Reclassification of the North Temperate Taxa Associated with Staphylinus Sensu Lato, Including Comments on Relevant Subtribes Of
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AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_1 Allen Press x DTPro System File # 01cc PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3287, 88 pp., 159 ®gures February 2, 2000 Reclassi®cation of the North Temperate Taxa Associated with Staphylinus Sensu Lato, Including Comments on Relevant Subtribes of Staphylinini (Coleoptera: Staphylinidae) ALESÏ SMETANA1, 2 AND ANTHONY DAVIES2 ABSTRACT This paper presents a reclassi®cation of the north temperate taxa associated with the genus Staphylinus sensu lato, and comments on relevant subtribes of the tribe Staphylinini. The treatment contains the following: (1) a historical introduction to the past and recent classi®- cation of these genera; (2) a discussion of the characters used herein, many of them being unconventional and used for the ®rst time, and many of which are illustrated either by SEM photomicrographs or by line drawings; (3) a key to the north temperate subtribes of the tribe Staphylinini; (4) a discussion of the diagnostic characters of each of the ®ve relevant subtribes (Philonthina, Quediina, Anisolinina, Staphylinina, and Xanthopygina); (5) a key to the north temperate generic- and subgeneric-level taxa associated with the genus Staphylinus sensu lato; (6) type species information and a discussion of diagnostic characters for each taxon at the generic and subgeneric level; (7) a discussion of reasons for taxonomic and nomenclatural steps taken at both generic and speci®c levels; (8) a checklist of north temperate taxa; and (9) a list of the references mentioned in the discussions in the text (not including those associated with citations of scienti®c names). At the subtribal level, Craspedomerina, originally erected by Bernhauer (1911: 88) as Cras- pedomeri, is placed in synonymy with Philonthina. Triacrina, originally erected as Triacri by Bernhauer (1931: 84), is placed in synonymy with Xanthopygina (both syn. nov.). At the generic level, new concepts are presented for Staphylinus, Dinothenarus, Parabemus, Ocypus, Matidus, Pseudocypus, Tasgius, and Rayacheila. 1 Research Associate, Division of Invertebrate Zoology, American Museum of Natural History. 2 Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Central Experimental Farm, Ottawa, Ontario K1A 0C6, Canada. Copyright q American Museum of Natural History 2000 ISSN 0003-0082 / Price $8.30 AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_2 Allen Press x DTPro System File # 01cc 2 AMERICAN MUSEUM NOVITATES NO. 3287 New synonymies at the generic level are as follows (synonyms in brackets): Thoracostron- gylus Bernhauer, 1915 (5 Parontholestes Coiffait, 1982), Platydracus Thomson, 1858 (5 Neotasgius J. MuÈller, 1925), Parabemus Reitter, 1909 (5 Parocypus Bernhauer, 1915; Hypa- bemus Scheerpeltz, 1966), Ocypus Leach, 1819 (5 Goerius Westwood, 1827; Xanthocypus J. MuÈller, 1925), Pseudocypus Mulsant and Rey, 1876 (5 Protocypus J. MuÈller, 1923; Nuda- bemus Coiffait, 1982), Agelosus Sharp, 1874 (5 Apecholinus Bernhauer, 1933), Tasgius Ste- phens, 1829 (5 Pseudotasgius Seidlitz, 1891; Paratasgius Jarrige, 1952), Rayacheila Mot- schulsky, 1845 (5 Anodus Nordmann, 1837 [nec Spix, 1829]; Alapsodus Tottenham, 1939; Allocypus Coiffait, 1964; Metocypus Coiffait, 1964; Paralapsodus Coiffait, 1974). At the speci®c level, lectotypes are designated for Dinothenarus insignis (J. MuÈller, 1926), Ocypus almorensis (Cameron, 1932), Ocypus fulvotomentosus Eppelsheim, 1889, and Ocypus lewisius Sharp, 1874. Nine replacement names, 58 new synonymies, 224 new combinations, and numerous new statuses at both generic and speci®c levels are established; the latter four categories are to be found in the checklist. INTRODUCTION Platydracus, and Dinothenarus (Fam. VIII). Ocypus included a heterogeneous group of The complex of the genera and subgenera species that were subsequently assigned to associated with the genus Staphylinus con- Goerius and Pseudocypus (Fam. I), as well tains species that are large to very large; as the former genera Physetops (Fam. II), some of them are in fact the largest repre- Tasgius (Fam. III), and Anodus (Fam. IV). sentatives of the family Staphylinidae. One Erichson's concept turned out to be very im- might therefore assume that the group is well portant and was in principle accepted by known and that its higher taxonomy was most subsequent classical authors, including worked out in a reasonable way long ago. Ganglbauer (see below). In 1845, Motschul- Nothing could be farther from the truth, how- sky established the enigmatic monotypic ge- ever. nus Rayacheila, the status of which remained Historically, most of the species in the area uncertain for a long time but was mostly list- covered by this paper were described in the ed as a synonym of Goerius or Ocypus;it genus Staphylinus. Then, in the early 1800s, becomes the valid name for the taxon known Leach (1819) started splitting the genus until now as Alapsodus. In 1857, Kraatz en- Staphylinus (by that time already in some- tirely accepted Erichson's concept of the two what restricted sense) by adding the genera genera Staphylinus and Ocypus, with the ex- Creophilus, Emus, and Ocypus. Shortly clusion of the genera Emus, Creophilus, thereafter, Westwood (1827) added the genus Leistotrophus Perty, 1830 (actually misiden- Goerius, Stephens (1829) added the genera ti®ed; see below), and Physetops. In 1858, Trichoderma and Tasgius, Mannerheim Thomson erected the genera Platydracus and (1830) added Physetops, and Nordmann Dinothenarus (the latter became the valid (1837) added Anodus (preoccupied, eventu- name for the preoccupied Stephens' name ally becoming a synonym of Rayacheila). In Trichoderma). In 1860, Motschulsky estab- 1840, in his prodigious work ``Genera et spe- lished the monotypic genus Matidus that be- cies staphylinorum, insectorum coleoptero- came the valid name for a large group of rum familiae,'' Erichson divided the genus species that were recently listed mostly under Staphylinus into two genera: Staphylinus and Ocypus. In 1874, Fauvel combined Creophi- Ocypus, each subdivided into several ``Fam- lus and Emus under the former name, con- iliae'' (12 in Staphylinus and4inOcypus). tinued to use Leistotrophus as applied by Staphylinus included, in addition to numer- Kraatz (1857), and combined all previously ous tropical groups, the former genera Emus established taxa under one name, Staphyli- (Fam. I), Creophilus (Fam. II), a group of nus, which included four different groups. It species that eventually became members of is worth mentioning that in his ``Groupe 4'' the genus Ontholestes (Fam. VI), and a large he included ``Anodus Nordm.ÐOcypus heterogeneous group containing species that Steph.ÐMatidus, Rhagochila Mots.'' This is were subsequently assigned to Staphylinus, signi®cant because it shows the dif®culties in AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_3 Allen Press x DTPro System File # 01cc 2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 3 interpreting some of the taxa, particularly group. Some authors continued to use the those described by Motschulsky, a situation system of the two genera Staphylinus and that persisted until quite recently. In 1876, Ocypus, with the other taxa recognized at the Mulsant and Rey recognized all previously subgeneric level (e.g., Szujecki, 1980); some described taxa (mentioned above) as separate authors accepted Coiffait's concept entirely genera and added the genus Abemus.In (e.g., Outerelo and Gamarra, 1985), while 1895, Ganglbauer in his classical work ``Die others continued to use some of the taxa as KaÈfer von Mitteleuropa,'' which became the subgenera, mostly of the genus Ocypus (e.g., standard reference and identi®cation aid for Pilon, 1998). Some criticisms of Coiffait's a long period, again essentially accepted Er- concept appeared, and some of his taxa were ichson's concept, as it was modi®ed by placed in synonymy (e.g., Atlantogoerius Kraatz (1857), except that he considered the with Pseudocypus and Metocypus with Alap- species assigned by Erichson to ``Fam. IV'' sodus [DvorÏaÂk, 1984]), but in general the in- (Anodus) as members of the subgenus Ocy- stability in the higher taxonomy of the group pus s. str.; he also recognized Kraatz's mis- persisted until now. identi®cation of Leistotrophus and estab- At the speci®c level, J. MuÈller in his series lished the genus Ontholestes for the north of papers mentioned above, started a trend to temperate species included in Leistotrophus split many species (mainly in Ocypus and by Kraatz. Ganglbauer's treatment, including Pseudocypus) into subspecies, based mostly the erroneous concept of Ocypus s. str., was on differences in the shape of the aedeagus. accepted by most subsequent authors, includ- While many of the subspecies were valid ing Reitter (1909) in his ``Fauna Germani- geographical races, others turned out to be ca.'' unacceptable, mainly because the variability It is apparent at this point that the higher of the characters on the aedeagus in many taxonomy of the group this paper deals with species was not known at that time. Unfor- was worked out based mostly on European tunately, the splitting of the species into sub- fauna, although some ``exotic'' elements, species was picked up by some modern au- such as Agelosus and Miobdelus of Sharp thors, particularly by Coiffait, who in nu- from Japan, or Naddia (5 Caranistes Erich- merous papers (mainly 1956, 1964, 1970, son, 1840) of Fauvel from the northern