than remaining tarsomeres combined.

Abdominal segments without imbricate macrosculpturing (cf. fig. 26). Segment III without paratergites; paratergal carina present at base laterad of spiracle; tergum and sternum III fused. Segment VII with tergum and sternum separated. Segments IV to VII without oval ‘‘windows’’ in intersegmental membrane (cf. fig. 24). Sternum I absent. Sterna IV to VII without glandular lobe or slit along anterior margin. Tergum IX (fig. 61) of male and female fused medially; emargination shallow, about one seventh of length of tergum, and narrow; lateroapical process (fig. 61) short, slender, slightly curved ventrally, and extending slightly beyond apical margin of tergum X; lateroapical process fused basally to remainder of segment. Tergum X (fig. 60) with apical margin rounded; base separated from tergum IX.

Genital sclerites fused to form broad, long median gonocoxal plate (fig. 64).

Aedeagus (figs. 62, 63) asymmetrical; parameres present, long, slender, with one or more apical setae, and separated from median lobe; basal piece present and divided into two sclerites at base of median lobe.

DISTRIBUTION AND HABITAT: Procirrus is a modest size group of 29 species found on the Canary Islands, across southernmost Europe and in Africa, eastward through southern Asia to Australia and Japan. Four fifths of the species are recorded from Africa (18) and Australia (6). In Africa 14 sub- Saharan species were described by Fagel (1971: 22–47) and in Australia four species are known from New South Wales, Victoria, and South Australia ( Lea, 1923: 8–10; Fauvel, 1878a: 509) and two from northwestern Western Australia ( Lea, 1923: 9–10). The remaining five species occur across southern Asia from Lebanon to Japan as follows. Procirrus hermani Drugmand occurs in Israel ( Drugmand, 1989: 108); P. saulcyi Fauvel in Israel ( Fauvel, 1873: 291), Lebanon ( Smetana, 2004: 624), and Turkey ( Assing, 2004: 683); P. feae Fauvel in India ( Cameron, 1931: 19), Myanmar ( Fauvel, 1895: 215), and Java ( Cameron, 1936: 42); P. fusculus Sharp in Bangladesh ( Sharp, 1889: 324); and P. lewisii Sharp in Thailand ( Last, 1961: 305), Hong Kong ( Rougemont, 2001: 43), and Japan ( Sharp, 1889: 324). Procirrus fusculus , originally described from Bangladesh and later reported in Japan ( Adachi, 1955: 14), was omitted from the most recent checklist of Japanese species of the genus ( Shibata, 1977: 19). Although Japan and Hong Kong share the species, none have been collected in mainland China. Note that in southwestern Asia the genus is reported only from sites near the Mediterranean; from there it next appears in India.

One of 18 African species, P. lefebvrei , is the only one of the genus in Europe and has been found only in southern Spain and southern Italy. Described originally from Sicily, it has since been recorded from Sardinia ( Porta, 1926: 67; Koch, 1934: 78), Calabria, Italy ( Porta, 1926: 67), Cádiz, Spain ( Outerelo, 1984: 291), the Canary Islands ( Fauvel, 1897: 270; Koch, 1934: 78; Machado and Oromi, 2000: 46), Algeria (P. Lucas, 1846: 122; Fauvel, 1873: 291; Koch, 1934: 78; Jarrige, 1952: 118), Morocco ( Fauvel, 1886: 32; Jarrige, 1952: 118), Tunisia ( Fauvel, 1902: 80; Normand, 1935: 365), Egypt ( Motschulsky, 1851: 656 [as P. niloticus ]; Koch, 1934: 77 [as P. lefebvrei macrops ]), Ethiopia ( Fauvel, 1876: 65), Sudan ( Scheerpeltz, 1974: 10 [as P. lefebvrei macrops ]), Java ( Fauvel, 1886: 32), and Borneo ( Scheerpeltz, 1933: 1212). The Ethiopian record is P. abyssinicus , which Fagel (1971: 30) described from a female deposited in the Fauvel collection and which Fagel stated that Fauvel (1876: 65) had confused with P. lefebvrei .

Little has been published concerning the habitat of species of Procirrus . Cameron (1931: 19) collected one specimen of an unnamed species in India in damp leaves, Fagel (1971: 27–47) cited African species collected in forest humus, leaf litter in the forest and near streams, plant detritus and at lights, Rougemont (2001: 43) reported P. lewisi from Hong Kong in damp leaf litter near a stream and in forest floor litter, and Assing (2004: 683) collected specimens of P. saulcyi in Turkey from oak and laurel litter. Species have been collected at elevations from 950 m ( P. strictus ) to as high as 1950 m ( P. bacillus ) in Africa, but most species were reported without elevational data ( Fagel, 1971: 45, 46); P. saulcyi was recorded at 400 and 920 m elevation in Turkey and 700 m in Israel ( Assing, 2004: 683).

SYNONYMY: Koch (1934: 79) described Procirrinus as a new subgenus for Procirrus saulcyi . To distinguish the monotypic subgenus from other species of the genus he relied on three elytral characters. The humeral angles are absent, the elytra are shorter than the pronotum, and they are narrower than the width of the apical region of the abdomen. Such reductions of the elytra are common to most staphylinid species that lack or have reduced wings. Species without wings tend to have more extreme reductions than those with merely shortened wings. Species are known in many genera with loss-of-flight reductions of the elytra and pterothorax and, although some authors have recognized genus-group taxa based solely on such adaptations, these features alone usually do not, perhaps never, define a monophyletic group. As there is only one species in Procirrus (Procirrinus) , the monophyly of the subgenus is a moot point. Because the stated features are unlikely to define a monophyletic group and I can find no others that do, the subgeneric name Procirrinus is placed as a junior synonym of Procirrus .

Procirrus Gray (1832: 294) is a junior synonymic homonym. His description of the genus and the only included species, P. lefebvrei (cited as P. lefeburi ), is an exact translation of the description by Latreille (1829: 436) and is partly the rationale for listing Gray’s name as a junior synonymic homonym of Latreille’s name ( Blackwelder, 1952: 322; Smetana, 2004: 624). The senior species-group name was originally spelled as P. lefeburi . Laporte (1840: 184, pl. 13, fig. 1) cited the name as P. lefeburi in the text, but as P. lefebvrei for the illustration. Thereafter, nearly all authors cited the name as Procirrus lefebvrei and that name was declared correct ( Herman, 2003: 3). Since Laporte and others used the name ‘‘ lefebvrei ’’ for the species, it is probable that the species was named for Monsieur A.L. Lefebvre, a well-known French entomologist of that era. Latreille cited the collector’s name as ‘‘Lefèvre.’’

Microphius Chevrolat (1846: 201) was published as a junior synonym of Procirrus , but was cited as ‘‘ Microphius , Dejean’’ with reference to the third edition of Dejean’s catalog (1836: 74), in which the generic name is listed with two unavailable species-group names. The generic name was cited by other authors thereafter, but without characters or available species (see Blackwelder, 1952: 246, for list of other citations of name). Finally, Chevrolat (1846: 201) listed it as a junior synonym of Procirrus , with the type species Procirrus lefebvrei Latreille , by objective synonymy ( Blackwelder, 1952: 322).

DISCUSSION: Most of the species have been described from one or a few individuals from one or a few localities. Specimens of the genus appear to be rarely collected. New records of some known species were usually published without access to the relevant type specimens. It is probable that revision of published material will reveal some misidentifications and the unpublished specimens and new collections will result in the discovery of additional African, Asian, and Australian species.

Fagel (1971: 27) presented a species group classification of five groups for 15 tropical African species, plus P. lefebvrei , based on the number and condition of the setae on the parameres. The classification has not been applied to non-African species.

Procirrus lefebvrei is currently comprised of two subspecies, the nominate from Italy, Spain, Morocco, Algeria, and Tunisia ( Fauvel, 1902: 80; Koch, 1934: 77–79; Outerelo, 1984: 291) and P. lefebveri macrops Koch , from the Canary Islands, Morocco, Algeria, Egypt, and Sudan ( Koch, 1934: 79; Jarrige, 1952: 118; Scheerpeltz, 1974: 10; Outerelo, 1984: 291–292). The species was recorded improbably from Java ( Fauvel, 1886: 32) and Borneo ( Scheerpeltz, 1933: 1212), but is almost certainly one or more other species. Most of the records for the species require confirmation and the species needs revision, both to evaluate the validity and distribution of the two subspecies, which are based largely on the size of the eyes, and to determine which species are represented in Java and Borneo. Fagel (1971) reported neither subspecies in Sudan. Only Jarrige (1952) report- ed the P. l. macrops from Morocco and Algeria; most authors who cited it reported it from Egypt and the Canaries ( Koch, 1934: 77–79; Coiffait, 1978b: 326; Smetana, 2004: 624). Outerelo (1984), who also listed it from Morocco and Algeria, was only summarizing the published records for the subspecies. If these two forms are subspecies and if Jarrige is erroneous in reporting it from Morocco and Algeria, then the curious disjunct distribution needs explanation. But if those records are accurate, then more details need be elicited about the distribution of both subspecies across North Africa.

Keys to species were published for the Mediterranean Region, including North Africa, by Coiffait (1978b: 325) and modified to include another species by Drugmand (1989). African species, except those of North Africa, can be identified with the key published by Fagel (1971: 25). The two species from India and Bangladesh were compared by Cameron (1931: 19). Fauvel (1878a: 509) provided a key for two of the Australian species and Lea (1923) described four more with comparisons among them, but published no key. Japan has only one species, which is also reported in China and Thailand.

Procirrus bicolor has Paederus -like coloration; the head, elytra, and apical abdominal segment are black, the pronotum and basal four abdominal segments are red. The color pattern of Procirrus allardianus may be similar ( Fagel, 1971: 42).

SPECIES INCLUDED AND

MATERIAL EXAMINED

abyssinicus Fagel – Lit. Att. [ Ethiopia]

allardianus Fagel – Lit. Att. [D.R. Congo]

antiquus Lea – Lit. Att. [ Australia] aristidis Fauvel – sp (FMNH)......... Egypt

hybridus Koch bacillus Fagel – H (MRAC)......... Tanzania bicolor Fagel – H (MRAC)...... D.R. Congo castelnaui Fauvel – Syn (IRSN, BMNH).....

........................... Australia congoensis Fagel – H (MRAC)... D.R. Congo crocodilus Bernhauer – Syn (FMNH)........

.......................... Zimbabwe dolichoderes Lea – sp (BMNH)...... Australia feae Fauvel – Syn (IRSN), sp (BMNH)......

...................... Myanmar, India ferrugineus Lea – Lit. Att. [ Australia] filiformis Fagel – Lit. Att. [ Kenya] fusculus Sharp – Syn (BMNH)......... India garambanus Fagel – H (MRAC)... D.R. Congo hermani Drugmand – Lit. Att. [ Israel] iti Drugmand – Lit. Att. [ Gabon] keanus Fagel – Lit. Att. [D.R. Congo] kwangensis Fagel – H (MRAC)... D.R. Congo latipennis Fagel – H (MRAC)....... Rwanda lefebvrei Latreille – sp (FMNH, SDEI).......

...... Algeria, Corsica, Egypt, Italy, Tunisia

lefebvrei Gray

niloticus Motschulsky

macrops Koch lewisii Sharp – H (BMNH), sp (FMNH).....

.................. Japan, China, Taiwan nimbaensis Fagel – sp (MRAC).... Ivory Coast opacus Lea – Lit. Att. [ Australia] saulcyi Fauvel – Syn (IRSN), sp (USNM, SDEI)

................................ Israel senegalensis Fagel – Lit. Att. [ Senegal] strictus Fagel – H (MRAC)...... D.R. Congo uniformis Fagel – H (MRAC).... D.R. Congo victoriae Fauvel – H (BMNH)....... Australia

UNDETERMINED SPECIMENS: Australia, Malaysia, Nepal, Tanzania, Zaire, Zimbabwe.

DISSECTIONS: Complete dissection: Procirrus lefebvrei (2 females, Tunisia) ; Abdominal dissection: (1 male, Egypt) .