AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_1 Allen Press • 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 ᭧ 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 • 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 (ϭ Parontholestes Coiffait, 1982), Platydracus Thomson, 1858 (ϭ Neotasgius J. MuÈller, 1925), Parabemus Reitter, 1909 (ϭ Parocypus Bernhauer, 1915; Hypa- bemus Scheerpeltz, 1966), Ocypus Leach, 1819 (ϭ Goerius Westwood, 1827; Xanthocypus J. MuÈller, 1925), Pseudocypus Mulsant and Rey, 1876 (ϭ Protocypus J. MuÈller, 1923; Nuda- bemus Coiffait, 1982), Agelosus Sharp, 1874 (ϭ Apecholinus Bernhauer, 1933), Tasgius Ste- phens, 1829 (ϭ Pseudotasgius Seidlitz, 1891; Paratasgius Jarrige, 1952), Rayacheila Mot- schulsky, 1845 (ϭ 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 • 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 (ϭ Caranistes Erich- merous papers (mainly 1956, 1964, 1970, son, 1840) of Fauvel from the northern Ori- 1971, 1974) continued to split many species, ental region, were also registered. In the ear- again mostly in Ocypus and Pseudocypus, ly 1920s, J. MuÈller started to publish his se- using almost exclusively characters on the ries of papers dealing with Staphylinus. This aedeagus. This approach was eventually crit- series of nine papers extended over 20 years icized by some authors. For example, Sme- (J. MuÈller, 1923, 1924, 1925, 1926a, b, c, d, tana (1965b: 44) demonstrated, using a long 1932, 1943), in which he frequently dealt series of males of O. picipennis collected in with the east Palearctic fauna. It is signi®cant one mountain pass in Anatolia, that the shape that MuÈller's concept was very conservative, of the apical area of the aedeagus varied such as shown by the fact that he recognized only that at least two ``subspecies'' had to be rec- one genus, Staphylinus. Since the publication ognized as occurring in this high pass. Sub- of MuÈller's papers, most subsequent authors sequently, some of the ``subspecies'' were followed a concept of either one genus, synonymized (e.g., Smetana, 1965b, 1967, Staphylinus, or two genera, Staphylinus and 1971; Gusarov, 1992; Drugmand, 1998), or Ocypus. This situation lasted until Coiffait in at least their validity was questioned (e.g., several papers between 1956 and 1974 not Pilon, 1998). Many additional new synony- only elevated all previously described taxa to mies are presented in this paper (see the the generic level, but also provided numerous checklist). additional generic-level taxa (some based en- As indicated above, the group was in need tirely on characters on the aedeagus), partic- of a new taxonomic revision (both at the ge- ularly in the group that is united here under neric and speci®c levels) that would be based the name of Tasgius (Coiffait, 1956, 1964, on new characters and on the reevaluation of 1974). The publication of Coiffait's papers characters already in use, as well as on a destabilized the higher taxonomy of the large number of specimens of as many taxa AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_4 Allen Press • DTPro System File # 01cc

4 AMERICAN MUSEUM NOVITATES NO. 3287

as possible. This paper does exactly that. It Our colleagues from the Biological Re- is a taxonomic revision, exploring the use- sources Program at the Eastern Cereal and fulness of some new, hitherto unused char- Oilseed Research Centre, Agriculture and acters. We intentionally do not provide a cla- Agri-Food Canada, Ottawa, provided impor- distic analysis, which is almost obligatory tant help. Yves Bousquet and LubomõÂr Mas- these days. This may be objectionable to ner read the original draft of the manuscript some researchers, but we feel that a mean- (or portions of it), and provided many helpful ingful cladistic analysis of this group must comments that improved the manuscript. Go include the instrumental, enormously diver- Sato inked and ®nished the line drawings and si®ed fauna that is indigenous to the eastern- provided additional artistic support. most parts of the Palearctic Region, particu- larly within the mountainous areas of main- DISCUSSION OF CHARACTERS land China. Our knowledge of this fauna is only fragmentary at present, but it is improv- The following section is not a comprehen- ing steadily thanks to the extensive recent sive discussion of all characters of higher collecting in that area. Once our knowledge taxa in Staphylinina. It deals only with those becomes comparable to that of the western characters that are important for understand- portions of the Palearctic Region, a cladistic ing the taxonomic changes that are proposed analysis will have a better chance of clari- in this paper and does not attempt to ac- fying the phylogeny of the group. Hopefully, knowledge and comment on all papers pub- such future study will support, at least in lished on the subject. The states of these general terms, our ®ndings presented here. characters in the other subtribes of Staphy- At any rate, we hope that the results pre- linini, and occasionally in other tribes, are sented in this paper will be considered a pos- mentioned where it is considered useful. The itive step toward the reliable taxonomic treat- sexual characters, both male and female, are ment of this group, which is quite interesting seldom used and therefore not discussed in not only taxonomically, but also zoogeo- this section. There is no doubt that such char- graphically. acters will prove important for the higher In the following text, the author and the taxonomy of the group; however, it is dif®- date of publication are given only for the ®rst cult to assess them properly, mainly because mention of the taxa not included in the our knowledge of the extremely rich eastern checklist at the end of the paper. Palearctic fauna (particularly that of main- land China), which is instrumental for un- ACKNOWLEDGMENTS derstanding the evolutionary trends and re- lationships within the group, is extremely Many people provided help at different poor (see also above). levels and by different means during the Many of the characters used are uncon- work on this paper. We acknowledge that ventional and have not been used before. without this assistance it would have been Their appreciation requires careful examina- almost impossible to conclude this paper in tion, since they are often inconspicuous. the form it is presented here. We thank all Most of them are on the ventral side of the involved. However, special thanks go to a body. It is therefore recommended that if the few colleagues who provided outstanding as- European method of mounting the specimens sistance, be it by their willingness to discuss on plates is used, at least one specimen of the taxonomic concepts and the selection and the series (when available) should be mount- importance of morphological characters, or ed upside down, with the mandibles widely by their allowing us to use the results of their open and the coxae of the legs lifted and own research and literature searches. These pushed aside so that the structures on thorac- individuals are Lee H. Herman, American ic sterna are easily observable. The same ap- Museum of Natural History, New York; Al- plies to pinned or pointed specimens, partic- fred F. Newton, Jr., Field Museum of Natural ularly as far as the sternal characters are con- History, Chicago; and Harald Schillhammer, cerned. Naturhistorisches Museum, Wien, Austria. MANDIBULAE (MANDIBLES) AND THEIR AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_5 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 5

Fig. 1. Cuticular ridges on the surface of the head of a hypothetical member of the tribe Staphylinini.

Fig. 2. Prosternum of a hypothetical member of the tribe Staphylinini. AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_6 Allen Press • DTPro System File # 01cc

6 AMERICAN MUSEUM NOVITATES NO. 3287

Fig. 3. Mesosternum of a hypothetical member of the tribe Staphylinini.

TEETH: In general, there is a great variation the shape varies from short and stout man- in the shape of the mandibles, as well as in dibles with short and blunt apices in Naddia the development of the teeth at their medial (®g. 9) through moderately long and robust edge. The two mandibles are mostly dissim- mandibles in Staphylinus or Ocypus (®gs. 8, ilar in the number and shape of the teeth 10) to slender, ¯at mandibles that eventually (there are exceptions to this; e.g., the medial become sickle-shaped (Tasgius [Rayacheila]) edge of each mandible in Protogoerius brul- (®g. 4). The latter type is apparently the most lei has one similarly shaped tooth and the derived state. In the Staphylinina and Xan- same occurs in some species of Pseudocy- thopygina the basolateral ridge on the man- pus). The shape of the teeth sometimes varies dible is usually markedly developed, distinct- within one species (e.g., in Ocypus af®nis). ly removed from the lateral margin, and bor- Rarely, each mandible is devoid of teeth dered by a deep depression (®g. 124). It is (Tasgius [Rayacheila]) (®g. 4), or there is also well developed in Philonthina, but tends only one simple, sometimes not clearly dif- to be closer to the lateral margin and not bor- ferentiated tooth present on each mandible dered by a deep depression, whereas it is (Tasgius [Tasgius]) (®g. 5). The teeth are much closer to the lateral margin and poorly mostly situated in one plane, usually dorsal developed to absent in almost all Quediina. (e.g., members of Ocypus); however, in some MANDIBULAR PROSTHECA: Three states of cases the teeth appear in two planes, dorsal the mandibular prostheca are present. (1) and ventral, either on both mandibles (e.g., Medial edge of the prostheca is copiously cil- most members of Platydracus, particularly iate along its entire length, with more exten- the New World species) (®gs. 6, 7) or at least sive basal group of ciliae becoming less on one mandible, usually the left one (e.g., dense and shorter toward apex, with prosthe- members of Staphylinus) (®g. 8). The shape ca often appearing as more or less bilobed, of the mandibles varies considerably and is or even multilobed, as in Platydracus, Din- also in some taxa (e.g., Creophilus) subject othenarus, Abemus, and others (®gs. 11, 12); to sexual dimorphism (with the apical por- this type may represent the primitive state tion longer and sharper in males). In general, and seems to be prevalent in Philonthina and AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_7 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 7

Xanthopygina (although it is more simple in like setae in the glandular openings on the both). (2) Supporting structure of the pros- surface (®gs. 26, 27), but it is mostly asetose theca is long and lanceolate, with shorter cil- (®gs. 22, 28); rarely it is variously setose (as iae along medial margin becoming gradually in Ocypus [Matidus], Physetops, and Tas- longer to moderately long near apex, as in gius, all with setae also on the apical segment Agelosus, Ocypus (Pseudocypus), and Tas- of the labial palpus) (®gs. 25, 29, 30); some- gius (Tasgius) (®gs. 13, 14). (3) Supporting times it has ®ne longitudinal furrows at the structure of prostheca is parallel-sided, nar- base (Ontholestes,asinPhilonthus Stephens, row, elongate, and sometimes rodlike, with 1829) (®gs. 31, 32). The character of the se- ciliae present only at the apex, as in Emus, tation (or its absence), on the last segment of Protogoerius, Staphylinus, and Tasgius (Ray- the maxillary palpus was largely ignored in acheila) (®gs. 15, 16). the past. Although this character was men- MAXILLA: The chaetotaxy of the maxillary tioned in some recent publications (e.g., Haya- sclerites, as well as the shape and setation of shi, 1993: 285, 299), its importance for the the segments of the maxillary palpus, are tax- higher taxonomy in Staphylinina was still not onomically important. The apex of the pal- properly assessed. Most recently, the absence pifer (markedly variable and more useful at of the setation on this segment was used by the generic level) may bear one strong lateral Drugmand (1998: 58) to de®ne the Ophthal- seta (Anisolinus Sharp, 1889; Erichsonius micus group of species in the genus Ocypus, Fauvel, 1874; some Xanthopygina [Plociop- but this character state is in fact a synapo- terus Kraatz, 1857]; and all Tachyporinae we morphy of the entire subgenus Ocypus. Sim- examined) (®gs. 17, 19), or one strong lateral ilarly, the presence and number of setae on and one shorter, ®ner medial seta (most Staph- the apical segment of the labial palpus vary ylinina [®gs. 121, 156], Xanthopygina, and widely, but in most cases it seems that those Philonthina [usually very short to minute; with many setae also have the apex dilated ®gs. 20, 33], as well as many Paederinae), or to various extents. one strong subapical and two or three shorter LABIUM: The chaetotaxy of the mentum is apical setae (e.g., Physetops, Protogoerius, of importance for distinguishing the subtribes but none of the Xanthopygina or Quediina of Staphylinini. In general, the mentum bears we examined [as in ®g. 149]); the Quediina either only one long seta (Philonthina [®g. have members in almost all of the above cat- 33] [Erichsonius and some Ca®us Stephens, egories and some others. The stipes usually 1829 have one or more shorter additional se- bears one strong lateral and one more or less tae along or removed from the anterior mar- ®ne medial seta (Philonthina, Anisolinina, gin] and all Tachyporinae examined), or one Staphylinina, Xanthopygina) (®gs. 17±19, short and one long anterolateral setae (Ani- 33), or two strong basal setae (Quediina) (®g. solinina, Staphylinina, Xanthopygina [Plo- 21). ciopterus has only one]) (®gs. 19, 34). Near- The maxillary palpus, or rather the shape ly all genera in the subtribe Staphylinina and condition of its last segment as related seem to have two of these setae, except for to those of the penultimate segment, is tax- Emus and Naddia, in which the conformation onomically important. In Staphylinina, the of the mentum is unusual. In Ontholestes the last segment is usually narrower, subequal in setae are near the lateral margin, and the lat- length to the penultimate segment, and is eral seta is minute or absent. However, with- generally fusiform (®gs. 22, 23); sometimes in Staphylinina this character is not reliable, it is variably shorter than the penultimate as the size and number of setae vary consid- segment, but is still more or less fusiform erably, even within a single species (e.g., (®g. 24) or almost cylindrical (®g. 25); it is Tasgius ater, T. pedator), and most often the rarely as broad at the apex as at the middle setae are broken. The sizes of the setae vary, (where the last segment of labial palpus is but the medial seta is usually markedly stron- broadly expanded, as in Agelosus, Physetops, ger and longer than the lateral seta (®g. 34) and Tasgius [Rayacheila]), or the last two (this seems to be reversed in Paederinae). segments are broadly expanded (Naddia). The mentum itself is fairly consistent in The last segment always bears modi®ed peg- shape, with the anterior margin being fairly AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_8 Allen Press • DTPro System File # 01cc

8 AMERICAN MUSEUM NOVITATES NO. 3287

distinct and broadly emarginate (obsolete to the dorsal portion of the postoccipital su- medially in Abemus), and the posterior mar- ture; this character has been largely over- gin follows the shape of the submentum (ex- looked because of its position beneath the an- cept, e.g., in Dinothenarus, Emus, and Ocy- terior margin of the pronotum, and it was pus [Pseudocypus], which have a semicir- earlier deemed to be present only in the Xan- cular, ®ne basal carina extending variably be- thopygina (where it is strongest and mostly tween the basolateral angles (®gs. 38, 119, unobscured by punctures); it is usually ex- 120). The lateral margins are very long in tended anteriad toward the nuchal constric- Agelosus and Ocypus (Matidus). The ligula tion laterally, and it sometimes meets the is mostly entirely bilobed (®g. 35) or at least base of the infraorbital ridge. The postgenal variably emarginate medioapically (®gs. 36, ridge is anteriad to the ventral basal ridge at 37) in members of Staphylinina, but it may the base of the postgena, which is apparently be occasionally virtually nonemarginate completely absent only in Thinopinus (al- (e.g., in almost all Dinothenarus [Dinothen- though it is sometimes confused or obsolete arus]) (®g. 38). In the past, the shape of the ventrally); it runs parallel to, or is con¯uent ligula was commonly used to separate the old with, the base of the infraorbital ridge later- tribes Philonthini (ligula solid) and Staphy- ally. The infraorbital ridge, which, when linini (ligula emarginate/bilobed). However, complete (as in most Quediina), begins on as shown above, this approach is not reliable, the dorsolateral area of the neck, passes ven- particularly since some members of Philon- trally along the lateral portion of the post- thina have a slightly emarginate ligula, even genal ridge, to the lateral end of the nuchal in some species of Philonthus (e.g., in P. ridge (which may join it or become con¯uent caeruleipennis [Mannerheim, 1831]) (®g. with it, as in most Quediina and Glenus 20). Agelosus, most Dinothenarus, Miobde- Kraatz, 1857 [Xanthopygina]), and then it lus, Naddia, Ocypus (Ocypus, Pseudocypus), continues anteriad to near the base of the Protogoerius, and Wasmannellus have from mandible below the genal puncture. In the a few to many setae on the apical segment Xanthopygina it usually extends some dis- of the labial palpus, whereas Abemus, Cre- tance in front of the lateral bend of the post- ophilus, some Dinothenarus, Emus, Hadro- genal ridge (Elmas Blackwelder, 1952, some tes MaÈklin, 1852, Ontholestes, Platydracus, Eugastus Sharp, 1876, Gastrisus Sharp, Staphylinus, Thinopinus LeConte, 1852, and 1876), even more than in Quedionuchus Thoracostrongylus have no setae on either Sharp, 1884; in most Staphylinina it becomes apical segment. obsolete just in front of the lateral portion of SUBMENTUM: The anterior margin of the the postgenal ridge, and in most Philonthina submentum is usually straight or feebly out- it appears rudimentary, joining the postgenal wardly arcuate, with small lateral lobes. It is ridge immediately. The nuchal ridge follows broadly outwardly arcuate in Abemus, Cre- immediately behind the line of the nuchal ophilus, Miobdelus, and Tasgius (Rayachei- constriction dorsally; as mentioned above, it la), and broadly emarginate in Emus, Had- is very strong in most Quediina and becomes rotes, Physetops, Staphylinus, and some Ocy- con¯uent with the infraorbital ridge laterally, pus (Pseudocypus). It seems to be distinctly, whereas in most Staphylinina it reaches, or completely beaded only in Platydracus, Ocy- almost reaches, the base of the infraorbital pus (Pseudocypus), and Thoracostrongylus. ridge at various angles and distances from CUTICULAR RIDGES ON THE HEAD (®g. 1): the nuchal constriction (it is notably short lat- In the Staphylinini, there are several taxo- erally in some Ontholestes and Protogoerius, nomically important ridgesÐparticularly in and it is obsolete medially in Physetops, the postgenal area ventrally, and dorsally be- Creophilus, Hadrotes, and Thinopinus); it hind the nuchal constrictionÐthat are usually becomes obsolete laterally near the anterior consistent in their positions (when present). end of the dorsal basal ridge in Anisolinina, The ventral basal ridge anteriad to the con- most Philonthina (notably excepting Erich- tinuous gular/postoccipital suture, which is sonius), most Xanthopygina, and in all Pae- always present, originates near the base of derinae studied. The postmandibular ridge is the gula. The dorsal basal ridge is anteriad the most variable and most dif®cult to inter- AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_9 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 9

pret; when complete (as in many Xanthopy- visible punctures. In the Philonthina, the disc gina and some Philonthina), it extends from is virtually impunctate, bearing no more than just below the genal puncture behind the base microscopic micropunctulae (®g. 39). In of the mandible posteriad, very near the in- most Xanthopygina, it is only very ®nely, fraorbital puncture (often intermittently); al- sparsely punctate (though never only micro- ternatively, it may be de¯ected ventrad be- punctulate, as in Philonthina), with only a hind the base of the maxilla (as in many Phi- few moderate punctures at lateral margins. In lonthina). These two types are deemed to be Creophilus, Hadrotes, and Thinopinus, there homologous in origin, since they both often is a mixture of ®ne and moderate punctures, occur in one genus (notably Philonthus,in but they are broadly obsolete on the disc. In which both types and every extent of com- the rest of Staphylinina, and in Anisolinina, pleteness occur). The postmandibular ridge is the dorsal disc of the neck is usually densely also deemed to be distinct from the infraor- punctured, ranging from ®ne punctures (e.g., bital ridge, since the infraorbital ridge origi- in Abemus, Emus, and Protogoerius) (®g. 40) nates on the dorsum of the neck, and both to larger umbilicate punctures (Miobdelus), ridges are distinct in many Xanthopygina. In- as well as rugose punctures (Naddia) (®g. terestingly, the postmandibular ridge is the 41). only one present in the Tachyporinae (be- SUPERIOR AND INFERIOR MARGINAL LINES sides the postoccipital suture); it is complete OF PRONOTAL HYPOMERON: Two basically dif- in all Bolitobiini studied. In all Tachyporini ferent states are present in these lines: (1) the studied, there is a ridge originating behind two lines join at variable distance behind the the mandible that is de¯ected completely anterior angles of the pronotum as a result of around the base of the maxilla (note that Leu- de¯ection of the superior line (e.g., Dinoth- cotachinus Coiffait and Saiz, 1968 has a sep- enarus, Tasgius) (®gs. 42, 43) and continue arate postocular ridge). This ridge has been together in a broadly rounded arc along the interpreted by Stickney (1923: ®g. 173) as prosternal margin to the anterior margin of the remnant of the occipital suture, which the pronotum (Ontholestes is unusual in hav- may be correct; however, Xanthopygus ing the lines distinctly narrowly rounded at Kraatz, 1857 and some Philonthus (e.g., P. the anterolateral margin of the prosternum, longicornis Stephens, 1832) have a separate as in ®g. 44); or (2) the two lines never dis- ridge in this orientation beneath the maxilla, tinctly join because either the inferior line and the postmandibular ridge is distinct and (Anisolinina, some Philonthina, Agelosus, distant. Ocypus [Ocypus, Matidus]) (®gs. 45±48) or The gular sutures vary considerably, but the superior line (Creophilus, Hadrotes, Nad- usually they are moderately separated and dia miniata, Thinopinus, and Wasmannellus) most approximate somewhere in the basal (®g. 49) becomes indistinct or obsolete near half of the postgena, diverging anteriad to the the anterior angles; in the latter case, the su- submentum and posteriad to the base (except perior line is distinctly de¯ected under the in Ocypus [Matidus] and Physetops, where anterior angle (as in the Philonthina), and the they are distinctly more approximate before superior line is frequently de¯ected as a ®ne the middle). At the base of the gula there is line to meet the inferior line near the anterior usually a transverse impression that is usu- angle. This de¯ection is variable between ally distinct but poorly delimited anteriorly species (especially Creophilus and Naddia), (e.g., in Agelosus, Creophilus, Hadrotes, and sometimes even within a species. How- Thinopinus, Wasmannellus, and most Platyd- ever, there is signi®cant variability in the de- racus); it is delimited anteriorly by a narrow- velopment of the two lines throughout the ly rounded ridge in Abemus and Thoraco- Staphylinina. They may join in the anterior strongylus, and by a distinct transverse ridge half of the pronotal hypomeron at variable in Emus, Naddia, Ontholestes, and most Phi- distances from anterior angles, as in Staphy- lonthina. In most of the Quediina, it is visible linus dimidiaticornis (®g. 50) or Miobdelus only along the basal marginal carina. sp. (®g. 51); or they may run close to each PUNCTATION ON DORSUM OF NECK: In the other anteriorly, appearing as a more or less Quediina the dorsal disc of the neck bears no grooved carina, as in Platydracus stercorar- AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_10 Allen Press • DTPro System File # 01cc

10 AMERICAN MUSEUM NOVITATES NO. 3287

ius (®g. 52) or Ocypus tenebricosus (®g. 48); it apparently became fused with the peri- or they may remain variably widely separat- treme, as has the episternum (Acylophorus ed, as described above. Nordmann, 1837, Anchocerus Fauvel, 1905); In the Xanthopygina, on the other hand, it apparently is absent only in Heterothops the inferior line always continues separately Stephens, 1829 and is rudimentary in Hae- and uninterrupted along the prosternal mar- matodes Laporte, 1835 and Smilax Laporte, gin, usually to the anterior margin of the 1835. On the other hand, the proepimeron is pronotum, where it either meets the superior apparently completely absent in the Anisoli- line (most cases), or the superior line be- nina, Philonthina (except Erichsonius) (®g. comes obsolete near the anterior angles. This 45), and in the Bolitobiini (the hypomeron is state is considered to be one of the main sharply de¯ected and ¯attened against the characters of Xanthopygina (to the best of venter). The proepimeron ranges between our knowledge, there are no exceptions), al- these two extremes in the Xanthopygina and though additional characters need to be ap- Staphylinina, being always membranous plied (see below). Agelosus was treated re- when present. It varies from moderately de- cently as a member of the Xanthopygina veloped to absent in the Xanthopygina, and (Naomi, 1983: 582), based on the separation from well developed, broadly triangular (e.g., of the two lines: however, Hayashi (1991: Abemus [®g. 56], Creophilus) to poorly de- 179) correctly returned it to Staphylinina, veloped (e.g., Thoracostrongylus) to absent based essentially on arguments presented al- (e.g., Eucibdelus Kraatz, 1859, Physetops, ready by Sharp (1889: 111) when erecting Staphylinus [®g. 57]) in the Staphylinina, and the genus. As noted above, other members of it varies from moderate to absent in Dinoth- the Staphylinina, Anisolinina, and even the enarus and Tasgius. Philonthina (Hayashi, 1994: 127) (some Bis- PROSTERNUM (®g. 2): The anteromedial nius Stephens, 1829 and Leptopeltus Bern- portion of the prosternum varies greatly in hauer, 1906, and many Philonthus and Ra- the Staphylinina: in Dinothenarus (Dinoth- bigus Mulsant and Rey, 1876) also display enarus), Emus, and Ontholestes, it is highly this separation of the lines, although not so prominent, forming a rounded knob that widely. In all of these cases, however, it is slopes abruptly anteriad, and it markedly de- the inferior line that becomes indistinct or limits the transverse marginal depression; in obsolete near the coxal cavity. As a result of Abemus, Dinothenarus (Parabemus), Nad- these various conformations, the disc of the dia, and Thoracostrongylus (as in many Phi- pronotum is virtually invisible in ventral lonthina), it is moderately prominent, with a view in Xanthopygina (®g. 53) and is scarce- distinct, but broadly rounded slope anteriad; ly visible around the anterior angles in Ani- in Creophilus, Hadrotes, almost all Ocypus, solinina. In most Philonthina, where the su- and others, it slopes very gradually so that perior line is distinctly de¯ected under the the marginal depression is de®ned only by anterior angle, the de¯ected portion is visible its own posterior margin; it is highly variable from about the edge of the coxal insertion. in Platydracus and Staphylinus, although This pattern also exists in Staphylinina (e.g., never as prominent as in the ®rst group; the Naddia), or the de¯ected portion is visible Anisolinina have the whole anterior portion from the posterior edge of the coxal insertion of the prosternum almost ¯attened (Eucib- (e.g., Platydracus [®g. 52], and Protogoeri- delus in Staphylinina approaches this state). us) or even from closer to the posterior an- The prosternal keel is very variable in the gles of the pronotum (e.g., Dinothenarus [®g. Staphylinina, even within the genera, and 54], and Thoracostrongylus). thus only a few more-or-less consistent fea- PRONOTAL EPIMERON: The proepimeron is tures can be described: in Abemus and On- distinctly developed and highly sclerotized in tholestes, the keel forms a very strong carina, nearly all Quediina (®g. 55) (as well as in all convex in pro®le, starting between the mac- Tachyporini studied, except the proepimeron rosetae, with no tooth at the intercoxal ridge; is absent in Cileoporus Campbell, 1995, with in most Ocypus (especially Ocypus s. str.), its margin ¯attened against the venter, and in Naddia, Thoracostrongylus, and Wasmannel- Tachinoporus Cameron, 1928), except where lus (and less so in Platydracus), the medial AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_11 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 11

margin of the sternacostal ridge is extended In most genera the acetabulum behind the anteriad for some distance along the inter- intercoxal process is slightly to moderately coxal depression, often forming a highly depressed (in Physetops, there is a small prominent, ®nlike carina (®g. 58); in Physe- rounded carina behind it), but it is deeply tops, the intercoxal depression is very long, depressed in Miobdelus, Ocypus, Thinopinus, and the furcasternum is abruptly declivous, Protogoerius, and some Dinothenarus (Par- producing a sharp toothlike process at the abemus), and there are variously shaped ca- sternacostal ridge; in most other genera the rinae behind the projection in Ascialinus, carina is weak, con®ned to the area adjacent Hadrotes, Agelosus, Tasgius (®gs. 62, 154), to the sternacostal ridge, or it is obsolete, as and some Ocypus (especially Pseudocypus). in Creophilus, Emus, and Thinopinus. METASTERNUM: The anterior metasternal In most genera the sternacostal ridge is projection is very long in the Anisolinina and slightly oblique and relatively straight; in in Ocypus (Ocypus) weisei (®g. 63), Ontho- Emus, it is distinctly transverse and much lestes, and Thoracostrongylus, with the apex closer to the two macrosetae mesially; in ranging from narrowly to broadly rounded. Dinothenarus s. str. and Hadrotes, it is dis- The margins converge at an angle that is dis- tinctly arched posteriad near the carina, ap- tinctly less than rectangular in Abemus and proaching the posterior furcasternal margin most Platydracus, with the apex narrowly to (®g. 54); this pattern also occurs in most Tas- moderately rounded. They converge subrec- gius (Rayacheila), but the furcasternum is tangularly in Dinothenarus (Dinothenarus), long. with the apex broadly rounded (it is more MESOSTERNUM (®g. 3): In all Staphylinini variable in Dinothenarus [Parabemus]); or studied, the disc of the mesosternum bears a they converge rectangularly, with the apex series of long setae in various positions that broadly rounded in Wasmannellus.InMiob- are arranged in a subtransverse- to V-shaped delus, Physetops, and almost all Ocypus, pattern. In Ontholestes, they are in a sub- Protogoerius and Tasgius, the margins con- transverse arrangement on either side of a verge quite obtusely, with the apex very distinct longitudinal medial carina. The setae broadly rounded (except the apex is more are in moderately impressed punctures ar- narrowly rounded in some Ocypus [Matidus] ranged subtransversely in Creophilus, Emus, and Tasgius [Tasgius]). The apical and lat- Staphylinus, and Thinopinus. Setae are on the eral margins of the projection are obsolete in edge of large, rounded depressions forming Creophilus, Emus, Hadrotes, Platydracus an irregular, broadly U-shaped ridge in Din- (Chaetodracus), Staphylinus (®g. 64), and othenarus (®g. 59) and Physetops. They are Thinopinus. on small elevations in broadly U-shaped pat- APEX OF HIND COXAE: In the Staphylinini terns in Naddia or Platydracus, or are rough- the dorsal apicolateral lobe of the hind coxae ly parallel to the margins of the projection, bears a series of setae that are usually dis- as in Abemus, or are along or behind an el- tinctly coarser than the rest of the setae on evated ridge which may be feeble to well el- the surface, frequently aligning along a fee- evated, irregular to even, and subtransverse ble ridge. The number and size of these setae to V-shaped in most other genera of Staphy- vary considerably in the Staphylinina, but linina (®g. 60), as well as in Amichrotus (®g. there are a few notable trends: in most Din- 98) and Philonthina (®g. 87). The margins of othenarus (s. str. in particular), Emus, and the mesosternal intercoxal process converge some Platydracus, there are more than three acutely or subacutely in most Staphylinina distinctly coarser setae in the apical series (®gs. 59, 60), with the angle being most var- (®g. 65); in Naddia and Thoracostrongylus, iable in Ocypus and Platydracus (where the there appears to be only one distinct spine, apex can be roundly truncate); they converge usually located distinctly subapically in subrectangularly in Protogoerius, Creophi- Thoracostrongylus;inCreophilus, Eucibde- lus, Dinothenarus (Dinothenarus), and lus, Hadrotes, Leistotrophus, Ontholestes, Ontholestes;inEmus, the margins are broad- Staphylinus, and Thinopinus, they are almost ly arcuate and broadly truncate apically (®g. indistinguishable in size from the other setae, 101). or they are completely absent (®g. 66); in AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_12 Allen Press • DTPro System File # 01cc

12 AMERICAN MUSEUM NOVITATES NO. 3287

most of the rest of the Staphylinina there are ing onto anterior margin of pronotum (®g. usually two or three coarse spines. 75). Pronotal hypomeron strongly in¯exed EMPODIAL SETAE: The importance of the below anterior angle of pronotum, there- presence or absence of the pair of empodial fore meeting prosternum at a sharp angle setae between the tarsal claws for subtribal (up to 90Њ), with both sclerites separated by notosternal suture (®g. 76). Head al- taxonomy was recently published by Schill- most always with complete infraorbital hammer (1998b). The empodial setae are ridge, at least extending signi®cantly in present in the Anisolinina, Staphylinina, front of postgenal ridge (®g. 77). Neck Xanthopygina, and Quediina (®gs. 67±69), without dorsal basal ridge (®g. 78) ..... but not in the Philonthina (including Cras- ...... Quediina pedomerina) (®gs. 70, 71). A notable case is 4. Superior and inferior marginal lines of the genus Thinopinus, in which each claw pronotal hypomeron variably widely sep- may bear from 3 to 5 empodial setae. arated throughout, never joined (®gs. 46, 48,53) ...... 5 ± Superior and inferior marginal lines of KEY TO NORTH TEMPERATE SUBTRIBES OF pronotal hypomeron joined at variable dis- STAPHYLININI tance behind or at anterior angles of pron- 1. Tarsal formula 5, 4, 4. Maxillary and la- otum (®gs. 42±44, 50, 51) ...... bial palpus each conspicuously slender ...... Staphylinina (pars) and elongate. General habitus resembling 5. Superior marginal line of pronotal hypom- genus Tachyporus of Tachyporinae . . . eron not de¯exed, no part of de¯ected por- ...... Tanygnathinina tion of pronotal disc visible in ventral view ± Tarsal formula 5, 5, 5. Maxillary and la- (®g. 79). Punctures on disc of neck ®ne, bial palpus each no more than moderately sparse, with only a few moderate punctures slender and long. General habitus not re- laterally (®g. 80). Head almost always with sembling genus Tachyporus ...... 2 long postmandibular ridge (®g. 81) ..... 2. Middle portion of disc of neck virtually ...... Xanthopygina (Triacrina, syn. nov.) impunctate, with no more than microscop- ± Superior marginal line of pronotal hypom- ic micropunctulation (®g. 39) ...... 3 eron de¯exed; de¯ected portion of prono- ± Middle portion of disc of neck punctate, tal disc visible in ventral view, at least at with coarseness and density of punctation anterior angles (®g. 73). Punctures on disc variable, but always ranging above micro- of neck dense, moderate to coarse (®gs. scopic micropunctulation (®gs. 40, 41); if 40, 41). Head with short to rudimentary impunctate, then pronotum with roughly postmandibular ridge (®g. 82) ...... 6 punctate pit at each lateral margin in front 6. Superior marginal line of pronotal hypom- of basal margin (Creophilus of Staphyli- eron not distinctly de¯ected under anterior nina) ...... 4 angle of pronotum before continuing onto 3. Empodial setae between claws of all tarsi anterior margin of pronotum (®g. 83). absent (®gs. 70, 71). Superior marginal Ventral basal ridge extended anteriad for line of pronotal hypomeron de¯exed under considerable distance parallel to gular su- anterior angle of pronotum before continu- ture, at least as far as postgenal ridge. Sec- ing onto anterior margin of pronotum ond segment of maxillary palpus with me- (®gs. 72, 73). Pronotal hypomeron no dial margin obtusely rounded apically, more than moderately in¯exed below an- with insertion of third segment situated en- terior angle of pronotum, meeting proster- tirely in lateral half of apex (®g. 84) .... num at a very ¯at angle, with both scler- ...... Anisolinina ites fused, and notosternal suture therefore ± Superior marginal line of pronotal hypom- absent (®g. 73). Head without apparent in- eron de¯exed under anterior angle of pron- fraorbital ridge (®g. 74). Neck with dorsal otum before continuing onto anterior mar- basal ridge (®g. 39) (excluding Erichson- gin of pronotum (®gs. 54, 56). Ventral bas- ius, and Gabronthus Tottenham, 1955) . . al ridge not extended signi®cantly parallel ... Philonthina (Craspedomerina, syn. nov.) to gular suture, but meeting it transversely ± Empodial setae between claws of all tarsi or acutely. Second segment of maxillary present (®gs. 67±69). Superior marginal palpus with apex transversely truncate, line of pronotal hypomeron situated entire- with insertion of third segment situated for ly on lateral margin, not de¯exed under most part at middle of apex (®g. 85). . . . anterior angle of pronotum before continu- ...... Staphylinina (pars) AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_13 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 13

NORTH TEMPERATE SUBTRIBES OF (as in Xanthopygina). Neck with middle por- STAPHYLININI tion of disc virtually impunctate, with no more than microscopic micropunctulation The subtribe Tanygnathinina, containing (®g. 39). Nuchal ridge present mesially, usu- only the genus Atanygnathus Jakobson, ally becoming obsolete near apex of dorsal 1909, is quite distinctive and is therefore not basal ridge (®g. 39). Infraorbital ridge usu- discussed any further. For the remaining sub- ally rudimentary, extending at most very lit- tribes, some sets of characters that are con- tle in front of postgenal ridge. Dorsal basal sidered important for distinguishing them, ridge rarely absent, with lateral extensions are discussed further. It became apparent dur- usually directed almost directly anteriad (®g. ing this study that some subtribes and their 39), often weak and ending toward base of conventional characters needed to be reas- rudimentary infraorbital ridge anteriad. Ven- sessed, and that new characters needed to be tral basal ridge usually follows closely along established for a meaningful characterization base of gular sutures up to postgena (as in of these groups. Some of these characters Anisolinina). Gular sutures usually close to- show ranges of variation and apparently de- gether or united basally; transverse depres- veloped in a parallel fashion several times in sion at base of gula usually delimited by an- different lineages. This is particularly true for teriorly broadly arcuate ridge from lateral the con®guration of the two marginal lines margins of base. of the pronotal hypomeron (superior and in- Pronotal hypomeron with superior margin- ferior lines), or for the development of the al line distinctly de¯exed under anterior an- ligula (emarginate/bilobed versus solid). The gle of pronotum, so that de¯exed portion of importance of both characters was undoubt- pronotal disc visible in ventral view (®g. 45), edly overrated until now. On the other hand, and with inferior marginal line almost always the presence of rare exceptions to otherwise joining it behind or along prosternal lateral reliable characters does not really affect their margin; both lines continue together below importance. Such exceptions are in general anterior angle and along anterior margin of not rare and are well known to any practising pronotum; if two lines not joined, inferior taxonomist, especially where broadly line becomes obsolete near lateral margin of evolved characters are secondarily lost. procoxal cavity (as in some Staphylinina); hypomeron no more than moderately in- SUBTRIBE PHILONTHINA ¯exed before anterior angles of pronotum, All members of this subtribe are without meeting prosternum at very ¯at angle, with empodial setae between the claws of all tarsi both sclerites fused, and notosternal suture (®gs. 70, 71). This character state separates therefore absent (®g. 45) (as in Staphylinina them positively from members of any other and Xanthopygina). Proepimeron absent. subtribe. They also share the character states Prosternum with basisternum evenly round- delineated below. ed, only moderately convex, without carina Ligula entire, no more than slightly sinuate medially before sternacostal suture (®g. 86). medioapically (e.g., in Philonthus caerulei- Mesosternum with series of large setae situ- pennis) (®g. 20, as in members of Quediina), ated in broadly arcuate pattern, usually just with sinuation rarely more pronounced. Pal- behind well-formed, broadly arcuate trans- pifer with medioapical seta minute to less verse ridge near middle of its length (®g. 87); than half as long as long lateroapical seta mesosternum without medial carina. Meta- (®gs. 18, 20). Labial and maxillary palpus sternum with margin of anterior projection with apical segment asetose, elongate fusi- almost always very broadly rounded, short form, or shorter and tapering apicad, not between middle coxae (®g. 87). broadened apically. Mentum rarely with NOTES: Neck without dorsal basal ridge in more than one anterolateral seta on each side Erichsonius and Gabronthus. Proepimeron (®g. 20). Postmandibular ridge usually pre- present, and prosternum with short carina in sent (often obsolete only in males with swol- front of sternacostal ridge in Erichsonius. len genae), either de¯exed behind maxilla or Mesosternum without transverse ridge in following line of large punctures around eyes Craspedomerus Bernhauer, 1911 (®g. 61), AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_14 Allen Press • DTPro System File # 01cc

14 AMERICAN MUSEUM NOVITATES NO. 3287

Erichsonius, and some species of Hesperus Velleius Leach, 1819, and Velleiopsis Fair- Fauvel, 1874. Metasternum with anterior maire, 1882) or by several shorter setae projection more narrowly rounded in some (Hemiquedius Casey, 1915 ), by one minute species of Ca®us and Hesperus, rarely par- seta (Bolitogyrus Chevrolat, 1842, Heinzia tially obsolete in some species of Ca®us and Korge, 1972, Heterothops, Quedius [Que- Neobisnius Ganglbauer, 1895. dionuchus]), or apparently without additional The subtribe Craspedomerina, which was setae (Acylophorus, Anaquedius Casey, established by Bernhauer (1911: 88, as Cras- 1915, Anchocerus, Beeria Hatch, 1957, Que- pedomeri) for the genus Craspedomerus, was dius [Raphirus Stephens, 1829], Valdiviodes based on the supposed presence of an addi- Smetana, 1981). Mentum with two antero- tional oblique line connecting the superior lateral setae on each side (most Quedius [®g. and inferior lines of the pronotal hypomeron. 90], sometimes separated as in Hemiquedius, However, this line is in fact the ventrally de- Quedius [Paraquedius Casey, 1915, Raphi- ¯ected superior line, and the pronotal lateral rus]) or with several setae (Velleius), or one margin is narrowly, somewhat irregularly seta (Anaquedius, Beeria, Bolitogyrus, Het- swollen, imitating a xanthopygine-like su- erothops); there are no setae on the mentum perior line (®g. 88). The empodial setae are in Acylophorus. Maxillary palpus usually missing in Craspedomerus (®g. 71), and the without setae. Gula with transverse basal im- genus agrees with all characters listed above, pression very narrow, distinct only along con®rming the assignment of this genus to basal marginal carina, apparently absent in Philonthina, as suggested recently by Haya- Heterothops and Smilax. Mandible with dor- shi (1997: 491). solateral ridge ranging from poorly de®ned, to absent (Acylophorus, Anaquedius, Bolito- SUBTRIBE QUEDIINA gyrus, Hemiquedius, Heterothops, Indoque- dius Cameron, 1932, Valdiviodes), and al- All members of this subtribe have the ways very close to lateral margin when pre- pronotal hypomeron strongly in¯exed below sent (®g. 91). Postmandibular ridge absent; the anterior angles of the pronotum, meeting ventral basal and postgenal ridges present. the prosternum at a sharp angle (up to 90Њ), Neck with middle portion of disc impunctate. and separated from it by the notosternal su- Pronotal hypomeron with superior margin- ture (®g. 76); the anterior angles of the pro- al line situated entirely on lateral margin, not notum are produced beyond the anterior mar- de¯exed under anterior angle of pronotum gin of the pronotum (®g. 76) (the in¯ection before continuing onto anterior margin of is less apparent in some taxa, e.g., Quedio- pronotum (®g. 75); inferior marginal line of nuchus). The infraorbital ridge is usually pronotal hypomeron becoming obsolete complete ventrolaterally on the head (®g. along anterolateral margin of coxal cavity in 77); this ridge is never complete in any other typical members of Quediina (®g. 76). An- subtribe of Staphylinini. The nuchal ridge is terior marginal line of pronotum gradually usually very strong and becomes con¯uent becoming separated from the margin mediad with the infraorbital ridge, but the infraorbit- in many genera, also often becoming obso- al ridge often can be seen to join the post- lete. Proepimeron highly developed and well genal ridge laterally as a ®ne line. The dorsal sclerotized in almost all typical Quediina (in- basal ridge is absent from the neck, a char- cluding Beeria, and as in most Tachyporini). acter shared only with Tachyporinae, Erich- Prosternum with basisternum usually mod- sonius, and Gabronthus of Philonthina. The erately prominent, evenly convex anteriad, members of Quediina also share the character relatively ¯at behind two long setae, and usu- states delineated below. ally without carina (as in Philonthina). Ster- Ligula minutely emarginate medioapically nacostal ridge usually straight, distinctly (as in members of Anisolinina; it is distinctly oblique. Furcasternum usually relatively de- emarginate or bilobed in most Staphylinina; clivous, often moderately carinate. Mesoster- ®gs. 35±37), rarely entire (®g. 89). Palpifer num with long setae situated (often not in with one strong apical seta, accompanied by regular pattern) on and/or behind broadly one shorter seta (most Quedius, as well as raised discal area, which follows contours of AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_15 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 15

margin of projection, so that margins appear perior line is not distinctly de¯exed under the depressed, well de®ned, never with trans- anterior angles of the pronotum, and the verse carina. Metasternum projects very little pronotum is highly convex; as a result, only between middle coxae (as in most Philonthi- a very narrow portion of the dorsal surface na), or not at all (e.g., Acylophorus). Empo- of the pronotum is visible in ventral view at dial setae present between claws of all tarsi. the anterior angle. The members of this sub- NOTES: Maxillary palpus with setae in tribe also share the character states delineated some more unusual Quediina (Anchocerus, below. Astrapaeus Gravenhorst, 1802, Euryporus Ligula minutely emarginate medioapically Erichson, 1839, Indoquedius, Strouhalium (®g. 93). Palpifer with only one strong latero- Scheerpeltz, 1962). Infraorbital ridge strong- apical seta (Anisolinus) (®g. 17), or with one ly reduced or missing entirely in Astrapaeus, additional shorter, ®ner medioapical seta Beeria, Heinzia, Quedius (Quedionuchus, (Amichrotus Sharp, 1889) (®g. 95). Both la- some Microsaurus Dejean, 1833), and Val- bial and maxillary palpi with apical segment diviodes Smetana, 1981. Nuchal ridge pre- asetose, elongate, distinctly fusiform; second sent only dorsally in Beeria, rudimentary in segment of maxillary palpus with medial Valdiviodes, and not distinctly converging margin obtusely rounded apically, with in- with infraorbital ridge in Anchocerus, Het- sertion of third segment entirely in lateral erothops, Quedius (Megaquedius Casey, half of apex (®g. 84). Mentum with two an- 1915, Microsaurus). terolateral setae on each side. Mandibles Pronotal epimeron apparently fused with long, sickle-shaped (®g. 94), each with one peritreme in Acylophorus and Anchocerus; broad tooth at middle (®g. 96), with dorso- hypomeron de¯exed against margin of peri- lateral ridge well developed, close to lateral treme and proepimeron apparently absent in margin; mandibular prostheca long, lanceo- Heterothops. Sternacostal ridge moderately late, with single series of long, individual produced anteriad along keel in Anaquedius, plumose ciliae clearly discernible, shortening Beeria, and Quedius (Megaquedius, Para- near apex. Gula with transverse basal im- quedius, Quedionuchus, some Microsaurus). pression well delimited anteriad, either by Empodial setae sometimes unevenly devel- narrowly rounded ridge or by abrupt impres- oped (Acylophorus lineage). sion of gula in front. Postmandibular ridge In the above diagnosis, several genera rudimentary to short; ventral basal ridge fol- have been omitted because they have very lowing base of gular sutures closely up to few characters in common with the Quedi- postgena; base of infraorbital ridge extending ina; on the other hand, many are typical of little beyond lateral bend of postgenal ridge, Philonthina or Xanthopygina (e.g., Glyphes- arching toward nuchal ridge dorsad on neck, thus Kraatz, 1858 and Moeocerus Fauvel, but (together with basal ridge) obscured me- 1899, which are actually members of Philon- sad by punctures on disc (especially in Ami- thina, and Haematodes and Scariphaeus Er- chrotus); nuchal ridge present dorsally, be- ichson, 1839, which will be dealt with at a coming obsolete near apex of arms of dorsal later date [in prep.]). basal ridge (as in Philonthina). Neck with moderately dense, moderate punctures on SUBTRIBE ANISOLININA disc (as in Ocypus s. str., Platydracus, and others of Staphylinina). In all north temperate members of this Pronotum with hypomeron visible in lat- subtribe, the superior and inferior marginal eral view for most of its length except below lines of the pronotal hypomeron are not anterior angle of pronotum, where it is slight- joined, and the inferior line becomes obsolete ly in¯exed, therefore meeting prosternum at along the lateral margin of the prosternum ¯at angle; both sclerites fused, notosternal (®g. 92) (as in Agelosus or Ocypus [Matidus] suture therefore absent (as in Philonthina, of Staphylinina), but the hypomeron clearly Staphylinina, and Xanthopygina). Proepime- does not extend anteriad along the anterior ron absent. Prosternum with entire basister- margin of the pronotum. In the Anisolinina, num depressed or ¯attened, with very little this conformation may occur because the su- convexity anteromedially, so that intercoxal AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_16 Allen Press • DTPro System File # 01cc

16 AMERICAN MUSEUM NOVITATES NO. 3287

depression well delimited anteriad, and/or ward subocular puncture. This ridge is lon- well-de®ned transverse ridge present at var- gest in Ontholestes, distinct and almost iable distance behind anterior margin (®g. reaching infraorbital puncture in Abemus, 97). Metasternum with anterior projection Thoracostrongylus, and Platydracus, and very long, apex narrowly (Anisolinus)to variably short to rudimentary in Dinothena- moderately rounded (Amichrotus) (®g. 98). rus, Ocypus (Pseudocypus), and Staphylinus. Empodial setae present between claws of all Neck always with dorsal basal ridge (as in tarsi, but usually very small (®g. 67). Philonthina and Xanthopygina), although NOTES: Many members of this subtribe re- usually with anterior extensions confused by semble in general habitus, and even in the dense punctation on disc. Gula with trans- shape of the mandibles and in the develop- verse impression at base usually distinct, ment of the mouthparts, those of the subtribe usually not well delimited anteriad. Base of Philonthina, particularly some species of infraorbital ridge extends at most slightly in Philonthus from the Cyanipennis group front of postgenal ridge laterally, or is en- (Schillhammer, 1998a: 102). The two antero- tirely rudimentary where postgenal ridge ex- lateral setae on each side of the mentum, the tends laterad (as in Platydracus), apparently densely punctate surface of the neck, the only more signi®cantly extended anteriad in long metasternal intercoxal process, and Agelosus. Nuchal ridge almost always com- presence of the empodial setae differentiate plete, reaching or coming close to base of them immediately. infraorbital ridge laterally. Neck with disc densely punctate, with ®ne to umbilicate SUBTRIBE STAPHYLININA punctures (®gs. 40, 41) generally separated by at most one diameter of puncture. With the exception of some species of Pronotum with hypomeron no more than Dinothenarus s. str. (®g. 38), all members of moderately in¯exed at anterior angles of this subtribe have the ligula emarginate or pronotum, meeting prosternum at a very ¯at bilobed, although the emargination is occa- angle, with both sclerites fused, and noto- sionally more or less minute (®gs. 35±37). sternal suture therefore absent (®g. 56) (as in The superior marginal line of the pronotal Philonthina, Anisolinina, and Xanthopygina). hypomeron is de¯ected under the anterior an- Proepimeron, when present, always membra- gles of the pronotum, so that the de¯ected nous and varying from well to poorly devel- portion of the disc is visible in ventral view oped (®gs. 51, 56). Acetabulum of middle (®gs. 42±44); the superior and inferior mar- coxae often with deep depression behind in- ginal lines are almost always joined along or tercoxal process of mesosternum (unique behind the posterolateral margin of the pro- within Staphylinini), as in Miobdelus, Ocy- sternum (®gs. 42±44). Members of Staphy- pus, Thinopinus, Protogoerius, and some linina also share the character states delin- Dinothenarus (Parabemus), also with vari- eated below. ably shaped carinae behind projection in Palpifer with one strong lateral and one Agelosus, Ascialinus, Hadrotes, Tasgius (®g. ®ner medial seta apically, medial seta at least 154), and some Ocypus (especially Pseudo- half as long as lateral seta (®gs. 121, 156); cypus). Metasternum usually distinctly pro- or with one strong subapical and several jecting between middle coxae. Empodial se- shorter setae (Creophilus, Emus, Hadrotes, tae present between claws of all tarsi (®g. Miobdelus, Ocypus [most members of Ocy- 68). pus and Matidus; ®g. 149], Physetops, Pro- NOTES: Mentum with cluster of variable togoerius, Thinopinus). Mentum usually with anterolateral setae in Emus and Naddia. one long anterolateral seta and one shorter Mandible with dorsolateral ridge weaker and seta laterad to it (®g. 99). Mandible usually closer to margin in Tasgius (Rayacheila), in with well-developed dorsolateral ridge, well which mandible is unusually narrow. Man- removed from lateral margin and bordered dibular prostheca highly variable in Staphy- by deep depression (as in Xanthopygina). linina, but unusual developments (e.g., dis- Postmandibular ridge almost always of xan- tinctly bilobed supporting structure as in thopygine type when present, directed to- Abemus [®g. 12] or Dinothenarus, or sup- AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_17 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 17

porting structure long, lanceolate or parallel- joins the superior line. In cases where the sided with setae only near the apex [®gs. 13± inferior line becomes obsolete at the antero- 15]) have not been seen in the other subtribes lateral angle of the prosternum (as in Algon of Staphylinini. Postmandibular ridge de- Sharp, 1874, Allostenopsis Bernhauer, 1921, ¯ected behind maxilla in Eucibdelus. Gula Eugastus, and Tympanophorus Nordmann, with transverse basal impression indistinct in 1837), the two lines are widely separated and Eucibdelus, weak in Protogoerius and Tas- the superior line is never de¯ected below the gius (Tasgius), in which gula tumid to near anterior angles of the pronotum, nor is the base; delimited anteriad by narrowly rounded disc of the pronotum visible in any part in arcuate ridge in Ontholestes and Thoraco- ventral view. These and the other members strongylus; limited by broader, more poste- of Xanthopygina also share the character riad depression in gula in Abemus, Emus, and states delineated below. Naddia. Nuchal ridge obsolete laterally in Ligula entire or no more than slightly sin- Miobdelus, some Ontholestes, and Protogoe- uate medioapically (Plociopterus, ®g. 19). rius, and obsolete mesially in Creophilus, Palpifer with one strong lateral, slightly sub- Hadrotes, and Thinopinus. Neck with punc- apical seta, and usually one ®ner medioapical tures entirely obsolete on disc in Creophilus, seta only up to half length of long seta. Men- Hadrotes, and Thinopinus. Pronotal epime- tum with two anterolateral setae: one long ron absent in Eucibdelus, Ocypus, Physetops, medial seta, one ®ne lateral seta. Mandible Protogoerius, and Staphylinus (®g. 57), Was- with basolateral ridge usually highly devel- mannellus, and in some Dinothenarus and oped, distinctly removed from lateral margin, Tasgius. and bordered by deep depression. Gula with Superior line of pronotal hypomeron be- transverse impression at base almost always coming variably indistinct or obsolete near well developed, delimited anteriorly by ei- anterior angle in Creophilus, Hadrotes, Thin- ther a narrowly rounded, anteriorly arcuate opinus, and Wasmannellus (®g. 49); inferior ridge (Eugastus, Nordus Blackwelder, 1952, line very close and parallel to superior line Philothalpus Kraatz, 1857, Elmas, Plociop- along prosternal margin in Ocypus (Ocypus terus, Styngetus Sharp, 1884, Tympanopho- and Matidus) (®g. 48); superior line poorly rus), or more typically by a distinct trans- de¯ected and two lines widely separated in verse carina that limits base of depressed Agelosus (®g. 46), but Agelosus shares many gula. Infraorbital ridge extending signi®cant- characters with Ocypus (Ocypus). Metaster- ly anteriad of lateral bend of postgenal ridge num with anterior projection very broadly (as in some Quediina, but unlike Philonthina rounded in Ascialinus, Physetops, and Pro- and Staphylinina). Neck with disc only ®nely togoerius. punctate (though never only micropunctulate as in Philonthina), with moderate punctures SUBTRIBE XANTHOPYGINA only laterally (®g. 80). Hypomeron of pronotum no more than Most members of this subtribe have a moderately in¯exed below anterior angles of complete postmandibular ridge on the head pronotum, meeting prosternum at a ¯at angle (®g. 81), although frequently irregular, al- (often angled slightly anteriad), with both ways distinct anteriorly, and usually reaching sclerites fused, and notosternal suture there- past subocular puncture, with at least curved fore absent (®g. 53). Proepimeron present in ridges extending to near postocular puncture. relatively few genera, always membranous The dorsal basal ridge is always present on and never more than moderately developed the neck (®g. 80). The superior and inferior (unlike in most Quediina). Prosternum with marginal lines of the pronotal hypomeron are basisternum rarely carinate, and then only not joined and are well separated. The infe- weakly so in front of sternacostal ridge, oth- rior line always continues uninterrupted and erwise sternacostal ridge usually strong and separately along the prosternal margin and projecting medially. Mesosternum never with almost always continues around the anterior distinct, arcuate ridge before series of long angles of the pronotum to the anterior margin setae (unlike Philonthina), with highly vari- of the pronotum (®g. 79), where it usually able placement. Acetabulum of middle coxae AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_18 Allen Press • DTPro System File # 01cc

18 AMERICAN MUSEUM NOVITATES NO. 3287

never more than moderately depressed be- they also differ in their ecology. Members of hind mesosternal intercoxal process. Meta- the Eucibdelus lineage are arboricol±¯oricol sternum with anterior projection always well (there are a few exceptions, e.g., Trichocos- developed, margins always complete and metes Kraatz, 1859), and members of the re- never converging more than rectangularly maining genera are restricted to saline sea along coxal margins, although apex some- beaches, with both of these habitats being times broadly rounded. Empodial setae al- strictly avoided by members of the Staphy- ways present between tarsal claws. linus complex. NOTES: Postmandibular ridge short and de- 1. Disc of pronotum and most of disc of ¯ected slightly ventrad behind maxilla in El- neck virtually impunctate and asetose . . mas, Glenus, and Tympanophorus; Triacrus ...... Creophilus Leach, 1819 Nordmann, 1837 has short sections near ± Disc of pronotum and disc of neck vari- postocular puncture. Pronotal hypomeron ably punctate and setose ...... 2 with superior line becoming obsolete near 2. Intercoxal process of mesosternum wide, anterior angles in Dysanellus transverseru- transverse apically (®g. 101), middle gosus Bernhauer, 1921, some Gastrisus, coxae therefore widely separated. First most Plociopterus, and most Xanthopygus ®ve antennal segments lacking dense ap- (this is clearly a secondary fading of the pressed pubescence, last six segments forming slightly differentiated, loose line). Palpifer with one long and two shorter club. Hind trochanters sexually dimor- setae apically in Trigonopselaphus Gemmin- phic; in male, very long and apically ex- ger and Harold, 1868, with several shorter tended into long, curved apophyse. En- setae in Triacrus and only one long seta in tire head, most of pronotum, and apical Elmas and Plociopterus. Mentum with an- portion of abdomen with conspicuous, terolateral setae shifted mediad in Tympan- long, dense golden-yellowish pubescence ophorus. Gula with weak basal impression in ...... Emus Leach, 1819 Glenus. ± Intercoxal process of mesosternum nar- Pronotal epimeron present in Algon, some row, acute to broadly rounded apically Dysanellus Bernhauer, 1911, Gastrisus, and (®gs. 59, 60, 63), middle coxae therefore contiguous or narrowly separated; rarely, Trigonopselaphus, and reduced in Nausico- if middle coxae widely separated (subgen. tus Sharp, 1884, Paraxenopygus Bernhauer, Chaetodracus of Platydracus), then other 1911, Polyphematiana Strand, 1914, Xantho- characters of the couplet agree. At most pygus, and Xenopygus Bernhauer, 1906 (®g. ®rst four antennal segments lacking dense 53). Prosternum weakly carinate in front of appressed pubescence, last six segments sternacostal ridge in some Philothalpus, not forming club. Hind trochanters simple, some Plociopterus, some Styngetus, and not sexually dimorphic. Pubescence of some Tympanophorus. Mesosternum with body different ...... 3 medial carina in Algon, Allostenopsis, Eu- 3. Puncture bearing postocular seta on head gastus, some Gastrisus, and Plociopterus. situated distinctly closer to posterior mar- gin of head than to posterior margin of eye (®g. 102). Head more or less dilated KEY TO NORTH TEMPERATE GENERIC-LEVEL posteriad (very slightly so in P. brevi- TAXA ASSOCIATED WITH STAPHYLINUS cornis). Submentum with anterior margin SENSU LATO beaded throughout. Paramere of aedea- gus somewhat dorsoventrally ¯exible, The following key includes only taxa that lacking sensory peg setae ...... are relevant for the understanding and proper ...... Platydracus Thomson, 1858 assessment of the Staphylinus complex of a. Middle coxal cavities margined pos- genera, and thus the Palearctic taxa of Staph- teriorly ...... ylinina of Eucibdelus lineage (with dilated . . Subgen. Platydracus Thomson, 1858 ± Middle coxal cavities not margined front tibiae), Liusus Sharp, 1889, and Had- posteriorly ...... ropinus Sharp, 1889, and the Nearctic taxa . . . Subgen. Chaetodracus J. MuÈller, 1926 Hadrotes and Thinopinus are not included. ± Puncture bearing postocular seta on head These genera are easily distinguishable from situated distinctly, or at least appreciably, any genus of the Staphylinus complex, and closer to posterior margin of eye than to AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_19 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 19

posterior margin of head (®gs. 103, 104). moderately and rather gradually de¯ected Head not appreciably dilated posteriad ventrad at variable distance between mid- (slightly so in some Ontholestes), with dle of pronotum and anterior angles of posterior corners variably, obtusely pronotum, anterior portion of superior rounded, or if dilated posteriad, then pos- line therefore situated only slightly to terolateral portions of head each extend- moderately below basal portion in lateral ed posteriad as rounded lobe (®g. 105). view (®gs. 113, 114). Puncture bearing Submentum with anterior margin usually ®rst lateral macroseta on pronotum close not beaded throughout. Paramere of ae- to superior line, separated from it by no deagus to great extent immovably at- more than two diameters of puncture tached to median lobe, with or without (®gs. 113, 114) ...... 13 sensory peg setae, or if somewhat dor- 6. Dorsal portion of neck continuous with soventrally ¯exible and without sensory dorsal surface of head due to absence of peg setae, then posterolateral portions of dorsal portion of nuchal ridge (®g. 115). head each extended posteriad as rounded Medial edge of each mandible with only lobe (®g. 105) ...... 4 one simple tooth. Body conspicuously 4. Posterolateral portions of head each ex- elongate, appearing cylindrical ...... tended posteriad as rounded lobe (®g...... Physetops Mannerheim, 1830 105). Eyes almost perfectly round to ± Dorsal portion of neck separated from scarcely oval. Mandibles each markedly dorsal surface of head by nuchal ridge short with very wide basal portion, no (®gs. 40, 116). Medial edge of at least more than twice as long as width at base one mandible with more than one tooth. and usually less so (®gs. 9, 105). Width Body not conspicuously elongate and cy- of mentum less than three times its max- lindrical ...... 7 imum length. Pronotum with lateral mar- 7. Mesosternum with ®ne, short, medial gins abruptly explanate starting a short longitudinal carina at base, or with fully distance behind anterior angles, giving developed, long medial carina (®gs. 117, appearance of an indentation (®g. 106). 118) ...... 8 Paramere of aedeagus somewhat dorso- ± Mesosternum without medial longitudi- ventrally ¯exible, without sensory peg nal carina (®gs. 59, 60) ...... 9 setae ...... Naddia Fauvel, 1867 8. Anterior angles of pronotum extended ± Posterolateral portions of head not ex- angulately (®g. 44). Mesosternum with tended posteriad (®gs. 103, 104), slightly fully developed, long medial carina (®g. so in some Ontholestes, in which eyes are 117) ...... Ontholestes Ganglbauer, 1895 longer than tempora (®g. 107). Eyes oval ± Anterior angles of pronotum obtuse, not or ovate. Mandibles each variably long extended angulately. Mesosternum with with moderately wide basal portion, dis- ®ne, short, medial carina at base (®g. 118) tinctly (more than twice) to considerably ...... Thoracostrongylus Bernhauer, 1915 longer than width at base (®gs. 8, 10, 9. Medial edge of mandibular prostheca co- 108). Width of mentum at least three piously ciliate along entire length, with times its maximum length. Pronotum in extensive basal group of long ciliae and dorsal view with lateral margins not ap- sparser distal group originating on sepa- pearing indented behind anterior angles rate, elongate supporting arm, appearing (®gs. 109, 110, 159). Paramere of aedea- therefore as more or less bilobed, or even gus to great extent immovably attached multilobed (®g. 12) ...... 10 to median lobe, with or without sensory ± Medial edge of mandibular prostheca peg setae ...... 5 variably, usually much less copiously, 5. Superior line of pronotal hypomeron ciliate, without extensive basal group of markedly and rather abruptly de¯ected ciliae originating on separate supporting ventrad, before or at most at middle of arm, distinctly not appearing bilobed, pronotum, anterior portion of superior sometimes narrow and elongate with cil- line therefore situated markedly below iae present only near or at apex (®gs. 13± basal portion in lateral view (®gs. 111, 16) ...... 11 112). Puncture bearing ®rst lateral ma- 10. Long serial setae on mesosternum (in croseta on pronotum distant from supe- more or less transverse or arcuate pat- rior line, separated from it by at least four tern) each originating in enlarged, more diameters of puncture (®gs. 111, 112) . . 6 or less pitlike puncture (®g. 59). De¯ect- ± Superior line of pronotal hypomeron ed portion of pronotum extensively ex- AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_20 Allen Press • DTPro System File # 01cc

20 AMERICAN MUSEUM NOVITATES NO. 3287

posed in ventral view, from at least near visible abdominal tergites simple, straight posterior margin of pronotal epimeron ...... Ascialinus Bernhauer, 1933 (®g. 54). Mentum with anterolateral, se- 13. Vertex of head with slight, obtusely ele- tae-bearing portions each at same level as vated carina, branching anteriorly in form disc of mentum and not separated from of a T. Pronotum distinctly narrowed an- it by obtuse carina (®g. 120). Mandibles teriad, disc with smooth, slightly elevated each without deep subrectangular inden- (gradually more so posteriorly) medial tation, with or without small subbasal line, separating two inde®nite, shallow tooth (®g. 121). Pronotal epimeron impressions in front of posterior margin broadly rounded or absent ...... of pronotum ...... Dinothenarus Thomson, 1858 ...... Apostenolinus Bernhauer, 1934 a. Ligula entire, not bilobed, rarely in- ± Vertex of head and pronotum without cised apically (®g. 38). Mandibles structures described above, although each with small subbasal tooth. Body pronotum often with impunctate median extensively covered with lush, var- strip ...... 14 iegate tomentose pubescence ..... 14. Superior and inferior lines of pronotal . . Subgen. Dinothenarus Thomson, 1858 hypomeron distinctly, widely separated at ± Ligula emarginate or more or less bi- anterior angle, with inferior line disap- lobed (®g. 119). Mandibles each with- pearing along prosternal margin (®g. 46) out small subbasal tooth. Body with- ...... Agelosus Sharp, 1889 out lush, variegate tomentose pubes- ± Superior and inferior lines of pronotal cence, although some variegate pu- hypomeron not widely separated near an- bescence may be rarely present .... terior angle, usually connected before or ..... Subgen. Parabemus Reitter, 1909 at anterior angle of pronotum, although ± Long serial setae on mesosternum (in sometimes running close to each other more or less V-shaped pattern), each anteriorly and not distinctly merging originating behind a small elevation, not (®gs. 48, 50, 52) ...... 15 in an enlarged puncture (®g. 122). De- 15. Mandibles each with only one simple ¯ected portion of pronotum moderately tooth on medial margin, or falciform, exposed near anterior angles in ventral without tooth on medial margin (®gs. 4, view, from about anterior edge of pron- 5). Last segment of maxillary palpus se- otal epimeron (®g. 56). Mentum with an- terolateral, setae-bearing portions each tose (®g. 29) ..... Tasgius Stephens, 1829 obliquely de¯ected at an angle and there- a. Mandibles each with one simple tooth fore not at same level as disc of mentum, on medial margin (®g. 5) ...... separated from it by obtuse carina (®g...... Subgen. Tasgius Stephens, 1829 123). Mandibles each with small subbas- ± Mandibles each simple, falciform, al tooth in deep subrectangular emargi- without tooth on medial margin (®g. 4) nation (®g. 124). Pronotal epimeron well . . Subgen. Rayacheila Motschulsky, 1845 developed, projecting triangularly ..... ± Mandibles each, or at least left mandible, ...... Abemus Mulsant and Rey, 1876 with two or more teeth on medial margin 11. Middle coxal cavities not margined pos- (®gs. 127, 128); if each mandible with teromedially (®g. 64). Body with isolated only one simple tooth, then last segment patches of yellow tomentose pubescence of maxillary palpus asetose (®g. 129) 16 on head and usually also on pronotum 16. Mandibles each with only one simple ...... Staphylinus LinneÂ, 1758 tooth on medial margin ...... 17 ± Middle coxal cavities margined postero- ± Mandibles each, or at least left mandible, medially (®gs. 62, 63). Body without iso- with two or more teeth on medial margin lated patches of yellow tomentose pubes- (®gs. 127, 128) ...... 18 cence on head and pronotum ...... 12 17. Posterior basal line on ®rst three visible ab- 12. Head with small, round, shallow, smooth, dominal tergites (best visible on tergite II) pitlike depressions among usual punctation obtusely extended posteriad on each lateral (®g. 125). Posterior basal line on ®rst three portion, base of tergite between extensions visible abdominal tergites gently bisinuate depressed, depression delimited laterally by (®g. 126) ...... Miobdelus Sharp, 1889 short oblique carina situated at middle of ± Head with usual punctation, without shal- each lateral extension of basal line. Last seg- low, smooth pitlike depressions (®g. ment of labial palpus short, truncate apical- 103). Posterior basal line on ®rst three ly, either more or less parallel-sided or var- AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_21 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 21

iably dilated apicad. Dorsal portion of ae- anteriad, widely truncate, securiform . . . deagus variably sclerotized . . . Ocypus Leach, ...... Wasmannellus Bernhauer, 1920 1819, pars (Aulacocypus J. MuÈller, 1925) ± Posterior basal line on ®rst three visible ab- THE GENERA dominal tergites simple, not extended pos- teriad on each lateral portion, bases of ter- The following treatment of each taxon lists gites without short oblique carina. Last seg- the type species and, if required, discusses ment of labial palpus distinctly fusiform. taxonomic and other information important Dorsal portion of aedeagus almost entirely for justi®cation and understanding of the new membranose ...... concepts presented. No full formal descrip- ...... Protogoerius Coiffait, 1956 tions and detailed information about the type 18. Bases of ®rst four visible abdominal tergites species are given, since they do not ®t in the not appreciably transversely impressed. intended scope of this paper. Hind tibia with spines on dorsolateral face (®g. 130). Last segment of labial palpus of variable shape, in general fusiform, or more Creophilus Leach, 1819 or less parallel-sided and truncate apically TYPE SPECIES: Staphylinus maxillosus Lin- (®gs. 131, 132) ...... neÂ, 1758...... Ocypus Leach, 1819 COMMENTS: Members of this genus may be a. Posterior basal line on ®rst three vis- ible abdominal tergites (best visible easily recognised by the following shared on tergite 2) obtusely extended pos- character states: (1) palpifer with one long teriad on each lateral portion, base of subapical seta and three to four shorter apical tergite between extensions de- setae; (2) disc of pronotum and most of dor- pressed, with depression delimited sal face of pronotum virtually impunctate laterally by short oblique carina sit- and asetose; (3) middle coxae widely sepa- uated at middle of each lateral exten- rated by broadly rounded apical margin of sion of basal line ...... mesosternum; (4) pronotal hypomeron with .... Subgen. Aulacocypus J. MuÈller, 1925 superior line becoming obsolete near anterior ± Posterior basal line on ®rst three vis- angles of pronotum and not joining the in- ible abdominal tergites simple, not ferior line (®g. 49); (5) presence, on both extended posteriad on each lateral dorsal and ventral faces of body, of long, portion ...... b more-or-less variegate pubescence varying in b. Last segment of maxillary palpus se- color from black to grayish-silver, grayish- tose (®g. 25) ...... Subgen. Matidus Motschulsky, 1860 golden, yellowish-red, or brownish-red (such ± Last segment of maxillary palpus as- pubescence not present in a few species oc- etose (®g. 133) ...... c curring elsewhere); (6) anterior projection of c. Last segment of both maxillary and metasternum very broad, with margins ob- labial palpus elongate, longer than solete; and (7) dorsal apicolateral lobe of wide, in general fusiform, narrowly hind coxa without distinct spines, or spines subtruncate apically (®gs. 23, 131). very ®ne. Palpifer usually with two apical setae Further character states include: mandi- (®g. 128) ...... Subgen. Pseudocypus bles each with microsculpture on mediobasal Mulsant and Rey, 1876 portion; mandibular prostheca copiously cil- ± Last segment of at least labial palpus iate along entire length, with long, dense bas- short, no more than moderately lon- al ciliae gradually shortening apicad; post- ger than wide, broadly truncate api- mandibular ridge absent; de¯ected portion of cally (®gs. 132, 133). Palpifer usu- pronotal disc visible in ventral view from ally with three apical setae (as in ®g. 149) ..... Subgen. Ocypus Leach, 1819 about anterior edge of pronotal epimeron, ± Bases of ®rst four visible abdominal ter- which is well developed and membranous; gites markedly transversely impressed metasternum oblique, highly convex between and with punctation different from that coxae, then horizontal. on rest of tergites. Hind tibia without This genus is assigned by some authors spines on dorsolateral face. Last segment (e.g., Coiffait, 1974: 557; Moore and Legner, of labial palpus short, distinctly dilated 1979: 32) to the subtribe Xanthopygina AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_22 Allen Press • DTPro System File # 01cc

22 AMERICAN MUSEUM NOVITATES NO. 3287

based on the con®guration of the superior maximum length; (5) lateral margins of pro- and inferior lines of the pronotal hypomeron notum abruptly explanate a short distance be- (see above). Although Creophilus (with Had- hind anterior angles, giving appearance of an rotes and Thinopinus) seems to be interme- indentation (®g. 106); and (6) paramere of diate between the two subtribes, characters aedeagus somewhat dorsoventrally ¯exible, 1, 6, and 7 above are shared exclusively with without sensory peg setae. members of the Staphylinina; together with Further character states include: outer an- the absence of the postmandibular ridge, tennal segments slightly asymmetrical to these con®rm their closer af®nity with the subserrate; mentum with cluster of variable Staphylinina. anterolateral setae; maxillary palpus with apical segment stoutly ¯attened, with last Emus Leach, 1819 two broadening to apex; last segment of la- bial palpus short, fusiform, setose; mandib- TYPE SPECIES: Staphylinus hirtus LinneÂ, ular prostheca lanceolate, with short ciliae 1758. along medial margin; postmandibular ridge COMMENTS: Species of this genus are quite rudimentary; postgenal ridge present only distinctive by the long, matted, black, gold- laterally; superior line of pronotal hypome- en-yellow, and whitish-gray pubescence of ron moderately de¯exed ventrad between the body, and by the characters used in the middle of pronotum and anterior angles of key to genera. pronotum, with anterior portion of superior Further character states include: mandible line therefore situated only moderately below with small rectangular tooth basad of large basal portion in lateral view; de¯ected por- middle tooth; mandibular prostheca long, tion of pronotal disc visible in ventral view parallel-sided, with ciliae only near apex; only from edge of coxal insertion; hypome- postmandibular ridge rudimentary; palpifer ron very broad at middle; proepimeron well with several apical and many discal, coarse developed, membranous; series of long setae setae; mentum with arcuate carina between on mesosternum situated on small tubercles basal angles, separating ellipsoidal membra- in a very broadly U-shaped pattern (shared nous basal portion from small anterolateral with members of Platydracus); mesosternal sclerotized portions, which are de¯ected an- intercoxal process with margins slightly con- terolaterad and bear a cluster of setae; su- cave, narrowly rounded apically; metaster- perior and inferior lines of pronotal hypo- num oblique, highly convex between coxae, meron connected just behind posterolateral horizontal posteriorly; dorsal apicolateral angle of prosternum; de¯ected portion of lobe of metacoxa with one short, ®ne spine. pronotal disc visible in ventral view from near posterior margin of proepimeron, which is well developed and membranous; sterna- Ontholestes Ganglbauer, 1895 costal ridge bisinuate, transverse, roughly TYPE SPECIES: Staphylinus murinus LinneÂ, parallel to anterior prosternal margin; ante- 1758. rior projection of metasternum very broad, COMMENTS: The species of this genus margins obsolete. share four distinctive synapomorphies: (1) anterior angles of pronotum angulately ex- Naddia Fauvel, 1867 tended anteriad (®g. 44); (2) marginal line of TYPE SPECIES: Caranistes westermanni Er- pronotal hypomeron narrowly rounded at ichson, 1840. prosternal margin (®g. 44); (3) mesosternum COMMENTS: Members of this genus share with complete, medial longitudinal carina the following character states: (1) posterolat- (®g. 117); and (4) mentum with long antero- eral portions of head extended posteriad as lateral seta near lateral margin, lateral seta rounded lobes (®g. 105); (2) eyes distinctly minute to absent. shorter than tempora, almost perfectly round Further character states include: outer seg- to scarcely oval; (3) mandibles short and ments of antenna variably asymmetrical; wide basally (®g. 9) (see key for details); (4) maxillary and labial palpus both with last width of mentum less than three times its segment asetose, fusiform; mandibles each AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_23 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 23

with dorsolateral ridge distinct, removed fore situated markedly below basal portion in from margin, bordered by deep, broad de- lateral view; de¯ected portion of pronotal pression basally; mandibular prostheca co- disc visible from distinctly behind epimeron piously ciliate along entire length, with long, in ventral aspect; proepimeron poorly devel- irregular, dense basal ciliae gradually short- oped, very membranous; prosternum very ening apicad; apical segments of both max- long (length about three-fourths width), with illary and labial palpus distinctly elongate fu- distinct, straight carina from behind macro- siform, without setae; postmandibular ridge setae, continuing on furcasternum; serial present; postgenal ridge arched to base of long setae on mesosternum arranged in gula; superior line of pronotal hypomeron broadly U-shaped pattern behind an elevated rather abruptly de¯exed ventrad around mid- arcuate ridge behind median carina; meso- dle of pronotum, with anterior portion of su- sternal intercoxal process very narrowly perior line therefore situated markedly below rounded apically; metasternum as in Ontho- basal portion in lateral view; pronotal epi- lestes; dorsal apicolateral lobe of metacoxa meron long, subtriangular; prosternum high- with one short, ®ne spine located distinctly ly prominent, sloping abruptly to anterior subapically; scutellum very long, almost ex- margin, with strong, convex carina starting tending halfway to apex of elytra. between macrosetae; series of long setae on We were able to study the holotype of Pa- mesosternum arranged in subtransverse row rontholestes nepalicus, deposited in the col- on each side of median carina; mesosternal lection of the Forschungsinstitut Sencken- intercoxal process angulate to broadly trun- berg (SMF C 15051), Frankfurt a. M., Ger- cate apically; metasternum oblique, highly many. It has all the characters of the genus convex between coxae, anterior projection Thoracostrongylus, including the presence of with acutely converging margins (®g. 117); a ®ne short carina at the base of mesoster- dorsal apicolateral lobe of metacoxa without num, despite the statement ``MeÂsosternum apparent spines. non careÂneÂ'' in the original description (Coif- fait, 1982: 72). The name Parontholestes Thoracostrongylus Bernhauer, 1915 therefore becomes a junior synonym of Thoracostrongylus. TYPE SPECIES: Ontholestes javanus Bern- hauer, 1915. We also studied the holotype of Thoraco- strongylus martensi Coiffait, 1982, deposited Parontholestes Coiffait, 1982 (syn. nov.) (type in the same institution (SMF C 15143). The species: Parontholestes nepalicus Coiffait, holotype cannot be distinguished from spec- 1982). imens of Abemus olivaceus Cameron, 1928. COMMENTS: Species of this genus share the The name T. martensi must therefore be synapomorphy of a short, medial carina at transferred to Abemus, where it becomes a base of mesosternum (®g. 118). They also junior synonym of A. olivaceus. share a submentum with apical margin bead- ed. Abemus Mulsant and Rey, 1876 Further character states include: eyes large, prominent, tempora short, evenly TYPE SPECIES: Staphylinus chloropterus rounded toward narrow neck, which is little Panzer, 1796. over one-third total width of head; maxillary COMMENTS: Members of this genus share and labial palpus both with last segment as- the following character states: (1) mandibles etose, fusiform; mandibular prostheca short, with similarly developed teeth, each with copiously ciliate along entire length, with large tridentate middle tooth (individual long, dense basal ciliae gradually shortening dents not at same plane) and a small tooth apicad; mentum with two lateral setae; post- basad of it in a deep, subrectangular inden- mandibular ridge present; postgenal ridge tation (®g. 124); (2) mandibular prostheca bent toward gula from lateral bend; superior appearing bilobed or even multilobed, with line of pronotal hypomeron rather abruptly extensive basal group, and long distal group de¯exed ventrad around middle of pronotum, of ciliae originating on separate supporting with anterior portion of superior line there- arm (®g. 12); (3) mentum with anterolateral AMNH NOVITATES Tuesday Dec 11 2001 10:10 AM 2000 novi 99163 Mp_24 Allen Press • DTPro System File # 01cc

24 AMERICAN MUSEUM NOVITATES NO. 3287

portions (bearing two anterolateral setae) with apical margin beaded throughout; pos- each obliquely de¯ected at an angle and teroventral margin of eye distinctly oblique, therefore not at same level as disc of men- postmandibular ridge rapidly diverging from tum, separated from it by obtuse carina (®g. it and bordered by even series of setae ba- 123); and (4) postmandibular ridge almost sally, reaching about halfway or more to sub- reaching infraorbital puncture, running close ocular puncture, which is separated from to eye basally, infraorbital puncture situated margin of eye by at least four, and usually three to four diameters from margin of eye. more than eight diameters of puncture; nu- Further character states include: maxillary chal ridge complete, directed toward trans- and labial palpi both with last segment ase- verse postgenal ridge laterally, and only tose, fusiform; mandibles each with dorso- moderately removed from nuchal constric- lateral ridge as in Ontholestes or Thoraco- tion; pronotal hypomeron markedly broader strongylus; submentum with anterior margin at middle, with superior line distinctly round- not beaded; lateral end of nuchal ridge dis- ed anteriorly, joining inferior line well be- tinctly, but moderately, removed from nuchal hind posterolateral angle of prosternum, de- constriction, directed toward level of trans- ¯ected portion of pronotal disc visible from verse postgenal ridge; epimeron of prothorax at, or behind, posterior edge of coxal inser- well developed, projecting triangularly (®g. tion (®g. 136); pronotal epimeron well de- 111); prosternum moderately prominent with veloped, triangular (®g. 52); prosternum broadly rounded slope anteriad, with promi- sloping moderately, never abruptly in front of nent, arcuate carina continuing evenly onto macrosetae, sternacostal ridge almost straight, furcasternum; mesosternum with long serial slightly arched posteriad; mesosternum with- setae originating behind small elevations in out ridges on intercoxal projection, with series more-or-less V-shaped pattern (®g. 122), me- of macrosetae on small elevations in broadly sosternal intercoxal process rather acute api- U-shaped pattern; mesocoxal acetabulum cally, mesocoxal acetabulum moderately im- weakly to moderately impressed behind, nev- pressed behind; metasternum more or less er with medial carinae extending from projec- horizontal, with little difference in convexity tion; anterior metasternal projection with mar- between projections and disc. gins complete (except in Chaetodracus), con- verging less than rectangularly; metasternum Platydracus Thomson, 1858 more or less horizontal, with little difference in convexity between anterior projection and TYPE SPECIES: Staphylinus stercorarius disc; dorsal apicolateral lobe of metacoxae Olivier, 1795. with two to seven coarse spines. COMMENTS: Members of this genus share the following character states: (1) puncture Subgenus Platydracus Thomson, 1858. bearing postocular seta on head situated dis- Neotasgius J. MuÈller, 1925 (syn. nov.) (type spe- tinctly closer to posterior margin of head cies: Ocypus brevicornis Weise, 1877; nec Mot- than to posterior margin of eye (®g. 102); schulsky, 1862 ϭ P. brachycerus, nom. nov.). and (2) paramere of aedeagus devoid of any sensory peg setae and attached to median COMMENTS: Members of this subgenus lobe in such a way that it is somewhat dor- share the characters of posteromedially mar- soventrally ¯exible. gined middle coxal cavities (®g. 137) and of Further character states include: mandib- the narrow intercoxal mesosternal process, ular teeth (on both mandibles) appearing in no more than moderately widely separating two planes, dorsal and ventral, in most spe- the middle coxae. cies (®g. 7); mandibular prostheca with dense Neotasgius was established by J. MuÈller patches of ciliae, ciliae long at base, gradu- (1925: 41) as a subgenus of Staphylinus. ally shortening apicad, with prostheca there- However, the type species of Neotasgius in fore appearing as more or less bilobed (®g. fact shows all typical character states of Pla- 11); maxillary and labial palpi with apical tydracus, although its narrow body shape, segments moderately elongate, fusiform, with the narrow head not much dilated pos- without setae (®gs. 134, 135); submentum teriad, renders it an unusual habitus. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_25 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 25

Subgenus Chaetodracus J. MuÈller, 1926 (type study of Mannerheim's original material. We species: Staphylinus patricius Bernhauer, 1915). therefore continue to use the longstanding Nordmann name and consider Staphylinus COMMENTS: The type species of this sub- genus is distinctive not only by the postero- immaculatus a doubtful name. medially margined middle coxal cavities (a 5. Platydracus yunnanensis (Bernhauer, character shared with the members of Staph- 1943: 76), described in Staphylinus, is pre- ylinus, see ®g. 64), but also by the wide in- occupied by Platydracus yunnanensis (Bern- tercoxal process of the mesosternum that hauer, 1933: 48), also described in Staphyli- very widely separates the middle coxae, sim- nus. A replacement name, Platydracus yun- ilar to that in Emus. Additional species be- nanicus, nom. nov., is proposed for Platyd- longing to this subgenus may ultimately be racus yunnanensis (Bernhauer, 1943). found among the east Palearctic species of Platydracus. Staphylinus LinneÂ, 1758 (stat. nov.)

TYPE SPECIES: Staphylinus erythropterus REMARKS ON SOME SPECIES OF PLATYDRACUS LinneÂ, 1758. 1. Platydracus brachypterus (Kraatz, COMMENTS: All members of this genus 1859: 76), described in Staphylinus, is pre- share the synapomorphy of the middle coxal occupied by Staphylinus brachypterus Geof- cavities not margined posteromedially (®g. froy, 1785: 167, now a synonym of Aleo- 64). In addition, they share (1) superior line chara curtula (Goeze, 1777: 730); by Staph- of pronotal hypomeron markedly and rela- ylinus brachypterus Marsham, 1802: 510, tively abruptly de¯exed ventrad, before or at now a synonym of Aploderus caelatus most at middle of pronotum, with its anterior (Gravenhorst, 1802); and by Staphylinus bra- portion situated markedly below basal por- chypterus BrulleÂ, 1839: 59, now in Proto- tion in lateral view (®g. 112); (2) bicolored goerius. A replacement name, Platydracus black body with red elytra and legs red ex- brevipennis, nom. nov., is hereby proposed cluding coxae; and (3) presence of yellow for Platydracus brachypterus (Kraatz). tomentose pubescence on head and usually 2. Platydracus oculatus (Bernhauer, 1929: also on pronotum, and paired yellow tomen- 110), described in Staphylinus, is preoccu- tose patches on abdominal tergites. pied by Staphylinus oculatus Fabricius, Further character states include: right man- 1775: 265, now a valid name in Creophilus; dible with one stout, apically truncate, sub- and by Staphylinus oculatus O. F. MuÈller, emarginate or slightly emarginate tooth, left 1776: 99. A replacement name, Platydracus mandible with two dorsal teeth, and with one oculosus, nom. nov., is hereby proposed for somewhat smaller tooth in more ventral Platydracus oculatus (Bernhauer). plane (®g. 138); mandibular prostheca nar- 3. Platydracus violaceus (Gravenhorst, row, elongate, with ciliae present only at 1802: 162), described in Staphylinus, is pre- apex (®g. 15); last segment of maxillary pal- occupied by Staphylinus violaceus Olivier, pus fusiform, narrower and markedly shorter 1795: 8 (now in Plochionocerus). The name than segment three, asetose (®g. 24); ligula Platydracus cupripennis (Melsheimer, 1844: emarginate; last segment of labial palpus fu- 35) becomes the valid name for this species siform, narrower and markedly longer than (stat. nov.). segment two, asetose (®g. 139); anterior mar- 4. Platydracus vulpinus (Nordmann, 1837: gin of submentum inwardly arcuate, without 53) has an older synonym, Platydracus im- bead; postmandibular ridge rudimentary to maculatus (Mannerheim, 1830: 22). Both very short; base of infraorbital ridge on the species were described in Staphylinus. Erich- neck distinctly recurved toward nuchal ridge son (1839: 379) treated Staphylinus imma- dorsally, sometimes reaching it; superior and culatus tentatively as identical with S. vul- inferior lines of pronotal hypomeron con- pinus, but all subsequent authors simply list- nected well before anterior angles of prono- ed Mannerheim's name as a synonym, de- tum (®g. 57); epimeron of prothorax absent spite its seniority. The status of these two (®g. 50); prosternum with middle portion of species can only be established after the basisternite broadly rounded or gradually AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_26 Allen Press • DTPro System File # 01cc

26 AMERICAN MUSEUM NOVITATES NO. 3287

sloping anteriad; serial long setae on meso- terior margin of epimeron (especially in sternum originating in moderately impressed members of Parabemus) to near the posterior punctures in subtransverse to broadly arcuate pronotal angles (especially in Dinothenarus arrangement; spines on dorsal apicolateral [Dinothenarus]); pronotal epimeron ranging lobe of hind coxae small, almost indistin- from moderately developed to virtually ab- guishable in size from other setae. sent; dorsal apicolateral lobe of hind coxa of- This is a new concept of this genus, re- ten with more than three (up to six) distinctly stricted to the species around S. erythropte- spinelike coarser setae. rus and S. caesareus. Six Palearctic and one Nearctic species are at present included in Subgenus Dinothenarus Thomson, 1858. the genus. Protabemus Scheerpeltz, 1966 (type species: Staphylinus xanthocephalus Kraatz, 1859).

Dinothenarus Thomson, 1858 (stat. nov.) COMMENTS: Members of this subgenus share the following character states: body ex- TYPE SPECIES: Staphylinus pubescens tensively covered with lush, variegated pu- DeGeer, 1774. bescence of tomentose character (®gs. 140, COMMENTS: Members of this genus share 141); at least mandibles and adjacent areas, the following character states: (1) mandibular and usually large portions of ventral surface prostheca bilobed, with basal group of long of head bicolored, with distinct testaceous to ciliae and sparser distal group originating on rufotestaceous areas; ligula virtually entire separate supporting arm; and (2) long serial (®g. 38), rarely incised apically (e.g., in D. setae on mesosternum situated on anterior xanthocephalus); anteromedial portion of edges of enlarged, more-or-less pitlike punc- prosternum highly prominent, forming a tures, in subtransverse to broadly U-shaped rounded knob which slopes abruptly anteriad pattern (®g. 59). and markedly delimits the transverse margin- Further character states include: right man- al depression; sternacostal ridge distinctly dible with one broad tooth at middle, left arched posteriad toward keel, usually coming mandible with two teeth or with one stout, close to posterior margin of furcasternum apically emarginate tooth at middle, with all (®g. 54); mesosternal intercoxal process with teeth in same (dorsal) plane; maxillary pal- margins more or less straight, converging pus with apical segment moderately elon- subrectangularly until just before rectangular gate, distinctly fusiform, narrower and sub- apex, margins without bead apically, slightly equal in length to segment three, asetose explanate; mesocoxal acetabulum only (®gs. 23, 28); apical segment of labial palpus slightly depressed behind projection; margins distinctly fusiform, narrower and markedly of anterior metasternal projection never con- longer than segment three, usually with a verge at less than right angle, although apex few, sometimes with many, ®ne setae (®g. varies from moderately to broadly rounded; 38); mentum usually with semicircular ®ne disc of metasternum always horizontal, on basal carina extending variably between ba- same plane as body. solateral angles (®gs. 119, 120); postmandib- The subgenus contains six Palearctic and ular ridge rudimentary to extended less than one Nearctic species at present. halfway to infraorbital puncture, infraorbital puncture from 4 to 10 diameters distant from Subgenus Parabemus Reitter, 1909 (stat. nov.) margin of eye; postgenal ridge usually broad- (type species: Staphylinus fossor Scopoli, ly obsolete and often de¯ected obliquely an- 1772). teriad toward gula, especially in members of Parocypus Bernhauer, 1915 (syn. nov.) (type spe- Parabemus; superior line of pronotal hypo- cies: Staphylinus dehradunensis Bernhauer, meron markedly and relatively abruptly de- 1915). ¯ected ventrad, before or at most at middle Hypabemus Scheerpeltz, 1966 (syn. nov.) (type of pronotum, with its anterior portion situ- species: Staphylinus chrysocomus Mannerheim, 1830). ated markedly below basal portion in lateral view (®g. 111), de¯ected portion of pronotal COMMENTS: Members of this subgenus disc visible in ventral view from behind pos- share the following character states: body AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_27 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 27

with patches of variegated pubescence; ven- ``China Prov. Fo-Kien G. Siemssen.'' We tral surface of head piceous to dark rufous, con®rm that Staphylinus fokiensis is a mem- without distinctly bicolored areas; ligula ber of the subgenus Parabemus of Dinoth- emarginate or bilobed (®g. 119), anterome- enarus. The above two specimens were com- dial portion of prosternum usually only mod- pared to the lectotype of Dinothenarus insig- erately prominent, with distinct, but broadly nis (J. MuÈller, 1926) (see below) and were rounded, slope anteriad; sternacostal ridge found to be conspeci®c. The name D. fokien- slightly oblique, relatively straight to mod- sis therefore becomes a junior synonym of erately arched posteriad toward keel; meso- D. insignis (syn. nov.). sternal intercoxal process with margins con- 2. Dinothenarus fossor (Scopoli, 1772: cave, converging distinctly less than rectan- 109) (Staphylinus). Fabricius (1792: 523) de- gularly, with apex acute to narrowly rounded, scribed Staphylinus fossor based on speci- margins distinctly beaded, at least near apex men(s) of unknown provenience. Erichson (®g. 142); mesocoxal acetabulum slightly de- (1839a: 377) cited S. fossor Fabricius as a pressed to deeply depressed behind projec- subsequent reference of Staphylinus fossor tion, with concavity below apex; some spe- Scopoli, 1772. However, Fabricius (loc. cit.) cies with metasternum short and at an redescribed the species, using the same name oblique angle to plane of body. S. fossor as had Scopoli (loc. cit.) before The type species of both Parocypus and him. The name S. fossor Fabricius, 1792 Hypabemus share the character states of Par- therefore becomes a junior synonym of abemus, as they were presented above, and Staphylinus fossor Scopoli, 1772 (syn. nov.). therefore there is no justi®cation for keeping 3. Dinothenarus ganglbauerianus (Bern- them as separate subgenera. hauer, 1938: 101) (Staphylinus). Bernhauer This is a new, expanded concept of this (loc. cit.) described this species as a member subgenus that includes 12 east Palearctic/ of the subgenus Xanthocypus of Staphylinus. Oriental species previously assigned to Par- Recently, Hayashi (1995: 45) studied the ho- ocypus, as well as 5 Nearctic species that lotype of this species from Kamikochi, Japan were previously listed as members of the ge- (see Hayashi, loc. cit., for the detailed label nus ``Staphylinus.'' One of the Nearctic spe- data), and redescribed it. He considered the cies (D. pleuralis) was assigned by Smetana species to be a member of the genus Ocypus, (1965a: 10) to Abemus (as a subgenus of very closely related to ``Ocypus similis,'' but Staphylinus at that time), but Frank (1979: at the same time he pointed out some unique 236) assigned it correctly to Parabemus, characteristics of the species. We were able based on larval characters. Additional Pale- to study the holotype and concluded, based arctic and Oriental species will likely be add- mainly on thoracic structures, on the shape ed to this subgenus in the future. of the mandibular prostheca, and on the con- ®guration of the aedeagus, that it belongs to REMARKS ON SOME SPECIES OF the subgenus Parabemus of the genus Din- DINOTHENARUS othenarus and is related to D. insignis. Din- othenarus insignis occurs in southern China 1. Dinothenarus fokiensis (Bernhauer, (Fujian) and supposedly also in Japan (see 1933: 32) (Staphylinus). Bernhauer (loc. cit.) below). However, D. insignis, which is a described the species as a member of the large and conspicuous species, was appar- subgenus Parocypus of Staphylinus and ently never rediscovered in Japan (see Shi- compared it to ``Staph. dehradunensis.'' bata, 1984: 88), which may indicate that one Since Parocypus becomes a junior synonym of MuÈller's original specimens may have of Parabemus (see above), Bernhauer's as- been mislabelled, and that the species does signment of this species was quite correct. not really occur in Japan. The same actually We were able to study the holotype from applies also to D. ganglbauerianus (see Shi- ``Foo-chow'' in the Bernhauer collection in bata, 1984: 94); we therefore also believe Chicago and another male specimen, com- that the holotype of D. ganglbauerianus is pared with the holotype, from the Scheer- likely mislabelled and that this species in fact peltz collection, Wien, labelled as follows: does not belong to the Japanese fauna. This AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_28 Allen Press • DTPro System File # 01cc

28 AMERICAN MUSEUM NOVITATES NO. 3287

would con®rm the dif®culties Hayashi cal setae; apical segments of maxillary and (1995: 45) expressed, based on different labial palpi moderately elongate, distinctly grounds, about the reliability of the type lo- fusiform, with only that of labial palpus se- cality of D. ganglbauerianus. The aedeagi of tose (®gs. 144, 145); postmandibular ridge D. insignis and D. ganglbauerianus resemble absent; nuchal ridge present only dorsally, in general con®guration those of some mem- becoming obsolete well before base of infra- bers of the genus Ocypus, which contributed orbital ridge laterally; dorsal disc of neck to Hayashi's (1995: 56) conclusion that D. with umbilicate punctures without visible in- ganglbauerianus was closely related to terspaces; pronotal epimeron small; long se- ``Ocypus similis'' (see above). However, we rial setae on mesosternum originating on believe that this similarity in the con®gura- edge of feeble, irregular, very broadly U- tion of the aedeagus is convergent. shaped ridge; mesosternal intercoxal process 4. Dinothenarus insignis (J. MuÈller, acute apically, margin ®nely beaded through- 1926b: 41) (Staphylinus). MuÈller (loc. cit.) out (®g. 60); acetabulum of middle coxae described this species as a member of the deeply depressed, nearly vertical behind pro- subgenus Parocypus of the genus Staphyli- jection; anterior metasternal projection with nus, based on two specimens: one from margins converging quite obtusely, apex ``China merid.'' and one from ``Japonia.'' very broadly rounded. The specimen from China is apparently de- The genus at present contains only one posited in the Zoologisches Museum der species, M. brevipennis from Japan. How- Humboldt-UniversitaÈt, Berlin (not seen), the ever, the genus in fact includes many unde- other one in the Natural History Museum, scribed species in Taiwan and in mainland London. We were able to study the latter China, all of which live mostly at higher specimen (female), which is labelled as fol- mountain elevations. A review of these spe- lows: ``Ocypus n. sp./Type [round label with cies is under preparation by the senior author. red margin]/Typus/Japan. G. Lewis. 1910- 320/St. (Parocypus) insignis n. sp. Det. J. Physetops Mannerheim, 1830 MuÈller.'' The specimen is in fair shape (the right hind tibia and tarsus are missing, and TYPE SPECIES: Staphylinus tataricus Pallas, most of the pubescence of the dorsal side of 1773. the body is gone, but the remnants permit COMMENTS: Members of this genus share reconstruction of its original pattern). The the following character states: (1) dorsal por- specimen is hereby designated as the lecto- tion of neck continuous with dorsal surface type of D. insignis; the label ``LECTOTYPE of head, due to absence of dorsal portion of Staphylinus insignis J. MuÈller A. Smetana nuchal ridge (®g. 115); (2) disc of head and des. 1999'' was attached to it. pronotum micropunctulate between larger punctures (®g. 115); and (3) middle abdom- Miobdelus Sharp, 1889 inal tergites notably swollen, especially in apical half, as convex as sternites, together TYPE SPECIES: Miobdelus brevipennis with convexity of pronotum and length of Sharp, 1889. elytra and body, giving body an overall cy- COMMENTS: Members of this genus share lindrical appearance. the following character states: (1) head with Further character states include: mandibles macrosetae and some of other setae originat- long, each with one blunt tooth on medial ing in punctures expanded into round, shal- edge, dorsolateral ridge weak and relatively low, smooth depressions (®g. 125); (2) man- close to margin; prostheca long, thin, lance- dibular prostheca lanceolate, with moderately olate, with short ciliae along medial margin, long setae along entire medial margin (®g. becoming progressively longer to moderate 143); and (3) posterior basal line on ®rst near apex; palpifer with one long subapical three visible abdominal tergites gently bi- and two shorter apical setae; apical segment sinuate (®g. 126). of both labial and maxillary palpus setose; Further character states include: palpifer apex of last segment of labial palpus oblique- with one long subapical and two shorter api- ly widened; submentum broadly inwardly ar- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_29 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 29

cuate, without bead; postmandibular ridge long, lanceolate, with short ciliae along en- rudimentary; gular sutures most approximate tire medial edge, gradually longer to mod- distinctly anteriad of middle, gradually wide- erate at apex; apical segment of maxillary ly diverging posteriad; postgenal ridge dis- palpus asetose, with apex slightly wider than tinct only laterally; lateral portion of nuchal width at middle, apical segment of labial pal- ridge directed anteriad toward transverse pus with many ®ne setae, apex expanded to postgenal ridge; superior line of pronotal hy- about two times width at base, slightly pomeron markedly and relatively abruptly oblique; postmandibular ridge absent; post- de¯ected ventrad near middle of pronotum, genal ridge highly oblique, reaching gular de¯ected portion of pronotal disc visible suture, very broadly rounded laterally; su- from well behind broadest portion of hypo- perior and inferior lines of pronotal hypo- meron in ventral view; superior and inferior meron run closely anteriad for some distance, lines of pronotal hypomeron almost coalesce merge along prosternal margin; prothoracic at margin of coxal cavity, then run very epimeron moderately developed; prosternum closely parallel along entire lateral margin of long, more than half as long as maximum prosternum; pronotal epimeron absent; pro- width; sternacostal ridge medially extended sternal basisternite sloping very gradually anteriad into short, acute, slightly elevated from anterior margin almost to sternacostal process; elytra short, broadly depressed ridge medially, furcasternum abruptly decli- along base; anterior metasternal projection vous, forming a sharp toothlike process me- with margins converging obtusely, very dially at sternacostal ridge; intercoxal de- broadly rounded apically; metasternum very pression relatively long, shallow; long serial short, at distinctly oblique angle to plane of setae on mesosternum originating in large, body, not visibly canaliculate medially; dor- rounded depressions forming an irregular U- sal apicolateral lobe of hind coxa with very shaped ridge; apex of mesosternal intercoxal ®ne spinelike setae, almost indistinguishable process acute, forming a blunt toothlike pro- from other setae; empodial setae very long. cess, with a rounded carina extending into The species seems to be close to some the moderately depressed mesocoxal acetab- members of the subgenus Matidus of Ocy- ulum behind; anterior metasternal projection pus. Ascialinus may actually contain more with margins converging highly obtusely, species in the east Palearctic area, particular- very broadly rounded apically; dorsal api- ly in mainland China; the ®nely setose pro- colateral lobe of metacoxa usually with one notal hypomeron may be an autapomorphy distinctly subapical, coarse spine, and several of A. beckeri. ®ner ones in an uneven series basad to it. Apostenolinus Bernhauer, 1934 Ascialinus Bernhauer, 1933 TYPE SPECIES: Apostenolinus cariniceps TYPE SPECIES: Ascialinus beckeri Bern- Bernhauer, 1934. hauer, 1933. COMMENTS: In addition to its large size (23 COMMENTS: The only species of this genus mm), the single species of this genus may be is characterized by the following character easily recognized by the following character states: mandibles each similarly shaped, with states: (1) head, pronotum, and elytra dark deep subbasal indentation and two similar bluish; (2) outer four segments of antenna short, angular teeth at middle; base of gula whitish-yellow; and (3) pronotum markedly highly swollen, transverse impression very narrowed anteriad toward narrow head, ®ne, along margin; pronotal hypomeron ®ne- which is distinctly attenuated posteriad. ly setose (but see below); mesosternal inter- Further character states include: mandibles coxal process with margins obsolete, disc en- rather long and slender, with acute apices, tirely, irregularly tumid, with angulate ®nlike medial margin of each with only one tooth, process at apex, extended behind as a long tooth on left mandible short, broadly trun- straight carina into deeply depressed acetab- cate, tooth on right mandible broadly trian- ulum. gular, rather acute; eyes slightly shorter (not Further character states include: prostheca longer, as stated in the original description) AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_30 Allen Press • DTPro System File # 01cc

30 AMERICAN MUSEUM NOVITATES NO. 3287

than tempora (ratio 0.86); puncture bearing Protogoerius Coiffait, 1956 postgenal macroseta situated much closer to posterior margin of eye than to posterior TYPE SPECIES: Staphylinus brachypterus margin of head; superior line of pronotal hy- BrulleÂ, 1839 (nec Geoffroy, 1785) (ϭ Pro- pomeron moderately and rather gradually de- togoerius brullei, nom. nov.). ¯exed ventrad at about apical fourth of COMMENTS: The single species of this ge- length of pronotum, joining inferior line of nus has the habitus of an Ocypus; it has two pronotal hypomeron well before anterior an- main identifying character states: (1) gula tu- gles, anterior portion of superior line situated mid almost to ®ne basal carina, transverse only moderately below basal portion in lat- impression very faint (®g. 146); (2) dorsal eral view; pronotal epimeron absent; surface portion of median lobe of aedeagus almost of pronotum with microsculpture of ®ne, entirely membranous. short, mostly oblique striae; mesosternum Further character states include: mandibles without transverse carina; punctation of ely- each with one strong, broadly triangular tra very dense, corneous; surface with dense, tooth on medial edge in front of a deep, an- ®ne, rugose microsculpture, appearing dull; gular indentation, with dorsolateral ridge not abdominal tergites with posterior basal line distinctly removed from lateral margin (®g. very gently bisinuate. 147); palpifer with one subapical and three The status of this taxon cannot be estab- shorter apical setae; apical segment of max- lished with certainty at present. The only illary and labial palpi moderately elongate, known male specimen of A. cariniceps is distinctly fusiform, only that of labial palpus somewhat damaged (see below), and it has setose (®g. 148); prostheca parallel-sided, lost all mouthparts (including mandibular elongate with long ciliae only at apex; nuchal prostheca) due to dermestid damage. Bern- ridge present only dorsally, becoming obso- hauer (1934: 9) established it as a subgenus lete well before base of infraorbital ridge lat- of Staphylinus. The general habitus of the erally; dorsal basal ridge very short, ending species is certainly conspicuous, as noted al- at very base of infraorbital ridge; superior ready by Bernhauer (loc. cit.), mainly due to and inferior lines of hypomeron run very the combination of the narrow head and the closely parallel for some distance anteriorly, pronotum markedly narrowed anteriad. Bern- gradually merging along prosternal margin; hauer (loc. cit.) described the pronotum as prothoracic epimeron absent; prosternum ``vor dem Schildchen mit zwei grossen, lan- with short transverse ridge on keel behind gen LaÈngseindruÈcken.'' The impressions are two macrosetae; mesocoxal acetabulum in fact somewhat irregular, and since the deeply depressed, with straight carina slop- pronotum is de®nitely cracked anteriorly, ing obliquely behind intercoxal process; mar- there is a distinct possibility that the prono- gins of anterior metasternal projection con- tum may have been exposed to excessive verging at a highly obtuse angle, projection pressure that caused not only the cracking very short; metasternum very short, at an but also the two irregular impressions; in oth- oblique angle to plane of body, with no vis- er words, the impressions may be arti®cial ible medial line; dorsal apicolateral lobe of and not characteristic of the species. hind coxa with only one or two very ®ne Due to the con®guration of the pronotum, spines; empodial setae very long. particularly the development of the superior BrulleÂ's (1839: 59) name Staphylinus bra- and inferior lines, Apostenolinus is not a chypterus is preoccupied by Staphylinus bra- member of the ``Dinothenarus±Staphylinus'' chypterus Geoffroy, 1785: 176, now a syn- lineage. The impressions on the head and onym of Aleochara curtula (Goeze, 1777), pronotum (if really present on the latter) are and by Staphylinus brachypterus Marsham, likely an autapomorphy. Apostenolinus is 1802: 510, now a synonym of Aploderus considered here as a separate genus only ten- caelatus Gravenhorst, 1802. The replacement tatively, pending the availability of more name Protogoerius brullei, nom. nov.,is specimens for study. hereby proposed for it. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_31 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 31

Ocypus Leach, 1819 (stat. nov.) constriction, directed distinctly posteriad of transverse postgenal ridge. TYPE SPECIES: Staphylinus cyaneus Pay- The type species of Goerius and Xantho- kull, 1789 (ϭ Staphylinus ophthalmicus Sco- cypus ®t well within the subgenus Ocypus, poli, 1763). as it is characterized herein, particularly as COMMENTS: All members of this genus far as the characters on the mouthparts are share the following character states: mandib- concerned. Some species, such as O. weisei, ular dorsolateral ridge not distinctly removed O. quadrimaculatus, O. auroguttatus, and O. from lateral margin, especially in dorsal wasmanni, may appear conspicuous due to view; prostheca long, lanceolate, with short the presence of striking patches of bright to- setae along medial margin, becoming pro- mentose pubescence, but in all other respects gressively longer to moderate near apex; they are typical members of the subgenus postmandibular ridge very short to absent; Ocypus. last segment of labial palpus in general var- iably fusiform, or more or less parallel-sided Subgenus Matidus Motschulsky, 1860 (stat. nov.) and truncate apically (®gs. 131, 132); supe- (type species: Matidus for®cularius Motschul- rior and inferior lines of hypomeron run very sky, 1860). closely parallel for some distance anteriorly, COMMENTS: Members of this subgenus either never distinctly merging or gradually share the following character states: mandib- merging along prosternal margin, superior ular prostheca with ciliae very short; palpifer line moderately and rather gradually de¯ect- with one long subapical and one or two ed ventrad, with its anterior portion situated shorter apical setae (®g. 149); apical segment only slightly to moderately below basal por- of both maxillary and labial palpus with tion in lateral view (®gs. 113, 114); protho- many setae (®gs. 150, 151), apical segment racic epimeron absent (®g. 113); de¯ected of maxillary palpus not distinctly fusiform, portion of pronotal disc visible in ventral with apex broadened, almost as wide as view from along coxal insertion (a little far- width at middle of segment, that of labial ther posteriad in Pseudocypus); anterior me- palpus distinctly fusiform; mentum without dial portion of prosternum sloping very grad- arcuate ridge basally; submentum not dis- ually anteriad; mesocoxal acetabulum deeply tinctly beaded; eye rarely more than half impressed, vertical to convexly carinate be- length of tempus; genal seta removed from hind intercoxal process. margin of eye by at least four times diameter of its puncture; lateral end of nuchal ridge Subgenus Ocypus Leach, 1819. distinctly removed from nuchal constriction, Goerius Westwood, 1827 (syn. nov.) (type spe- directed distinctly posteriad of transverse cies: Staphylinus olens O. MuÈller, 1764). postgenal ridge. Xanthocypus J. MuÈller, 1925 (syn. nov.) (type species: Ocypus weisei Harold, 1877). Subgenus Pseudocypus Mulsant and Rey, 1876 (stat. nov.) (type species: Staphylinus mus COMMENTS: Members of this subgenus BrulleÂ, 1832). share the following character states: mandib- Protocypus J. MuÈller, 1923 (syn. nov.) (type spe- ular prostheca with long ciliae (®g. 13); pal- cies: Ocypus fulvotomentosus Eppelsheim, pifer with one long subapical and two shorter 1889). apical setae (as in ®g. 149); apical segment Atlantogoerius Coiffait, 1956 (type species: Ocy- of labial palpus with many setae (®g. 132), pus sylvaticus Wollaston, 1865). at least as large as that of maxillary palpus, Fortunocypus Coiffait, 1964 (type species: Ocy- which is asetose, fusiform; mentum without pus fortunatarum Wollaston, 1871). arcuate ridge basally; submentum not dis- Nudabemus Coiffait, 1982 (syn. nov.) (type spe- cies: Nudabemus caerulescens Coiffait, 1982). tinctly beaded; eye rarely more than half length of tempus, genal seta removed from COMMENTS: Members of this subgenus margin of eye by at least four diameters of share the following character states: mandib- its puncture, usually more; lateral end of nu- ular prostheca with long ciliae (®g. 13); pal- chal ridge distinctly removed from nuchal pifer with one long subapical and one shorter AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_32 Allen Press • DTPro System File # 01cc

32 AMERICAN MUSEUM NOVITATES NO. 3287

apical seta (®g. 128); apical segment of labial fer to unite Nudabemus with Pseudocypus,as palpus with many setae (®g. 131), that of characterized herein. maxillary palpus asetose, both distinctly fu- The type species of Protocypus is a typical siform; mentum almost always with ®ne, representative of mainly east Palearctic broadly arcuate ridge extending for variable groups of species of Pseudocypus that in distance from posterior margin between ba- general resemble members of the subgenus solateral angles; submentum with anterior Matidus (or even Ocypus). However, in all margin beaded (usually ®nely) throughout; other character states, particularly those of eye usually longer than tempus, less fre- the mouthparts, they ®t well within the sub- quently variably shorter; genal seta removed genus Pseudocypus, as characterized herein. from margin of eye by no more than three times diameter of its puncture; lateral end of Subgenus Aulacocypus J. MuÈller, 1925 (type spe- nuchal ridge only moderately removed from cies: Ocypus gloriosus Sharp, 1874). nuchal constriction, directed to level of trans- COMMENTS: Members of this subgenus verse postgenal ridge; superior and inferior share the following character states: (1) man- lines of hypomeron merging along prosternal dibles each with one simple tooth on medial margin. margin; (2) each lateral portion of posterior The type species of Nudabemus conforms, basal line on ®rst three visible abdominal ter- with a few exceptions, to the subgenus Pseu- gites obtusely extended posteriad, with base docypus, particularly in the con®guration of of tergite between extensions depressed, de- the mandibular teeth, palpifer (one short api- pression delimited laterally by short oblique cal seta), submentum (weakly beaded), and carina situated at middle of each lateral ex- prosternum; the mesocoxal acetabulum is tension of posterior basal line. deeply impressed and convexly carinate be- Further character states include: maxillary hind the intercoxal projection, and the me- palpus with apical segment short, as broad at tasternum is short and oblique. The species apex as at base; apical segment of labial pal- is somewhat unusual for Pseudocypus (but pus short, truncate apically, either more or similar to O. aethiops, or to the east Palearc- less parallel-sided, or variably dilated apicad, tic species previously assigned to Protocy- with apex up to almost three times its width pus); the apical segment of the labial palpus at base. is relatively broad and slightly obliquely J. MuÈller (1925: 40) erected Aulacocypus truncate apically (but as in O. fulvipennis or as a subgenus of Staphylinus to accommo- O. mus); the superior line of the pronotal hy- date two species (see the checklist) that share pomeron is rather abruptly de¯ected ventrad, the distinctive character state of the posterior at a point relatively close to the anterior basal line of the ®rst three visible abdominal pronotal angle, yet it merges with the inferior tergites (see the key and above). This char- line before the prosternal margin (as in O. acter state, combined with the simple tooth mus); the hind tibia has only one spine on its on medial margin of each mandible, justi®es dorsolateral face; the punctation on the head the retention of a separate subgenus within and pronotum is coarse and the elytra are Ocypus for these two species, and tentatively short (as in the east Palearctic species pre- for another species from China (see comment viously assigned to Protocypus). under O. kansuensis below). Ocypus quadrimaculatus, assigned by Coiffait (1982: 31) to Nudabemus, is quite REMARKS ON SOME SPECIES OF OCYPUS different from the type species of Nudabe- mus and belongs to the subgenus Ocypus s. 1. Ocypus almorensis (Cameron, 1932). str. (see above). Cameron (1932: 204) described the species Nudabemus may represent a separate lin- (in Staphylinus) from specimens from ``West eage of east Palearctic species, occurring in Almora: Kali Valley, alt. 9000 feet'' and the Himalayas and particularly in the moun- ``Punjab: Triun, alt. 9300 feet.'' We were tains of mainland China and it may deserve able to study one of the male syntypes in the a subgeneric status; however, until the tax- Natural History Museum, London, England, onomy of this fauna is better known, we pre- labelled as follows: ``Triun, 9,300 ft Dharm- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_33 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 33

sala. Punjab. H. G. C./M. Cameron. Bequest. 5. Ocypus hauseri (Bernhauer, 1933: 32). B.M. 1955-147./Staphylinus (Ocypus) al- The collection at the Field Museum of Nat- morensis.'' The specimen is in very good ural History, Chicago, Illinois, contains one shape. It was dissected, and the aedeagus and male specimen (holotype) under this name. the genital segment were glued to the plate It is labelled as follows: ``Kalgan Jnn-shan with the beetle. It is hereby designated as the 980±2100 m/285/Hauseri Brh. Typus unic./ lectotype of Staphylinus almorensis; the label Hauseri Bernh.Typus unic./Chicago NHMus ``LECTOTYPE Staphylinus almorensis M. Bernhauer Collection.'' The specimen is Cameron A. Smetana des. 1999'' was at- in very good condition. It was dissected, tached to it. The specimen cannot be distin- compared to the lectotype of O. fulvotomen- guished from those of Ocypus helleni J. tosus (see above), and was found to be con- MuÈller that occur in northern Pakistan and in speci®c with it. The name O. hauseri be- Kashmir. The name O. almorensis becomes comes a junior synonym of O. fulvotomen- a junior synonym of O. helleni (syn. nov.). tosus (syn. nov.); the holotype was labelled The corresponding determination label was accordingly. attached to the specimen. 6. Ocypus himalayicus (Cameron, 1935: 2. Ocypus auricomus (Lindberg, 1953: 4), 288) (Staphylinus). Cameron (loc. cit.) de- described in Staphylinus, is preoccupied by scribed the species from a single female from Staphylinus auricomus BrulleÂ, 1842: pl. 5, Karakorum range. The holotype, deposited at ®g. 6, now a synonym of Glenus chrysis the Natural History Museum, London, Eng- (Gravenhorst, 1806), and by Staphylinus au- land, is labelled as follows: ``Type [round ricomus Cameron, 1929: 65. The name Ocy- disc with red margin]/Sind valley 2000±2800 pus mateui (Coiffait, 1954: 170), listed at m 2±5.V.1929/Nederlandsche Karakorum- present as a junior synonym of O. auricomus, Expeditie J. A. Sillem leg./M. Cameron. Be- becomes the valid name of this species (stat. quest. B.M. 1955-147./Staphylinus himalay- nov.). icus Cam. TYPE.'' The specimen is in fair 3. Ocypus bimaculatus (Cameron, 1932: shape (last segment of left front tarsus and 207), described in Staphylinus, is preoccu- claws of middle right tarsus are missing, and pied by several names in Staphylinus, the most of the pubescence of the head and pro- oldest one being Staphylinus bimaculatus notum is missing as well). The specimen Schrank, 1798: 644, now a synonym of Lor- cannot be distinguished from those of Ocy- dithon trinotatus (Erichson, 1839b: 409). pus (Pseudocypus) helleni J. MuÈller that oc- The replacement name Ocypus cameroni, curs in northern Pakistan and in Kashmir. nom. nov., is hereby proposed for Ocypus The name O. himalayicus becomes a junior bimaculatus (Cameron, 1932). synonym of O. helleni (syn. nov.). The cor- 4. Ocypus fulvotomentosus Eppelsheim, responding determination label was attached 1889: 172. The collection at the Naturhisto- to the specimen. risches Museum in Wien, Austria, contains 7. Ocypus italicus (Aragona, 1830: 10). one male specimen under this name. It is la- By coincidence, Erichson (1840: 406) later belled as follows: ``Kan-ssa 1885 G. Patanin/ redescribed this species as new, using the c. Eppelsh. Steind. d./fulvotomentosus det. same name as Aragona. Subsequent authors Bernhauer/fulvotomentosus det. J. MuÈller/ considered Erichson's description as a sub- TYPUS.'' The specimen is in good condi- sequent citation of Aragona's species. How- tion; however, the entire left middle leg, out- ever, based on Erichson's treatment of his er three segments on left hind tarsus, and the Ocypus italicus, including his comments un- entire right middle tarsus are missing. It was der ``Obs.,'' there is little doubt that Erichson dissected and the aedeagus was glued to a considered it a new species. Therefore, the plate attached to the pin, with the beetle. The name Ocypus italicus Erichson, 1840 be- specimen is hereby designated as the lecto- comes a junior synonym of Ocypus italicus type of O. fulvotomentosus; the label ``LEC- (Aragona, 1830) (syn. nov.). TOTYPE Ocypus fulvotomentosus Eppel- 8. Ocypus kansuensis (Bernhauer, 1933: sheim A. Smetana des. 1997'' was attached 33) (Staphylinus). Bernhauer (loc. cit.) de- to it. scribed the species as a member of the sub- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_34 Allen Press • DTPro System File # 01cc

34 AMERICAN MUSEUM NOVITATES NO. 3287

genus Xanthocypus of Staphylinus. We were ``Ocypus lewisius Yokohama. Japan. Lewis able to study the male holotype deposited in [on plate with beetle]/Japan G. Lewis./Sharp the Bernhauer collection at the Field Muse- Coll. 1905-313.'' Spec. 3 (female): ``Ocypus um of Natural History, Chicago, Illinois. It lewisius Yokohama. 27.3.880. Lewis [on is labelled as follows: ``Kansou mer. Hoei- plate with beetle]/Japan. G. Lewis./Sharp Siee./286./Mittel-China don. Dr. G. Hauser/ Coll. 1905-313.'' Spec. 4 (male): ``Japan. G. kansuensis Brnh. typ. unic./Chicago NHMus Lewis. 1910-320./Iokii Tanaka/lewisius Shp M. Bernhauer Collection/kansuensis Brnh. det. J. MuÈller.'' The second male specimen typus un. Xanthocypus.'' The specimen is in (dissected) is hereby designated as the lec- fair condition. Only three basal segments of totype of Ocypus lewisius; the label ``Lec- the left antenna are present, the pronotum is totype Ocypus lewisius Sharp A. Smetana longitudinally cracked on the left side, and des. 1999'' was attached to it. the mesosternum and metasternum are dam- 11. Ocypus nero (Faldermann, 1835: 118) aged, again on the left side. The specimen was recently used by Pilon (1991: 105) as a was obviously exposed to dorsoventral pres- valid name to replace the misidenti®ed sure. It was dissected, and the aedeagus and Staphylinus similis Fabricius, 1792: 521. Fa- genital segment were glued to the plate with bricius (loc. cit.) did not describe Staphylinus the beetle. By the form of the mandibles, the similis as a new species; he attributed it to shape and setation of the last segment of the Paykull. The species Paykull (1789: 10) de- labial palpus, the thoracic structures, the scribed as Staphylinus similis is today in Tas- characteristic con®guration of the posterior gius (Rayacheila), so another name was basal line of the ®rst three visible abdominal needed for the species known for a long time tergites, as well as by the form of the aedea- as Staphylinus similis Fabricius. Pilon (loc. gus, the species ®ts the subgenus Aulacocy- cit.) chose Staphylinus nero as a name, with- pus of Ocypus. out giving any reasons. However, the name 9. Ocypus kobensis Cameron, 1930: 207. Staphylinus nitens Schrank, 1781: 231, used The collection at the Natural History Muse- for the species already by Fauvel (1874: um, London, England, contains one male 411), is older, and therefore Ocypus nitens specimen (holotype) under this name. It is (Schrank) becomes the valid name for this labelled as follows: ``Type [round label with species (stat. nov.). red margin]/X/JAPAN Kobe Harada 7.XI '19 12. Ocypus orientalis (Bernhauer and [last line illegible]/538/Ocypus kobensis Schubert, 1914: 389), established as a re- TYPE Cam./M. Cameron. Bequest. B.M. placement name for Staphylinus tomentosus 1955-147.'' The specimen was dissected Baudi, 1869: 384 (nec Gravenhorst, 1802: (apex of the abdomen was damaged before 161), is preoccupied by Staphylinus orien- the dissection), and the aedeagus and the talis Motschulsky, 1858: 67, now a synonym genital segment (damaged) were glued to the of Creophilus maxillosus. The replacement plate with the beetle. The specimen was name Ocypus orientis, nom. nov., is hereby compared to the lectotype of Ocypus lewisius proposed for Ocypus orientalis (Bernhauer (see below) and was found to be conspeci®c. and Schubert, 1914). The name Ocypus kobensis is therefore a ju- 13. Ocypus pullus Hochhuth, 1849: 121 nior synonym of Ocypus lewisius (syn. and Ocypus simulator Eppelsheim, 1878: nov.); the holotype was labelled accordingly. 420. These two species were declared as be- 10. Ocypus lewisius Sharp, 1874: 33. The ing possibly identical by Eppelsheim (1887: collection at the Natural History Museum, 432) and by J. MuÈller (1926d: 14); however, London, England, contains four conspeci®c MuÈller (loc. cit.) suggested this co-identi®- specimens of the original series under this cation only tentatively, and he pointed out name, as follows: spec. 1 (male): ``Type some differences between them. Both species [round label with red margin]/Beach Kobe belong to a dif®cult group, and their status Oct. 69 [on underside of plate with beetle]/ can only be determined by the study of Sharp Coll. 1905-313./Ocypus lewisius. type Hochhuth's original material, particularly of D.S.'' Spec. 2 (male, dissected, genital seg- the characters on the aedeagus. They very ment and aedeagus on plate with beetle): likely are not identical and are therefore both AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_35 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 35

listed in the checklist as separate species. The solete along prosternal margin, de¯ected por- aedeagus of the holotype of O. simulator tion of pronotal disc only narrowly visible (``Turcia Merkl'') was illustrated by Smetana near anterior angles in ventral view (®g. 46). (1965b: 40); the second specimen of the Further character states include: most oth- original series of O. simulator comes from er characters shared with species of Ocypus ``Acarnanien.'' Ocypus pullus was described (Ocypus), with following exceptions: labial from specimens ``Aus verschiedenen Gegen- palpus more broadly expanded apically, up den Kaukasiens.'' to 1.5 times width at base (®g. 153); lateral 14. Ocypus rossii (Jarrige, 1954: 164) margins of mentum relatively long, straight, (Staphylinus cupreus Rossi, 1790: 248; nec as in O. (Matidus) brenskei (Reitter, 1884); Staphylinus cupreus Fourcroy, 1785: 173). mesosternum with sharp, convex carina be- Coiffait (1974: 495, 513) recognized three hind intercoxal process, as in O. (Ocypus) subspecies of this species (under the name of af®nis Wollaston, 1864. P. cupreus): P. cupreus cupreus from west- The type species of Apecholinus agrees in ern and southwestern Europe, P. cupreus ser- all character states with the members of the iceicollis from the east Mediterranean region, genus Agelosus, except for the dual puncta- and P. cupreus fulvicupreus from the island tion (®ne and coarse) on the dorsal side of of Cyprus. The three subspecies were based the head and on the pronotum. The character on differences in the shape of the apical por- state of the dual punctation is not considered tion of the paramere of the aedeagus. In fact, to be suf®cient for retaining a separate taxon the apical portion of the paramere, as well as for the species. that of the median lobe, shows distinct var- iability that is not geographically delimited. Wasmannellus Bernhauer, 1920 The differences given in the key by Coiffait (1974: 495) cannot be con®rmed (based on TYPE SPECIES: Wasmannellus tristis Bern- substantial material from the entire geo- hauer, 1920. graphical range of the species). Therefore, COMMENTS: Members of this genus share the two subspecies are placed in synonymy the following character states: (1) superior with the nominal species. Jarrige (loc. cit.) line of pronotal hypomeron moderately sep- proposed the name O. rossii as a replacement arate from inferior line anteriorly, becoming name (see above). However, three older syn- obsolete near anterior angle of pronotum; (2) onyms are available: Goerius con®nis Ste- bases of ®rst four visible abdominal tergites phens, 1832: 211, Ocypus angustatus Ste- markedly transversely impressed, with punc- phens, 1832: 212, and Staphylinus sericei- tation different from that on rest of tergites; collis MeÂneÂtrieÂs, 1832: 143. Based on the (3) hind tibia without spines on dorsolateral concept discussed above, the latter name by face. MeÂneÂtrieÂs was chosen. The name Ocypus Further character states include: apical sericeicollis (MeÂneÂtrieÂs) is therefore the valid segment of maxillary palpus asetose, almost name of this species (stat. nov.). as wide apically as at middle; apical segment of labial palpus setose, apex expanded to Agelosus Sharp, 1889 about 1.5 times width of base, slightly TYPE SPECIES: Goerius carinatus Sharp, obliquely securiform; lateral end of nuchal 1874. ridge distinctly removed from nuchal con- striction, directed distinctly posteriad of Apecholinus Bernhauer, 1933 (syn. nov.) (type transverse postgenal ridge; pronotal hypo- species: Apecholinus kaiseri Bernhauer, 1933). meron very broad at middle, de¯ected por- COMMENTS: Members of this genus share tion of pronotal disc visible only from ante- the following two character states: (1) infra- rior edge of coxal insertion in ventral view; orbital ridge extended signi®cantly anteriad prothoracic epimeron absent; medial margin of postgenal ridge (®g. 152); and (2) superior of sternacostal ridge extended anteriad into and inferior lines of pronotal hypomeron dis- sharp, ®nlike carina; serial setae on meso- tinctly, widely separated at anterior angles of sternum inserted along edge of elevated, ir- pronotum, with inferior line becoming ob- regular ridge parallel to margins; mesosternal AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_36 Allen Press • DTPro System File # 01cc

36 AMERICAN MUSEUM NOVITATES NO. 3287

intercoxal process with margins converging able, from as broad at apex as at base to 1.5 acutely, acute at apex, distinctly margined times as broad; apical segment of labial pal- throughout; margins of anterior metasternal pus with apex broadened from about 1.5±2 projection converge rectangularly, very times its width at base. broadly rounded at apex. Further character states include: mandib- The genus at present contains only two ular dorsolateral ridge rarely represented by species, but there is at least one additional more than a ®ne line, always very close to undescribed species in the eastern part of the lateral margin; mentum frequently with more Palearctic region (mainland China). than two setae near anterolateral angles along anterior margin; submentum with anterior Tasgius Stephens, 1829 (stat. nov.) margin straight; postmandibular ridge absent, rarely weak but reaching almost halfway to TYPE SPECIES: Astrapaeus ru®pes Latreille, infraorbital puncture; gula tumid to near 1806 (ϭ Staphylinus pedator Gravenhorst, base, with basal transverse impression very 1802). ®ne and along margin; postgenal ridge di- COMMENTS: Members of this genus share rected obliquely anteriad; inferior line of the following character states: (1) mandibles pronotal hypomeron runs close to superior without teeth, or each with only one poorly line for some distance anteriad, never dis- differentiated tooth (®gs. 4, 5); (2) apical tinctly merging with it and becoming obso- segment of all palpi with many setae; (3) me- lete along prosternal margin (T. pedator), or sosternal intercoxal process highly acute api- merging just behind prosternal margin (T. cally, extended posteriad into very deeply ater) (®g. 155); sternacostal ridge not dis- depressed acetabulum as sharp, straight to tinctly arched posteriad, but straight and highly convex, or even angulate carina (®g. slightly oblique toward keel; anterior meta- 154). sternal projection with margins converging Further character states include: palpifer highly obtusely. with one long lateral subapical seta and one Pseudotasgius is based on the same type shorter medioapical seta; base of infraorbital species. Paratasgius was based on a single ridge distinctly recurved toward nuchal ridge character: the presence of the prothoracic dorsally; prothoracic epimeron ranging from epimeron. Since the development of the pro- absent to moderately developed, broadly thoracic epimeron varies within Tasgius (see rounded in all subgenera; prosternum long, above), there is no justi®cation for recogniz- at least half as long as maximum width (®g. ing Paratasgius as a separate taxon, as was 155); series of long setae on mesosternum done, for example, by Coiffait (1974: 517). inserted along edge of a feeble, irregular, broadly U-shaped ridge, close to margins of Subgenus Rayacheila Motschulsky, 1845 (stat. projection; dorsal apicolateral lobe of hind nov.) (type species: Rayacheila inderiensis coxa with two to three very short, relatively Motschulsky, 1845). ®ne spines. Anodus Nordmann, 1837 (nec Spix, 1829) (syn. nov.) (type species: Staphylinus morio Grav- Subgenus Tasgius Stephens, 1829. enhorst, 1802). Pseudotasgius Seidlitz, 1891 (syn. nov.) (type Alapsodus Tottenham, 1939 (syn. nov.) (type spe- species: Staphylinus pedator Gravenhorst, cies: Staphylinus morio Gravenhorst, 1802). 1802). Allocypus Coiffait, 1964 (syn. nov.) (type species: Paratasgius Jarrige, 1952 (syn. nov.) (type spe- Staphylinus winkleri Bernhauer, 1906). cies: Staphylinus ater Gravenhorst, 1802). Metocypus Coiffait, 1964 (syn. nov.) (type spe- cies: Staphylinus globulifer Geoffroy, 1785). COMMENTS: Members of this subgenus Paralapsodus Coiffait, 1974 (syn. nov.) (type share the following character states: (1) man- species: Staphylinus solskyi Fauvel, 1875). dibles each with only one poorly differenti- ated tooth, moderately narrow basally (®g. COMMENTS: Members of this subgenus 5); (2) prostheca long, lanceolate, with short share the following character states: (1) man- ciliae along medial margin, becoming grad- dibles each without teeth, falciform, ¯at, ually longer to moderate at apex (®g. 14); (3) very narrow to near base (®g. 4); and (2) maxillary palpus with apical segment vari- mandibular prostheca elongate, narrow, par- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_37 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 37

allel-sided, with ciliae only near apex (®g. 2. Tasgius bicolor (Cameron, 1944: 11). 16). The collection at the Natural History Muse- Further character states include: maxillary um, London, England, contains one male palpus with apical segment variable, from as specimen (holotype) under this name. It is broad at apex as at base to 1.5 times as labelled as follows: ``Type [round label with broad; apical segment of labial palpus with red margin]/Govt. Entomologist, Punjab CI/ apex broadened from about 1.5 to more than Under leaves in sweet potato ®eld Ayallpur 4 times its width at base (variable within spe- 23.6.1939 M. A. Ghani/Pres. by Imp. Inst. cies, with base sometimes very narrow); Ent. B.M. 1940/S. (Ocypus) bicolor TYPE mentum short, with an even, broadly rounded Cam.'' The mouthparts (including both pal- transverse discal swelling between basolater- pi) and mandibles con®rm the assignment of al angles so that basal portion depressed and this species to Tasgius (Tasgius). A label anterior portion slightly de¯ected at an angle ``belongs to Tasgius (Tasgius) Det. A. Sme- to submentum (®g. 156); submentum with tana 1999'' was attached to the specimen. anterior margin broadly, arcuately rounded 3. Tasgius eppelsheimianus (Jakobson, (®g. 156); postmandibular ridge absent; post- 1909: 510). This is a replacement name for genal ridge directed obliquely anteriad, Ocypus ru®pes Eppelsheim, 1884: 15 (nec broadly obsolete or confused near gula; lat- Astrapaeus ru®pes Latreille, 1806: 285), eral end of nuchal ridge directed toward, or commonly used for the species. At the time anteriad of, transverse postgenal ridge; infe- of the replacement, both names were listed rior line of pronotal hypomeron runs close to in Staphylinus. Although both are now in dif- superior line for some distance anteriad, ferent genera, a junior secondary homonym merging with it along prosternal margin; replaced before 1961 is permanently invalid prosternum with long, broadly arcuate ®ne (Art. 59[b] of the Code). Bernhauer (1900: carina behind macrosetae medially (®g. 157); 55) described Ocypus ru®pes var. obscuripes, sternacostal ridge distinctly arched posteriad, which predates Jakobson's replacement name or distinctly oblique toward keel; anterior and may be in fact the valid name of this metasternal projection with margins converg- species. However, the status of this ``variety'' ing highly obtusely, apex broadly rounded is not clear at present, and therefore we con- (®g. 158). sider it as a doubtful name pending a revision Until recently, this subgenus was known of Bernhauer's original material. under the name Alapsodus, a name provided by Tottenham (1939: 225) to replace the pre- CHECKLIST OF NORTH TEMPERATE occupied Nordmann name Anodus. However, TAXA BohaÂcÏ (1988: 553) revised the type specimen of the type species of Rayacheila (R. inder- This checklist cites, at the generic level, iensis, incorrectly cited by BohaÂcÏ [loc. cit.] only valid taxa, new synonyms, and from as described in Ocypus) and recognized it as previously recognized synonyms only those a species of Metocypus. Since Metocypus, that are for some reason important; similarly, along with Allocypus and Paralapsodus,isa at the speci®c level, only currently recog- super¯uous taxon based solely on aedoeagal nized, valid speci®c and subspeci®c (or characters, which was already placed in syn- ranked as subspeci®c at least once in the onymy with Alapsodus by DvorÏaÂk (1984: past) names are given, with the exception of 66), Rayacheila becomes the valid name of a few synonyms of importance, and the new the taxon. synonyms. Valid names are in italics; syno- nyms in roman type. A complete listing of REMARKS ON SOME SPECIES OF TASGIUS all names, at both generic and speci®c levels, including complete references, will be avail- 1. Tasgius amoenus (Reitter, 1909: 122), able in the upcoming world catalog of Staph- described in Staphylinus, is preoccupied by ylinidae by Herman. Staphylinus amoenus Olivier, 1795. The re- As speci®ed in the title of this paper, the placement name Tasgius amoenanus, nom. species listed occur only in the northern tem- nov., is proposed for it. perate zone, that is in the Palearctic and Ne- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_38 Allen Press • DTPro System File # 01cc

38 AMERICAN MUSEUM NOVITATES NO. 3287

arctic Regions, with some overlap into the the checklist: Pa (ϭ Palearctic); Ne (ϭ Ne- northern areas of the Oriental Region and arctic); Nt (ϭ Neotropical); Or (ϭ Oriental). into the Neotropical Region. The limits of the Palearctic Region are identical, mainly CREOPHILUS LEACH, 1819 for practical reasons, with those speci®ed for the contributors to the Catalogue of the Co- C. maxillosus (LinneÂ, 1758) ± Pa, Or, Ne leoptera of the Palearctic Region, which is C. maxillosus maxillosus (LinneÂ, 1758) ± Pa in preparation. The southern limits of the Pa- C. maxillosus villosus (Gravenhorst, 1802) ± learctic Region are, from west to east, as fol- Ne lows: Atlantic Islands (including Azores), C. sikkimensis Wendeler, 1927 ± Pa Africa north of Sahara (Morocco, Algeria, C. villipennis Kraatz, 1859 ± Pa, Or Tunisia, Egypt), the entire Arabian Peninsula (including Socotra), all of Pakistan, areas EMUS LEACH, 1819 along the base of the Himalayas (Kashmir, Himachal Pradesh, Uttar Pradesh, Nepal, E. griseosericans Fairmaire, 1894 ± Pa Sikkim and Darjeeling, Bhutan, Arunachal E. hirtus (LinneÂ, 1758) ± Pa Pradesh), all of mainland China (including Hainan), Taiwan, Japan (including Ryukyus NADDIA FAUVEL, 1867 and Bonin Isles), and the Kuril Islands. The taxa at the subspeci®c level are listed N. assamensis Cameron, 1932 ± Pa, Or as they are recognized at present; however, N. atripes Bernhauer, 1939 ± Pa many of them, in addition to those presented N. chinensis Bernhauer, 1929 ± Pa in this paper, will be placed in synonymy N. decipiens Cameron, 1932 ± Pa, Or once the complexes are properly studied (see N. ishiharai Shibata, 1994 ± Pa also the Introduction). This is particularly N. malaisei Scheerpeltz, 1965 ± Or, Pa true for those erected for Ocypus (Ocypus) N. miniata Fauvel, 1895 ± Pa, Or ophthalmicus, even quite recently (Drug- N. monticola Shibata, 1994 ± Pa mand, 1998), and, to a lesser extent, for Ocy- N. ru®pennis Bernhauer, 1915 ± Or, Pa pus picipennis (see the new synonymies un- N. taiwanensis Shibata, 1979 ± Pa der this species). The new synonymies at the N. wittmeri Coiffait, 1982 ± Pa, Or subspeci®c level presented in the following checklist are those that were established ONTHOLESTES GANGLBAUER, 1895 based on the study of much material (not necessarily type material), including the var- O. aurosparsus (Fauvel, 1895) ± Pa, Or iability of the apical portion of the median O. callistus (Hochhuth, 1849) ± Pa lobe and the paramere of the aedeagus. When O. chalcopygus (Hochhuth, 1849) ± Pa additional details (e.g., data on the original O. cingulatus (Gravenhorst, 1802) ± Ne material studied) were deemed necessary O. dieckmanni Smetana, 1958 ± Pa (this information is given for all new syn- O. gracilis (Sharp, 1874) ± Pa onymies at speci®c level, but see Platydracus O. hairaerensis Li and Chen, 1993 ± Pa below), these were given in ``Remarks on O. haroldi (Eppelsheim, 1884) ± Pa Some Species'' under the proper genus/sub- O. hayashii Li, 1992 ± Pa genus. The synonymies presented in the ge- O. inauratus (Mannerheim, 1830) ± Pa nus Platydracus are based on the results of O. marginalis (GeneÂ, 1836) ± Pa the study of the original material by Newton O. murinus (LinneÂ, 1758) ± Pa, Ne (in litteris), and he should be credited with O. oculatus (Sharp, 1874) ± Pa them. The species described by Li (1992) O. orientalis Bernhauer, 1906 ± Pa and by Li and Chen (1993) from China were O. paramurinus Li and Chen, 1993 ± Pa left in the genera to which the authors as- O. proximus Kirschenblat, 1936 ± Pa signed them; these assignments may not be O. simulator Kirschenblat, 1936 ± Pa correct. O. tenuicornis (Kraatz, 1859) ± Pa The following abbreviations are used in O. tessellatus (Geoffroy, 1785) ± Pa AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_39 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 39

THORACOSTRONGYLUS BERNHAUER, 1915 P. centralis (Sharp, 1884) ± Ne, Nt ± (comb. (Parontholestes Coiffait, 1982) nov.) (from Staphylinus) T. birmanus (Fauvel, 1895) ± Pa, Or P. modestus (Sharp, 1884) (comb. nov., syn. T. costatus (Fauvel, 1895) ± Or, Pa nov.) (from Staphylinus) T. formosanus Shibata, 1982 ± Pa P. fusiformis (Casey, 1915) (comb. nov., syn. T. malaisei Scheerpeltz, 1965 ± Or, Pa nov.) (from Staphylinus) T. miyakei Bernhauer, 1943 ± Pa P. chalcescens (Sharp, 1889) ± Pa ± (comb. nov.) T. nepalicus (Coiffait, 1982) ± Pa ± (comb. nov.) (from Staphylinus) (from Parontholestes) P. chalcocephalus (Fabricius, 1801) ± Pa T. velutinus Scheerpeltz, 1965 ± Or, Pa P. chapmani (Bernhauer, 1933) ± Pa ± (comb. nov.) (from Staphylinus) ABEMUS MULSANT AND REY, 1876 P. chinensis (Bernhauer, 1914) ± Pa ± (comb. nov.) (from Staphylinus) A. chloropterus (Panzer, 1796) ± Pa P. cinnamopterus (Gravenhorst, 1802) ± Ne A. hebraeus (Smetana, 1978) ± Pa ± (comb. nov.) P. coeruleipennis Coiffait, 1983 ± Pa (from Staphylinus) P. collaris Coiffait, 1977 ± Pa A. olivaceus (Cameron, 1928) ± Pa ± (comb. P. comes (LeConte, 1863) ± Ne ± (comb. nov.) nov.) (from Staphylinus) (from Staphylinus) A. martensi (Coiffait, 1982) (comb. nov., syn. P. consularis (Bernhauer, 1915) ± Pa ± (comb. nov.) (from Thoracostrongylus) nov.) (from Staphylinus) P. cupripennis (Melsheimer, 1845) ± Ne ± (comb. PLATYDRACUS THOMSON, 1858 nov., stat. nov.) (from Staphylinus) Subgenus Platydracus Thomson, 1858 P. violaceus (Gravenhorst, 1802) (nec Olivier, (Neotasgius J. MuÈller, 1925) 1795) (syn. nov.) P. aeneoniger (Bernhauer, 1933) ± Pa ± (comb. P. fenyesi (Bernhauer, 1917) (comb. nov., syn. nov.) (from Staphylinus) nov.) (from Staphylinus) P. amamiensis Ito, 1982 ± Pa P. dauricus (Mannerheim, 1830) ± Pa P. asemus (Kraatz, 1859) ± Pa ± (comb. nov.) P. demissus (J. MuÈller, 1925) ± Pa ± (comb. nov.) (from Staphylinus) (from Staphylinus) P. aeneicollis (Bernhauer, 1911) (comb. nov.) P. dudgeoni (Cameron, 1932) ± Pa ± (comb. nov.) (from Staphylinus) (from Staphylinus) P. aureofasciatus (Motschulsky, 1862) ± Pa ± P. exulans (Erichson, 1839) ± Ne ± (comb. nov.) (comb. nov.) (from Staphylinus) (from Staphylinus) P. teter (Bondroit, 1913) (comb. nov.) (from P. femoratus (Fabricius, 1801) ± Ne, Nt ± (comb. Staphylinus) nov.) (from Staphylinus) P. aurosericans (Fairmaire, 1891) ± Pa ± (comb. P. varipes (Sachse, 1852) (comb. nov., syn. nov.) (from Staphylinus) nov.) (from Staphylinus) P. becquarti (Bernhauer, 1938) ± Pa ± (comb. P. ¯avopunctatus (Latreille, 1804) ± Pa nov.) (from Staphylinus) P. formosae (Bernhauer, 1933) ± Pa P. brachycerus (nom. nov.)±Pa P. fossator (Gravenhorst, 1802) ± Ne P. brevicornis (Weise, 1877) (nec Motschulsky, P. fulvipes (Scopoli, 1763) ± Pa 1862) (comb. nov.) (from Ocypus) P. brevicornis (Motschulsky, 1862) ± Pa P. fuscolineatus (Bernhauer, 1934) ± Pa ± (comb. P. paganus Sharp, 1874 nov.) (from Staphylinus) P. brevipennis (nom. nov.)±Pa P. goryi (Laporte, 1835) ± Pa, Or ± (comb. nov.) P. brachypterus (Kraatz, 1859) (nec Geoffroy, (from Staphylinus) 1785; nec Marsham, 1802; nec BrulleÂ, P. auripennis (Kraatz, 1859) (comb. nov.) (from 1839) (comb. nov.) (from Staphylinus) Staphylinus) P. caliginosus (Erichson, 1839) ± Ne, Nt P. hauserianus (Bernhauer, 1933) ± Pa ± (comb. P. ejulans (Tottenham, 1939) nov.) (from Staphylinus) P. campestris Coiffait, 1977 ± Pa P. hypocrita (J. MuÈller, 1925) ± Pa P. catalonicus Coiffait, 1967 ± Pa P. immaculatus (Mannerheim, 1830) (see com- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_40 Allen Press • DTPro System File # 01cc

40 AMERICAN MUSEUM NOVITATES NO. 3287

ment under P. vulpinus under Platydracus P. pinorum (Casey, 1915) ± Ne ± (comb. nov.) above) (from Staphylinus) P. imperatorius (Bernhauer, 1916) ± Pa ± (comb. P. plebejus (Bernhauer, 1915) ± Pa nov.) (from Staphylinus) P. praelongus (Mannerheim, 1830) ± Ne P. impotens (Eppelsheim, 1889) ± Pa ± (comb. P. pratti (Scheerpeltz, 1962) ± Pa ± (comb. nov.) nov.) (from Staphylinus) (from Staphylinus) P. indicus (Kraatz, 1859) ± Pa, Or ± (comb. nov.) P. pseudopaganus (Bernhauer, 1914) ± Pa ± (from Staphylinus) (comb. nov.) (from Staphylinus) P. lineatus (Walker, 1859) (comb. nov.) (from P. pseudopaganus pseudopaganus (Bern- Staphylinus) hauer, 1914) P. inornatus (Sharp, 1874) ± Pa P. pseudopaganus pseudopatricius (J. MuÈll- P. kasyi (Scheerpeltz, 1962) ± Pa er, 1926) (comb. nov.) (from Staphylinus) P. kiulungensis (Bernhauer, 1933) ± Pa ± (comb. P. reitterianus (Bernhauer, 1933) ± Pa ± (comb. nov.) (from Staphylinus) nov.) (from Staphylinus) P. kiushiuensis (Bernhauer, 1939) ± Pa ± (comb. P. riojanus Hozman, 1977 ± Pa nov.) (from Staphylinus) P. rutilicauda (Horn, 1879) ± Ne ± (comb. nov.) P. latebricola (Gravenhorst, 1806) ± Pa (from Ocypus) P. lomii (Cerruti, 1951) ± Pa ± (comb. nov.) (from P. sachalinensis (Matsumura, 1911) ± Pa ± Staphylinus) (comb. nov.) (from Staphylinus) P. maculipennis (Kraatz, 1859) ± Pa ± (comb. P. semipurpureus (Kraatz, 1859) ± Pa ± (comb. nov.) (from Staphylinus) nov.) (from Staphylinus) P. meridionalis (Rosenhauer, 1847) ± Pa P. sepulchralis (Erichson, 1839) ± Ne ± (comb. P. montanus Coiffait, 1977 ± Pa nov.) (from Staphylinus) P. mortuorum (Bernhauer, 1912) ± Pa ± (comb. P. modestus (Fall, 1907) (nec Sharp, 1884) nov.) (from Staphylinus) (comb. nov., syn. nov.) (from Staphyli- P. muellerianus (Scheerpeltz, 1933) ± Pa ± nus) (comb. nov.) (from Staphylinus) P. neomexicanus (Bernhauer and Schubert, P. banghaasi (J. MuÈller, 1932) (nec Staphylinus 1914) (comb. nov., syn. nov.) (from Sta- banghaasi Bernhauer, 1915) (comb. nov.) phylinus) (from Staphylinus) P. sharpi (Fauvel, 1901) ± Pa ± (comb. nov.) P. mysticus (Erichson, 1840) ± Ne ± (comb. nov.) (from Staphylinus) (from Staphylinus) P. sparsus (Cameron, 1932) ± Pa ± (comb. nov.) P. ¯uviaticus (Casey, 1915) (comb. nov., syn. (from Staphylinus) nov.) P. speculifrons (Bernhauer, 1939) ± Pa ± (comb. P. nepalensis (Scheerpeltz, 1976) ± Pa ± (comb. nov.) (from Staphylinus) nov.) (from Staphylinus) P. stercorarius (Olivier, 1795) ± Pa P. oculosus (nom. nov.)±Pa P. stercorarius stercorarius (Olivier, 1795) P. oculatus (Bernhauer, 1929) (nec Fabricius, P. stercorarius fuscofemoratus (J. MuÈller, 1775; nec O. F. MuÈller, 1776) (comb. nov., 1923) syn. nov.) (from Staphylinus) P. subaeneipennis (Scheerpeltz, 1976) ± Pa ± P. opaciceps (Scheerpeltz, 1976) ± Pa ± (comb. (comb. nov.) (from Staphylinus) nov.) (from Staphylinus) P. subaeneus (Roth, 1851) ± Pa P. parvulus (nom. nov.) ± Pa, Or P. subauronotatus Coiffait, 1977 ± Pa P. parvus (Cameron, 1932) (nec Solier, 1849) P. submarmorellus (Schubert, 1908) ± Pa (comb. nov., syn. nov.) (from Staphyli- P. subviridis (Bernhauer, 1933) ± Pa ± (comb. nus) nov.) (from Staphylinus) P. perniger (Scheerpeltz, 1976) ± Pa ± (comb. P. sumakowi (Bernhauer, 1911) ± Pa nov.) (from Staphylinus) P. tarsalis (Mannerheim, 1843) ± Ne ± (comb. P. perreaui Coiffait, 1984 ± Pa nov.) (from Staphylinus) P. phoenicurus (Nordmann, 1837) ± Ne ± (comb. P. submetallicus (LeConte, 1861) (comb. nov.) nov.) (from Staphylinus) (from Staphylinus) P. macgregori (Cooper, 1933) (comb. nov., syn. P. lucanus (Horn, 1894) (comb. nov., syn. nov.) nov.) (from Staphylinus) (from Staphylinus) AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_41 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 41

P. curticollis (Bernhauer, 1917) (comb. nov., 1802; nec Latreille, 1804; nec O. Costa, syn. nov.) (from Staphylinus) 1839) P. temporalis (Casey, 1915) ± Ne ± (comb. nov.) (from Staphylinus) DINOTHENARUS THOMSON, 1858 (stat. P. tomentosus (Gravenhorst, 1802) ± Ne nov.) P. cubae (Jacquelin du Val, 1857) (comb. nov., Subgenus Dinothenarus Thomson, 1858 syn. nov.) (from Ocypus) (Protabemus Scheerpeltz, 1966) P. ussuriensis (Solsky, 1871) ± Pa ± (comb. nov.) D. capitatus (Bland, 1864) ± Ne (from Staphylinus) D. choui Smetana, 1992 ± Pa P. vicarius (Sharp, 1889) ± Pa ± (comb. nov.) D. ¯avocephalus (Goeze, 1777) ± Pa (from Staphylinus) D. ¯avocephalus ¯avocephalus (Goeze, P. viduatus (Fabricius, 1801) ± Ne 1777) P. maculosus (Gravenhorst, 1802) D. ¯avocephalus adonis (Coiffait, 1956) P. viridanus (Horn, 1879) ± Ne ± (comb. nov.) D. pubescens (DeGeer, 1774) ± Pa (from Staphylinus) D. ru®ceps (Cameron, 1932) ± Pa P. triplicans (Casey, 1924) (comb. nov., syn. D. sagaris Smetana, 1992 ± Pa nov.) (from Staphylinus) D. xanthocephalus (Kraatz, 1859) ± Pa P. vulpinus (Nordmann, 1837) ± Ne P. wittmeri Coiffait, 1977 ± Pa Subgenus Parabemus Reitter, 1909 (stat. nov.) P. yolensis (Cerruti, 1951) ± Pa ± (comb. nov.) (Parocypus Bernhauer, 1915) (from Staphylinus) (Hypabemus Scheerpeltz, 1966) P. yunnanensis (Bernhauer, 1933) ± Pa ± (comb. nov.) (from Staphylinus) D. arrosus (Eppelsheim, 1890) ± Pa ± (comb. P. yunnanicus (nom. nov.)±Pa nov.) (from Parabemus) P. yunnanensis (Bernhauer, 1943) (nec Bern- D. badipes (LeConte, 1863) ± Ne ± (comb. nov.) hauer, 1933) (comb. nov., syn. nov.) (from Staphylinus) (from Staphylinus) D. chrysocomus (Mannerheim, 1830) ± Pa ± P. zonatus (Gravenhorst, 1802) ± Ne (comb. nov.) (from Parabemus) P. badius (Mannerheim, 1830) (syn. nov.) D. coiffaiti Bordoni, 1977 ± Pa ± (comb. nov.) P. quadraticeps (Casey, 1924) (comb. nov., syn. (from Parabemus) nov.) (from Staphylinus) D. dehradunensis (Bernhauer, 1915) ± Pa ± P. caseyi (Scheerpeltz, 1933) (comb. nov., syn. (comb. nov.) (from Ocypus) nov.) (from Staphylinus) D. fossor (Scopoli, 1772) ± Pa ± (comb. nov.) (from Parabemus) Subgenus Chaetodracus J. MuÈller, 1926 D. fossor (Fabricius, 1792) (comb. nov., syn. nov.) P. patricius (Bernhauer, 1915) ± Pa D. baderlei (Scheerpeltz, 1966) (comb. nov.) D. ganglbauerianus (Bernhauer, 1939) ± Pa ± STAPHYLINUS LINNEÂ , 1758 (stat. nov.) (comb. nov.) (from Ocypus) D. insignis (J. MuÈller, 1926) ± Pa ± (comb. nov.) S. caesareus Cederhjelm, 1798 ± Pa (from Staphylinus) S. caesareus caesareus Cederhjelm, 1798 D. fokiensis (Bernhauer, 1933) ± Pa ± (comb. S. caesareus corporaali Sainte Claire De- nov., syn. nov.) (from Staphylinus) ville, 1927 D. kiautschauensis (Bernhauer, 1916) ± Pa ± S. daimio Sharp, 1889 ± Pa (comb. nov.) (from Staphylinus) S. dimidiaticornis Gemminger, 1851 ± Pa D. luteipes (LeConte, 1861) ± Ne ± (comb. nov.) S. erythropterus LinneÂ, 1758 ± Pa (from Staphylinus) S. erythropterus erythropterus LinneÂ, 1758 D. nigrellus (Horn, 1879) ± Ne ± (comb. nov.) S. erythropterus springeri J. MuÈller, 1923 (from Ocypus) S. medioximus Fairmaire, 1852 ± Pa D. pleuralis (LeConte, 1861) ± Ne ± (comb. nov.) S. ornaticauda (LeConte, 1863) ± Ne (from Staphylinus) S. rubricornis (AdaÂm, 1987) ± Pa D. prainae (Eppelsheim, 1895) ± Pa ± (comb. S. ru®cornis Bernhauer, 1913 (nec Gravenhorst, nov.) (from Staphylinus) AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_42 Allen Press • DTPro System File # 01cc

42 AMERICAN MUSEUM NOVITATES NO. 3287

D. saphyrinus (LeConte, 1861) ± Ne ± (comb. O. kyproticus Coiffait, 1964 (as subsp. of O. nov.) (from Ocypus) gridellii) D. sibiricus (Gebler, 1830) ± Pa ± (comb. nov.) O. pseudolens Coiffait, 1964 (as subsp. of O. (from Parabemus) gridellii) D. szechuanensis (Bernhauer, 1935) ± Pa ± O. cyaneopubens (Reitter, 1913) ± Pa (comb. nov.) (from Staphylinus) O. festae (J. MuÈller, 1926) ± Pa O. miwai (Bernhauer, 1943) ± Pa MIOBDELUS SHARP, 1889 O. olens (O. MuÈller, 1764) ± Pa, Ne O. olens olens (O. MuÈller, 1764) M. brevipennis Sharp, 1889 ± Pa O. olens azoricus (MeÂquignon, 1942) O. ophthalmicus (Scopoli, 1763) ± Pa PHYSETOPS MANNERHEIM, 1830 O. ophthalmicus ophthalmicus (Scopoli, P. ahrendti Wendeler, 1921 ± Pa 1763) P. giganteus Semenov, 1906 ± Pa O. cyanochloris Hochhuth, 1849 P. herculeanus Semenov, 1906 ± Pa O. baicalensis (Eppelsheim, 1887) (as va- P. spilleri Bernhauer, 1923 ± Pa riety of O. ophthalmicus) P. tataricus (Pallas, 1773) ± Pa O. ophthalmicus atrocyaneus (Fairmaire, 1860) ASCIALINUS BERNHAUER, 1933 O. ibericus Coiffait, 1964 (as subsp. of O. ophthalmicus) A. beckeri Bernhauer, 1933 ± Pa O. pernigrus Coiffait, 1964 (as subsp. of O. ophthalmicus)(syn. nov.) APOSTENOLINUS BERNHAUER, 1934 O. ophthalmicus hypsibatus (Bernhauer, A. cariniceps (Bernhauer, 1934) ± Pa 1899) O. ophthalmicus balcanicus (J. MuÈller, 1923) PROTOGOERIUS COIFFAIT, 1956 O. ophthalmicus balearicus (J. MuÈller, 1926) O. ophthalmicus rodopensis Coiffait, 1971 P. brullei (nom. nov.)±Pa O. ophthalmicus benoiti Drugmand, 1998 P. brachypterus (BrulleÂ, 1839) (nec Geoffroy, O. ophthalmicus brigitteae Drugmand, 1998 1785; nec Marsham, 1802) O. quadrimaculatus (Cameron, 1932) ± Pa, Or ± (comb. nov.) (from Nudabemus) OCYPUS LEACH, 1819 (stat. nov.) O. rhaeticus Eppelsheim, 1873 ± Pa Subgenus Ocypus Leach, 1819 O. wasmanni (Bernhauer, 1920) ± Pa ± (comb. (Goerius Westwood, 1827) nov.) (from Nudabemus) (Xanthocypus J. MuÈller, 1925) O. weisei Harold, 1877 ± Pa ± (comb. nov.) (from O. af®nis Wollaston, 1864 ± Pa Agelosus) O. al®erii (Bernhauer, 1925) ± Pa O. auroguttatus (Cameron, 1932) ± Pa, Or ± Subgenus Matidus Motschulsky, 1860 (stat. (comb. nov.) (from Nudabemus) nov.) O. baronii (Coiffait, 1982) ± Pa ± (comb. nov.) (from Nudabemus) O. albanicus (J. MuÈller, 1943) ± Pa O. caerulescens (Coiffait, 1982) ± Pa ± (comb. O. alpestris (Erichson, 1840) ± Pa nov.) (from Nudabemus) O. bernhaueri (J. MuÈller, 1925) ± Pa O. cameroni (nom. nov.) O. biharicus (J. MuÈller, 1926) ± Pa O. bimaculatus (Cameron, 1932) (nec Schrank, O. brenskei (Reitter, 1884) ± Pa 1798; nec Gravenhorst, 1802; nec Mar- O. bisetosus (J. MuÈller, 1943) (as form of O. sham, 1802) (comb. nov.) (from Staphy- brenskei)(syn. nov.) linus) O. holosetosus (J. MuÈller, 1943) (as subsp. of O. curtipennis Motschulsky, 1849 ± Pa O. brenskei)(syn. nov.) O. integer Abeille, 1900 O. brevipennis (Heer, 1839) ± Pa O. gridellii (J. MuÈller, 1924) O. brevipennis brevipennis (Heer, 1839) O. libanoticus (J. MuÈller, 1950) (as subsp. of O. O. brevipennis pseudoalpestris (J. MuÈller, gridellii)(syn. nov.) 1926) AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_43 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 43

O. brunnipes (Fabricius, 1781) ± Pa O. pedemontanus pedemontanus (J. MuÈller, O. brunnipes brunnipes (Fabricius, 1781) 1924) O. brunnipes intermedius (J. MuÈller, 1911) O. pedemontanus pyrenaeus (J. MuÈller, O. caroli (Jarrige, 1943) ± Pa 1924) O. cerceticus Coiffait, 1964 ± Pa O. pedemontanus cantabricus (J. MuÈller, O. chevrolatii (Baudi, 1848) ± Pa 1926) O. alsaticus Coiffait, 1964 (as subsp. of O. O. ponticus Smetana, 1968 ± Pa chevrolatii)(syn. nov.) O. pullus Hochhuth, 1849 ± Pa O. pseudobrevipennis Coiffait, 1964 (as subsp. O. quadraticeps (MeÂneÂtrieÂs, 1832) ± Pa of O. chevrolatii)(syn. nov.) O. reimoseri (Bernhauer, 1906) ± Pa O. vesubiensis Coiffait, 1964 (as subsp. of O. O. simulator Eppelsheim, 1878 ± Pa chevrolatii)(syn. nov.) O. solarii (J. MuÈller, 1923) ± Pa O. zimmermanni Korge, 1965 (as subsp. of O. O. syriacus Baudi, 1848 ± Pa chevrolatii)(syn. nov.) O. syriacus syriacus Baudi, 1848 O. coreanus (J. MuÈller, 1925) ± Pa O. syriacus primigenius (J. MuÈller, 1923) O. deuvei Coiffait, 1978 ± Pa O. tenebricosus (Gravenhorst, 1846) ± Pa O. for®cularius (Motschulsky, 1860) ± Pa O. depolii (J. MuÈller, 1924) O. korgei Smetana, 1965 O. torvus Smetana, 1965 ± Pa O. frater Smetana, 1965 ± Pa O. trapezensis Coiffait, 1964 ± Pa O. heinzi Smetana, 1965 ± Pa O. turcicus (Bernhauer, 1923) ± Pa O. hissaricus DvorÏaÂk, 1984 ± Pa O. hochhuthi Eppelsheim, 1878 ± Pa Subgenus Pseudocypus Mulsant and Rey, 1876 O. italicus (Aragona, 1830) ± Pa (stat. nov.) O. italicus (Erichson, 1840) (syn. nov.) (Protocypus J. MuÈller, 1923) O. garganicus Fiori, 1894 (syn. nov.) (Atlantogoerius Coiffait, 1956) O. silensis Fiori, 1894 (as variety of O. italicus) (Fortunocypus Coiffait, 1964) (syn. nov.) (Nudabemus Coiffait, 1982) O. mirtensis J. MuÈller, 1943 (as subsp. of O. O. addendus (Lindberg, 1953) ± Pa ± (comb. italicus)(syn. nov.) nov.) (from Pseudocypus) O. khnzoriani Coiffait, 1967 ± Pa O. aeneocephalus (DeGeer, 1774) ± Pa, Ne ± O. kuntzeni (J. MuÈller, 1926) ± Pa (comb. nov.) (from Pseudocypus) O. longimanus Smetana, 1965 ± Pa O. aereus Cameron, 1928 ± Pa O. macrocephalus (Gravenhorst, 1802) ± Pa O. aethiops (Waltl, 1835) ± Pa ± (comb. nov.) O. manceps Smetana, 1965 ± Pa (from Pseudocypus) O. matilei Jarrige, 1971 ± Pa O. luigionii (J. MuÈller, 1926) (as subsp. of O. O. iranicus (Smetana, 1971) aethiops)(comb. nov., syn. nov.) (from O. megalocephalus (Nordmann, 1837) ± Pa Pseudocypus) O. milleri Quedenfeldt, 1882 ± Pa O. almensis Coiffait, 1967 ± Pa O. milleri milleri Quedenfeldt, 1882 O. angustulus Eppelsheim, 1888 ± Pa O. milleri robustus Coiffait, 1964 O. anophthalmicus (HernandeÂz and Aguiar, 1988) O. nitens (Schrank, 1781) (stat. nov.) ±Pa±(comb. nov.) (from Pseudocypus) O. similis (Fabricius, 1792) (misidenti®cation, O. apterus (Scheerpeltz, 1976) ± Pa ± (comb. nec Paykull, 1789) nov.) (from Staphylinus) O. nero (Faldermann, 1835) (syn. nov.) O. canariensis Gemminger and Harold, 1868 ± Pa O. nitens nitens (Schrank, 1781) ±(comb. nov.) (from Pseudocypus) O. nitens semialatus (J. MuÈller, 1904) O. densissimus (Bernhauer, 1933) ± Pa ± (comb. O. nitens grigiensis (Reitter, 1918) nov.) (from Staphylinus) O. nitens ochropus (J. MuÈller, 1950) O. dorsalis Sharp, 1889 ± Pa O. nubigena Smetana, 1965 ± Pa O. eppelsheimi Reitter, 1887 ± Pa ± (comb. nov.) O. ormayi (Reitter, 1887) ± Pa (from Pseudocypus) O. ottomanus Fauvel, 1900 ± Pa O. excisus (J. MuÈller, 1950) ± Pa ± (comb. nov.) O. pedemontanus (J. MuÈller, 1924) ± Pa (from Pseudocypus) AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_44 Allen Press • DTPro System File # 01cc

44 AMERICAN MUSEUM NOVITATES NO. 3287

O. libanicus Coiffait, 1956 (as subsp. of O. pi- O. montanus (Cameron, 1942) ± Pa ± (comb. cipennis)(comb. nov.) (from Pseudocy- nov.) (from Staphylinus) pus) O. mus (BrulleÂ, 1832) ± Pa ± (comb. nov.) (from O. fortunatarum Wollaston, 1871 ± Pa ± (comb. Pseudocypus) nov.) (from Pseudocypus) O. transadriaticus (J. MuÈller, 1926) (as subsp. O. fujiensis Naomi, 1992 ± Pa of O. mus)(comb. nov., syn. nov.) (from O. fulvipennis Erichson, 1840 ± Pa ± (comb. nov.) Pseudocypus) (from Pseudocypus) O. tauricus (J. MuÈller, 1932) (as subsp. of O. O. fulvotomentosus Eppelsheim, 1889 ± Pa mus)(comb. nov.) O. hauseri (Bernhauer, 1933) (comb. nov., syn. O. nemotoi Naomi, 1992 ± Pa nov.) (from Staphylinus) O. nepalicus (Coiffait, 1981) ± Pa ± (comb. nov.) O. fuscatoides (Coiffait, 1964) ± Pa ± (comb. (from Pseudocypus) nov.) (from Pseudocypus) O. nigroaeneus Sharp, 1889 ± Pa O. fuscatus (Gravenhorst, 1802) ± Pa ± (comb. O. rambouseki (J. MuÈller, 1925) nov.) (from Pseudocypus) O. obscuroaeneus Fairmaire, 1852 ± Pa ± (comb. O. abbreviatus (Motschulsky, 1858) (comb. nov.) (from Pseudocypus) nov.) (from Pseudocypus) O. obscuroaeneus obscuroaeneus Fairmaire, O. fuscoaeneus Solsky, 1871 ± Pa ± (comb. nov.) 1852 (from Pseudocypus) O. algiricus (J. MuÈller, 1932) (as subsp. of O. graeseri Eppelsheim, 1887 ± Pa ± (comb. O. obscuroaeneus)(comb. nov.) nov.) (from Pseudocypus) (from Pseudocypus) O. graeseri graeseri Eppelsheim, 1887 O. atlasicus (Coiffait, 1974) (as subsp. of O. graeseri brunneopubens (J. MuÈller, 1926) O. obscuroaeneus)(comb. nov., syn. nov.) (from Pseudocypus) (comb. nov.) (from Pseudocypus) O. horsti (Coiffait, 1974) (as subsp. of O. O. hagai Naomi, 1992 ± Pa obscuroaeneus)(comb. nov., syn. O. hakusanus Naomi, 1992 ± Pa nov.) (from Pseudocypus) O. helleni (J. MuÈller, 1926) ± Pa ± (comb. nov.) O. obscuroaeneus schatzmayri (J. MuÈller, (from Pseudocypus) 1923) (comb. nov.) (from Pseudocypus) O. almorensis (Cameron, 1932) ± Pa ± (comb. O. ohtsukaorum Naomi, 1992 ± Pa nov., syn. nov.) (from Pseudocypus) O. orientis (nom. nov.)±Pa O. himalayicus (Cameron, 1935) ± Pa ± (comb. O. orientalis (Bernhauer and Schubert, 1914) nov., syn. nov.) (from Staphylinus) (nec Motschulsky, 1858) (comb. nov., O. hidanus Watanabe, 1987 ± Pa syn. nov.) (from Pseudocypus) O. himalaycus (Coiffait, 1982) ± Pa ± (comb. O. picipennis (Fabricius, 1792) ± Pa ± (comb. nov.) (from Pseudocypus) nov.) (from Pseudocypus) O. impennis (Fauvel, 1882) ± Pa ± (comb. nov.) O. picipennis picipennis (Fabricius, 1792) (from Staphylinus) O. abbruzzensis (J. MuÈller, 1926) (as O. inexspectatus Eppelsheim, 1887 ± Pa ± (comb. subsp. of O. picipennis)(comb. nov.) nov.) (from Pseudocypus) (from Pseudocypus) O. iyonus Naomi, 1992 ± Pa O. aprutianus (J. MuÈller, 1926) (as subsp. O. izayoi Naomi, 1992 ± Pa of O. picipennis)(comb. nov.) (from O. jeannei Coiffait, 1980 ± Pa Pseudocypus) O. kendappa Naomi, 1992 ± Pa O. herzegovinensis (J. MuÈller, 1932) (as O. kiimontanus Naomi, 1992 ± Pa subsp. of O. picipennis)(comb. nov., O. kisonus Naomi, 1992 ± Pa syn. nov.) (from Pseudocypus) O. lewisius Sharp, 1874 ± Pa O. picipennis graecus (Scheerpeltz, 1958) O. kobensis Cameron, 1930 (syn. nov.) (comb. nov., syn. nov.) (from Pseu- O. mateui (Coiffait, 1954) ± Pa ± (comb. nov., docypus) stat. nov.) (from Pseudocypus) O. anatolicus (Coiffait, 1964) (as subsp. of O. auricomus (Lindberg, 1953) (nec BrulleÂ, O. picipennis)(comb. nov.) (from 1842; nec Cameron, 1929) (comb. nov., Pseudocypus) syn. nov.) (from Pseudocypus) O. andorranus (Coiffait, 1964) (as subsp. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_45 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 45

of O. picipennis)(comb. nov.) (from O. angustatus Stephens, 1832 (comb. nov., syn. Pseudocypus) nov.) (from Pseudocypus) O. pindensis (Coiffait, 1964) (as subsp. of O. con®nis Stephens, 1932 (comb. nov., syn. O. picipennis)(comb. nov.) (from nov.) (from Pseudocypus) Pseudocypus) O. rossii (Jarrige, 1954) (comb. nov., syn. O. bulgaricus (Coiffait, 1970) (as subsp. of nov.) (from Pseudocypus) O. picipennis)(comb. nov., syn. O. fulvicupreus (Coiffait, 1956) (comb. nov., nov.) (from Pseudocypus) syn. nov.) (from Pseudocypus) O. rodopensis (Coiffait, 1971) (as subsp. O. sericeomicans (Bernhauer, 1931) ± Pa ± of O. picipennis)(comb. nov., syn. (comb. nov.) (from Staphylinus) nov.) (from Pseudocypus) O. serotinus (AdaÂm, 1992) ± Pa ± (comb. nov.) O. ponticus (Coiffait, 1978) (as subsp. of (from Pseudocypus) O. picipennis)(comb. nov., syn. O. smetanai Naomi, 1992 ± Pa nov.) (from Pseudocypus) O. subaenescens Wollaston, 1864 ± Pa ± (comb. O. picipennis altaiensis (J. MuÈller, 1926) nov.) (from Pseudocypus) (comb. nov.) (from Pseudocypus) O. svozili DvorÏaÂk, 1984 ± Pa O. picipennis barbarus (J. MuÈller, 1926) O. sylvaticus Wollaston, 1865 ± Pa ± (comb. (comb. nov.) (from Pseudocypus) nov.) (from Pseudocypus) O. marocanus (Coiffait, 1964) (as subsp. O. umbricola Wollaston, 1864 ± Pa ± (comb. of O. picipennis)(comb. nov., syn. nov.) (from Pseudocypus) nov.) (from Pseudocypus) O. yamato Naomi, 1992 ± Pa O. picipennis caucasicus (J. MuÈller, 1926) O. yoroi Naomi, 1992 ± Pa (comb. nov.) (from Pseudocypus) O. yuinus Naomi, 1992 ± Pa O. picipennis fallaciosus (J. MuÈller, 1926) Subgenus Aulacocypus J. MuÈller, 1925 (comb. nov.) (from Pseudocypus) O. maritimus Coiffait, 1956 (as subsp. of O. gloriosus Sharp, 1874 ± Pa O. picipennis)(comb. nov.) O. kansuensis (Bernhauer, 1933) (comb. nov.) O. picipennis nevadensis (J. MuÈller, 1926) (from Staphylinus) (see comment under Ocypus above) (comb. nov.) (from Pseudocypus) O. parvulus Sharp, 1874 ± Pa O. asturicus Coiffait, 1956 (as subsp. of O. O. subtilis Tikhomirova, 1973 ± Pa picipennis)(comb. nov.) (from Pseu- docypus) AGELOSUS SHARP, 1889 O. cerdanicus Coiffait, 1956 (as subsp. of (Apecholinus Bernhauer, 1933) O. picipennis)(comb. nov.) (from A. carinatus (Sharp, 1874) ± Pa Pseudocypus) A. chinensis Li, 1992 ± Pa O. gallicus Coiffait, 1956 (as subsp. of O. A. fraternus (Fairmaire, 1891) ± Pa picipennis)(comb. nov.) (from Pseu- A. kaiseri (Bernhauer, 1933) ± Pa docypus) A. ohkurai Hayashi, 1978 ± Pa O. teruelensis (Coiffait, 1976) (as subsp. A. unicolor Naomi, 1983 ± Pa of O. picipennis)(comb. nov., syn. A. unicolor unicolor Naomi, 1983 nov.) (from Pseudocypus) A. unicolor masaoi Hayashi, 1991 O. plagiicollis (Fairmaire, 1891) ± Pa ± (comb. nov.) (from Staphylinus) WASMANNELLUS BERNHAUER, 1920 O. sabi Naomi, 1992 ± Pa W. stevensi (Cameron, 1932) ± Pa ± (comb. nov.) O. scutiger Sharp, 1889 ± Pa (from Staphylinus) O. semenowi Reitter, 1887 ± Pa W. tristis Bernhauer, 1920 ± Pa O. septentrionalis Watanabe, 1984 ± Pa O. sericeicollis (MeÂneÂtrieÂs, 1832) ± Pa ± (comb. TASGIUS STEPHENS, 1829 (stat. nov.) nov., stat. nov.) (from Pseudocypus) Subgenus Tasgius Stephens, 1829 O. cupreus (Rossi, 1790) (nec Fourcroy, 1785) (Pseudotasgius Seidlitz, 1891) (comb. nov., syn. nov.) (from Pseudocy- (Paratasgius Jarrige, 1952) pus) T. amiculus (J. MuÈller, 1925) ± Pa AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_46 Allen Press • DTPro System File # 01cc

46 AMERICAN MUSEUM NOVITATES NO. 3287

T. ater (Gravenhorst, 1802) ± Pa, Ne T. ensifer (J. MuÈller, 1932) (comb. nov.) (from T. atronitidus (Reitter, 1909) ± Pa Alapsodus) T. bicolor (Cameron, 1944) ± Pa ± (comb. nov.) T. herculeanus (Coiffait, 1964) ± Pa ± (comb. (from Staphylinus) nov.) (from Alapsodus) T. deuvei Coiffait, 1984 ± Pa T. inderiensis (Motschulsky, 1845) ± Pa T. pedator (Gravenhorst, 1802) ± Pa T. limbifrons (Hochhuth, 1849) ± Pa ± (comb. T. peyerimhof® (J. MuÈller, 1926) ± Pa nov.) (from Alapsodus) T. planipennis (AubeÂ, 1842) ± Pa T. lusitanicus (J. MuÈller, 1943) ± Pa ± (comb. T. praetorius (Bernhauer, 1915) ± Pa ± (comb. nov.) (from Alapsodus) nov.) (from Ocypus) T. maderae (Jarrige, 1943) ± Pa ± (comb. nov.) T. sikkimensis (Bernhauer, 1920) ± Pa (from Alapsodus) T. transversiceps (Luze, 1904) ± Pa T. melanarius (Heer, 1839) ± Pa, Ne ± (comb. nov.) (from Alapsodus) Subgenus Rayacheila Motschulsky, 1845 (stat. T. melanarius melanarius (Heer, 1839) nov.) T. melanarius sahlbergi (J. MuÈller, 1926) ± (Anodus Nordmann, 1837; nec Spix, 1829) Pa ± (comb. nov.) (from Alapsodus) (Alapsodus Tottenham, 1939) T. messor (Nordmann, 1837) ± Pa ± (comb. nov.) (Allocypus Coiffait, 1964) (from Alapsodus) (Metocypus Coiffait, 1964) T. minax (Mulsant and Rey, 1861) ± Pa ± (comb. (Paralapsodus Coiffait, 1974) nov.) (from Alapsodus) T. morsitans (Rossi, 1790) ± Pa ± (comb. nov.) T. algericus (Coiffait, 1964) ± Pa ± (comb. nov.) (from Alapsodus) (from Alapsodus) T. compressus (Marsham, 1802) (comb. nov.) T. amoenanus (nom. nov.)±Pa (from Alapsodus) T. amoenus (Reitter, 1909) (nec Olivier, 1795) T. fulvipes (Motschulsky, 1858) (comb. nov.) (comb. nov., syn. nov.) (from Alapsodus) (from Alapsodus) T. arrowi (J. MuÈller, 1932) ± Pa ± (comb. nov.) T. myops (J. MuÈller, 1925) ± Pa ± (comb. nov.) (from Alapsodus) (from Alapsodus) T. bellicosus (Fairmaire, 1855) ± Pa ± (comb. T. nigrinus (Lucas, 1849) ± Pa ± (comb. nov.) nov.) (from Alapsodus) (from Alapsodus) T. bucharicus (Bernhauer, 1912) ± Pa ± (comb. T. olympicus (Baudi, 1857) ± Pa ± (comb. nov.) nov.) (from Alapsodus) (from Alapsodus) T. bulgaricus (Coiffait, 1971) ± Pa ± (comb. nov.) T. pendjabensis (Bernhauer, 1915) ± Pa ± (comb. (from Alapsodus) nov.) (from Staphylinus) T. caspius (Bernhauer, 1906) ± Pa ± (comb. nov.) T. pliginskii (Bernhauer, 1915) ± Pa ± (comb. (from Alapsodus) nov.) (from Alapsodus) T. depressus (Hochhuth, 1849) ± Pa ± (comb. T. rubripennis (Reiche and Saulcy, 1856) ± Pa ± nov.) (from Alapsodus) (comb. nov.) (from Alapsodus) T. doriae (J. MuÈller, 1932) ± Pa ± (comb. nov.) T. solskyi (Fauvel, 1875) ± Pa ± (comb. nov.) (from Alapsodus) (from Alapsodus) T. eppelsheimianus (Jakobson, 1909) ± Pa ± T. solskyi solskyi (Fauvel, 1875) (comb. nov.) (from Alapsodus) (see com- T. solskyi bosphoranus (J. MuÈller, 1926) ment under Tasgius above for var. obscu- (comb. nov.) (from Alapsodus) ripes Bernhauer, 1900) T. tricinctus (Aragona, 1830) ± Pa ± (comb. nov.) T. falcifer (Nordmann, 1837) ± Pa ± (comb. nov.) (from Alapsodus) (from Alapsodus) T. tricolor (Coiffait, 1974) ± Pa ± (comb. nov.) T. falcifer falcifer (Nordmann, 1837) (from Alapsodus) T. falcifer aliquoi (Bordoni, 1976) (comb. T. walkeri (Fauvel, 1898) ± Pa ± (comb. nov.) nov.) (from Alapsodus) T. globulifer (Geoffroy, 1785) ± Pa ± (comb. T. winkleri (Bernhauer, 1906) ± Pa, Ne ± (comb. nov.) (from Alapsodus) nov.) (from Alapsodus) T. gracilicornis (Hochhuth, 1849) ± Pa ± (comb. nov.) (from Ocypus) The following Palearctic species of Staphylinus AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_47 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 47

sensu lato, unknown to us at present, cannot be 1917. Zur Staphylinidenfauna von Nordamer- placed with certainty in the new concepts of gen- ika. Koleopt. Rdsch. 6: 1±4. era and/or subgenera. They are listed alphabeti- 1929. Zur Staphylinidenfauna des chinesisch- cally (by species) in the original combination: en Reiches. Entomol. Nachrbl. 3: 109± 112. Ocypus (Goerius) aenescens Eppelsheim, 1889 1931. Eine neue Subtribus der Quediini Ocypus altissimus Coiffait, 1977 (Staphylinidae). Koleopt. Rdsch. 17: Ocypus ambiguus Coiffait, 1964 84. Ocypus bhutanicus Coiffait, 1977 1933. Neuheiten der chinesischen Staphylini- denfauna. Wiener Entomol. Ztg. 50: Ocypus bureschi Coiffait, 1971 25± 48. Staphylinus (Goerius) caeruleoides Scheerpeltz, 1934. Siebenter Beitrag zur Staphylinidenfau- 1976 na Chinas. Entomol. Nachrbl. 8: 1±20. Staphylinus (Goerius) coeruleus Cameron, 1932 1938. Zur Staphylinidenfauna von China und Ocypus cyclopus Peyron, 1858 Japan. (10. Beitr.). Ent. Nachrbl. 12: Staphylinus (Goerius) duplicatus J. MuÈller, 1943 97±109. Staphylinus griseipennis Fairmaire, 1889 1943. Neue Staphyliniden der palaÈarktischen Ocypus (Neotasgius) harbinensis Li and Chen, Fauna. Koleopt. Rundsch. 29: 71±76. 1993 Bernhauer, M., and K. Schubert Ocypus (Ocypus) japonicus Sawada, 1965 1914. Staphylinidae IV (pars 57). In W. Junk and S. Schenkling (eds.), Coleoptero- Parocypus krejcii Coiffait, 1984 rum catalogus, vol. V: 289±408. Berlin: Ocypus (Neotasgius) liaoningensis Li and Chen, W. Junk, 988 pp. 1993 BohaÂcÏ, J. Staphylinus (Abemus) marginatus Cameron, 1944 1988. New and little known Staphylinidae Ocypus murgulensis Coiffait, 1978 (Coleoptera, Staphylinidae). Ent. Ocypus nudicollis Coiffait, 1977 Obozr. 67: 549±557 [In Russian]. Ocypus opacus Roth, 1851 BrulleÂ, A. Staphylinus (Goerius) pecoudi Jarrige, 1954 1839. Entomologie. In P. B. Webb and S. Ber- Ocypus? queinneci Coiffait, 1984 thelot, Histoire naturelle des Iles Ca- Ocypus sikkimensis Coiffait, 1985 naries, vol. 2, pt. 2. Paris: Mellier, 119 Staphylinus (Ocypus) suturalis Matsumura, 1911 pp., 8 pls. 1845. Insectes. In A. d'Orbigny, Voyage dans Ocypus testaceipes Fairmaire, 1887 l'AmeÂrique meÂridionale (1837±43), tome IV, 2e partie. Paris: Bertrand, 222 REFERENCES pp., 32 pls. Cameron, M. Only references mentioned in the discussions 1929. New species of Staphylinidae from the in the text are included. References associated Belgian Congo. Rev. Zool. Bot. Afr. only with citations of scienti®c names do not ap- 18: 1±10. pear here, unless they were consulted for taxo- 1930. New species of Staphylinidae from Ja- nomic or distribution information. pan. Entomol. Mon. Mag. 67: 181±208. Aragona, L. A. 1932. The fauna of British India, including 1830. De quibusdam insectis Italiae novis aut Ceylon and Burma. Coleoptera. Staph- rarioribus. Ticini Reg.: 31 pp. ylinidae Ð vol. III. London: Taylor & Baudi, F. Francis, xiii ϩ 443 pp., 4 pls. 1869. Coleopterorum messis in insula Cypro 1935. Staphylinidae (Col.). In P. C. Visser et Asia Minore ab Eugenio Truqui con- (ed.), Wissenschaftliche Ergebnisse der gregatae recensitio: de Europaeis notis Niederlandischen Expedition in den quibusdam additis. Berliner Entomol. Karakorum und die angrenzenden Ge- Z. 13: 369±418. biete 1922, 1925 und 1929±30, vol. 1. Bernhauer, M. Leipzig: F. A. Brockhaus, xviii ϩ 499 1900. Neue Staphyliniden (Coleopt.) aus dem pp., 1 map, 8 pls. Kaukasus und den angrenzenden LaÈn- 1944. Descriptions of new Staphylinidae (Co- dern. Wiener Entomol. Ztg. 19: 46±55. leoptera). Proc. R. Entomol. Soc. Lon- 1911. Zur Staphylinidenfauna Ostindiens und don (B) 13: 11±15. der Sundainseln. Entomol. BlaÈtter 7: Coiffait, H. 55±62, 86±93. 1954. Contribution aÁ la connaissance des Sta- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_48 Allen Press • DTPro System File # 01cc

48 AMERICAN MUSEUM NOVITATES NO. 3287

phylinides des Canaries: reÂcoltes de J. 1839b. Die KaÈfer der Mark Brandenburg: 385± Mateu. Arch. Inst. Aclimat. Almeria 2: 740. Berlin: F. H. Morin. 161±177. 1840. Genera et species staphylinorum insec- 1956. Les Staphylinus et les genres voisins de torum coleopterorum familiae: 401± France et des reÂgions voisines. Essai de 945. Berlin: F. H. Morin. PaleÂobiogeÂographie. MeÂm. Mus. Nat. Fabricius, J. C. Hist. Nat. Paris (n. ser.), A, Zool. 8: 1775. Systema entomologiae, sistens insecto- 177±224. rum classes, ordines, genera, species, 1964. Note sur les Ocypus (sensu lato) avec adiectis synonymis, locis, descriptioni- description de formes nouvelles. Bull. bus, observationibus. Flensburgi et Soc. Hist. Nat. Toulouse 99: 81±106. Lipsiae: Libraria Kortii. 30 ϩ 832 pp. 1970. Staphylinidae nouveaux ou mal connus 1792. Entomologia systematica emendata et de la reÂgion paleÂarctique occidentale. aucta, secundum classes, ordines, gen- Ibid. 106: 99±111. era, species adjectis synonymis, locis, 1971. Staphylinides nouveaux ou mal connus observationibus, descriptionibus, tom. de Bulgarie. Nouv. Rev. Entomol. 1: I, pars I. Hafniae: C. G. Proft, xx ϩ 330 279± 286. pp. 1974. ColeÂopteÁres Staphylinidae de la reÂgion Faldermann, F. paleÂarctique occidentale. II. Sous-fam- 1835. Fauna Entomologica Transcaucasica, ille Staphilininae. Tribus Philonthini et vol. 1. Nouv. MeÂm. Soc. Nat. Moscou Staphylinini. Suppl. Nouv. Rev. Ento- 4: 1±314, 10 pls. mol., tome IV, fasc. 4, 593 pp., Tou- Fauvel, A. louse. 1874. In Faune gallo-rheÂnane ou species des 1982. Contribution aÁ la connaissance des Sta- insectes qui habitent la France, la Bel- phylinides de l'Himalaya (NeÂpal, La- gique, la Hollande, le Luxembourg, la dakh, Cachemire) (Insecta: Coleoptera: Prusse RheÂnane, le Nassau et le Valais Staphylinidae). Senckenbergiana Biol. avec tableaux synoptiques et planches 62: 21±179. graveÂes. ColeÂopteÁres, tome 2. III. Sta- Drugmand, D. phylinides: 391±544, CaeÈn: [s. n.], 4 1998. SysteÂmatique et biogeÂographie des pls. sous-espeÁces europeÂennes d'Ocypus Frank, J. H. ophthalmicus (Scopoli, 1763) (Coleop- 1979. Larval morphology and the classi®ca- tera: Staphylininae). Ann. Soc. Ento- tion of Staphylinus sensu lato (Col., mol. Fr. (n. ser.) 34: 45±61. Staphylinidae). Entomologist's Mon. DvorÏaÂk, M. Mag. 114: 235±238. 1984. Zwei neue Ocypus-Arten aus Mittela- Ganglbauer, L. sien und einige Bemerkungen zu den 1895. Die KaÈfer von Mitteleuropa. Die KaÈfer Angaben uÈber KoÈrperlaÈngen der Fami- der oÈsterreich-ungarischen Monarchie, lie Staphylinidae. Faun. Abhandl. Deutschlands, der Schweiz, sowie des Staatl. Mus. Tierkunde Dresden 12: franzoÈsischen und italianischen Alpen- 59±68. gebietes. Band 2. Familienreihe Sta- Eppelsheim, E. phylinoidea. 1. Theil. Staphylinidae, 1878. Neue Staphylinen. Stett. Entomol. Ztg. Pselaphidae. Wien: Carl Gerod's Sohn, 39: 417±424. vi ϩ 880 pp., 38 ®gs. 1884. Diagnosen neuer Staphyliniden aus Geoffroy, M. dem Kaukasus und aus Lenkoran. Verh. 1785. In A. F. Fourcroy, Entomologia Pari- Naturf. Ver. BruÈnn 22: 11±16. siensis; sive catalogus insectorum quae 1887. Synonymische Bemerkungen uÈber eu- in agro Parisiensi reperiuntur; secun- ropaÈische Staphylinen. Dtsch. Entomol. dum methodum Geoffraeanam in sec- Z. 31: 430±432. tiones, genera & species distributus: cui 1889. Insecta, a Cl. G. N. Potanin in China et addita sunt nomina trivialia & fere tre- in Mongolia novissime lecta. V. Neue centa novae species, pars 1. Parisiis: [s. Staphylinen. Horae Soc. Ent. Rossicae n.], viii ϩ 231 pp. 23: 169±184. Goeze, J.A.E. Erichson, W. F. 1777. Entomologische BeytraÈge zu des Ritter 1839a. Genera et species staphylinorum insec- Linne zwoÈlften Ausgabe des Natursys- torum coleopterorum familiae: 1±400. tems, 1. Theil. Leipzig: Weidmann, xvi Berlin: F. H. Morin. ϩ 736 pp. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_49 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 49

Gravenhorst, J.L.C. Leach, W. E. 1802. Coleoptera microptera Brunsvicensia 1819. [New genera.] In G. Samouelle. The nec non exoticorum quotquot exstant in entomologist's useful compendium, or collectionibus entomologorum bruns- an introduction to the knowledge of vicensium in genera familias et species British insects, comprising the best distribuit. Brunsvigae: Carolus Rei- means of obtaining and preserving chard, lxvi ϩ 206 pp. them, and a description of the apparatus Gusarov, V. generally used; together with the gen- 1992. New and little known Palaearctic era of LinneÂ, and the modern method Staphylinids (Coleoptera, Staphylini- of arranging the classes Crustacea, dae). Entomol. Obozr. 71: 775±788 [In Myriapoda, spiders, mites and insects, Russian]. from their af®nities and structure, ac- Hayashi, Y. cording to the views of Dr. Leach. Also 1991. Studies on Staphylinidae from Japan. an explanation of the terms used in en- III. Entomol. Rev. Japan 46: 179±185. tomology; a calendar of the times of 1993. Studies on the Asian Staphylininae, I appearance and usual situations of (Coleoptera, Staphylinidae). Elytra (To- 3,000 species of British insects; with kyo) 21: 281±301. instructions for collecting and ®tting up 1994. Studies on the Asian Staphylininae objects for the microscope. London: (Coleoptera, Staphylinidae). II. On the Longman, Hurst, Rees, Orme, Brown characteristics of the genus Philonthus & Green, 496 pp. Curtis, sensu stricto, with a redescrip- Li, J. tion of Philonthus splendens (Fabri- 1992. The Coleoptera fauna of northeast Chi- cius). Ibid. 22: 115±131. na. Jilin: Jilin Edu. Publ. House, 205 1995. Studies on Staphylinidae (Coleoptera) pp. from Japan, V. Revision on the type Li, J., and P. Chen specimens of Amichrotus excellens 1993. Studies of fauna and ecogeography of Bernhauer and Staphylinus (Xanthocy- soil animal. Chang Chun: Northeast pus) ganglbauerianus Bernhauer. En- Normal University Press, 265 pp. tomol. Rev. Japan 50: 45±57. Lindberg, H. 1997. Studies on the Asian Staphylininae 1953. Zweiter Beitrag zur Kenntnis der KaÈ- (Coleoptera, Staphylinidae). III. The ferfauna der Kanarischen Inseln. Com- characteristics of the Xanthopygini. El- mentat. Biolog. 13(12): 1±18. ytra (Tokyo) 25: 475±492. Mannerheim, C. G. Hochhuth, H. 1830. PreÂcis d'un nouvel arrangement de la 1849. Die Staphylinen-Fauna des Kaukasus famille des bracheÂlytres de l'ordre des und Transkaukasiens. Bull. Soc. Nat. insectes coleÂopteÁres. MeÂm. Acad. Imp. Moscou 22(1): 18±214. Sci. St. PeÂtersb. 1: 415±501. Sep. St. Jarrige, J. PeÂtersbourg, 87 pp. 1954. Les Staphylinus des õÃles Atlantides Marsham, T. (Col. Staphylinidae). Bull. Soc. Ento- 1802. Entomologia britannica, sistens insecta mol. France 58: 160±164. Britanniae indigena, secundum meth- Kraatz, G. odum Linnaeanam disposita. Tomus I. 1856±58. Staphylinii. Naturgeschichte der In- Coleoptera. Londini: Wilks & Taylor, secten Deutschlands. Erste Abtheilung. xxxi ϩ 549 pp. Coleoptera, 2. Band. Berlin: Nicolai, Melsheimer, F. E. viii ϩ 1080 pp. 1844. Descriptions of new species of Cole- 1859. Die Staphylinen-Fauna von Ostindien, optera of the United States. Proc. Acad. insbesondere der Insel Ceylan. Archiv Nat. Sci. Philadelphia 2: 26±47. Naturgesch. 25: 1±193, 3 pls. Sep. Ber- Moore, I., and E. F. Legner lin: Nikolai, 196 pp., 3 pls. 1979. An illustrated guide to the genera of the Latreille, P. A. Staphylinidae of America north of 1806. Genera crustaceorum et insectorum se- Mexico exclusive of Aleocharinae (Co- cundum ordinem naturalem in familias leoptera). Berkeley: Univ. California disposita, iconibus exemplisque plu- Publ. 4093, 332 pp. rimis explicata. Tomus primus. Parisiis Motschulsky, V. et Argentorati: Koenig, xviii ϩ 302 pp., 1845. Remarques sur la collection de ColeÂop- 16 pls. teÁres russes. Bull. Soc. Imp. Nat. Mos- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_50 Allen Press • DTPro System File # 01cc

50 AMERICAN MUSEUM NOVITATES NO. 3287

cou 18(1): 3±127 [Staphylinidae on pp. neÂe. ColeÂopteÁres, tome 3. Paris: Lan- 38±42]. neau. [Each genus treated is separately 1858. E tudes Entomologiques VI. Imprim. paginated.] Soc. LitteÂr. Finn. Helsingfors: 112 pp., Outerelo, R., and P. Gamarra 1 pl. 1985. Las familias y geÂneros de los esta®lõÂn- 1860. E numeration des nouvelles espeÁces de idos de la PenõÂnsula IbeÂrica. Claves coleÂopteÁres rapporteÂes de ses voyages. para la identi®cacion de la fauna Es- 3ieÁme article. IV. Staphylinides de Rus- panÄola 10. Madrid: Universidad Com- sie. Bull. Soc. Imper. Nat. Moscou plutense, 139 pp. 33(2): 539±588. Paykull, G. MuÈller, J. 1789. Monographia staphylinorum Sueciae. 1923. Contributo alla conoscenza del genere Upsaliae: J. Edman, xii ϩ 82 pp. Staphylinus L. Boll. Soc. Entomol. Ital. Pilon, N. 55: 135±144. 1991. Note sugli Staphylinini di un ambiente 1924. Secondo contributo alla conoscenza del collinare del Comasco. Boll. Soc. En- genere Staphylinus L. Ibid. 56: 19±40. tomol. Ital. 123: 100±106. 1925. Terzo contributo alla conoscenza del 1998. Atlante faunistico degli Staphylinini it- genere Staphylinus L. Ibid. 57: 40±48. aliani con note sinonimiche (Coleop- 1926a. Quarto contributo alla conoscenza del tera). Memor. Soc. Entomol. Ital. 76: genere Staphylinus L. Ibid. 58: 27±32. 61±129. 1926b. Quinto contributo alla conoscenza del Reitter, E. genere Staphylinus L. Ibid. 58: 41±48. 1909. Fauna Germanica. Die KaÈfer des 1926c. Sesto contributo alla conoscenza del Deutschen Reiches. Nach der analy- genere Staphylinus L. Ibid. 58: 73±75. tischen Methode bearbeitet. II. Band. 1926d. Untersuchungen uÈber europaÈische Sta- Stuttgart: K. G. Lutz, 392 pp., pls. 41± phylinus-Arten. Coleopt. Centralblatt 1: 80. 5±24. Schillhammer, H. 1932. Settimo contributo alla conoscenza del 1998a. Revision of the east Palaearctic and genere Staphylinus L. Boll. Soc. Ento- Oriental species of Philonthus Ste- mol. Ital. 64: 75±88. phens, pt.1. The cyanipennis group 1943. Ottavo contributo alla conoscenza del (Coleoptera: Staphylinidae, Staphylini- genere Staphylinus. Atti Mus. Civ. Stor. nae). Koleopt. Rundschau 68: 101±118. Nat. Trieste 15: 95±109. 1998b. Hybridolinus gen. n. (Insecta: Coleop- MuÈller, O. F. tera: Staphylinidae), a problematic new 1776. Zoologiae Danicae prodromus, seu an- genus from China and Taiwan, with de- imalium Daniae et Norvegiae indigen- scription of seven new species. Ann. arum characteres, nomina, et synonyma Naturh. Mus. Wien 100(B): 145±156. imprimis popularium. Hafniae: Halla- Schrank, F. von Paula geriis, xxxii ϩ 282 pp. 1781. Enumeratio insectorum Austriae indi- Mulsant, E., and C. Rey genorum. Augustae Vindelicorum: 1876. Histoire naturelle des coleÂopteÁres de Klett et Franck, xxiv ϩ 548 pp., 4 pls. France. Tribu des breÂvipennes. Staphy- 1798. Fauna Boica, vol. 1, pt. 2: 293±720. liniens. Annales Soc. Agric. Lyon (5e NuÈrnberg: [s.n.]. seÂr.) 8: 145±856. Scopoli, J. A. Naomi, S.-I. 1772. Observationes zoologicae. Ann. Hist. 1983. Revision of the subtribe Xanthopygina Nat. 5: 75±125. (Coleoptera, Staphylinidae) of Japan Sharp, D. III. Kontyuà (Tokyo) 51: 582±592. 1874. The Staphylinidae of Japan. Trans. R. Nordmann, A. Entomol. Soc. London: 1±103. 1837. Symbolae ad monographiam staphyli- 1889. The Staphylinidae of Japan III. Ann. norum. Petropoli: Ex Academiae Cae- Mag. Nat. Hist. (ser. 6) 3: 28±44, 108± sareae Scientiarum Tomo IV, 167 pp., 121, 249±267, 319±334, 406±419, 2 pls. 463±476. Olivier, A.G. Shibata, Y. 1795. Entomologie, ou histoire naturelle des 1984. Provisional check list of the family insectes, avec leurs caracteÁres geÂneÂ- Staphylinidae of Japan. IV. (Insecta: riques et speÂci®ques, leur description, Coleoptera). Ann. Bull. Nichidai Sanko leur synonymie, et leur ®gure enlumi- 22: 79±141. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_51 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 51

Smetana, A. ing a guide to their classi®cation. Lon- 1965a. Staphylinini und Quediini von Canada don: Baldwin & Cradock, 68 cols. und Alaska (Col., Staphylinidae). Acta Stickney, F. S. Lundensis (sect. II), 13: 1±18. 1923. The head-capsule of Coleoptera. Illi- 1965b. Zur Kenntnis der Staphylinus- und nois Biol. Monogr. 8: 1±105, 26 pls. Ocypus-Arten Nordanatoliens (Coleop- Szujecki, A. tera, Staphylinidae) (66. Beitrag zur 1980. Klucze do oznaczania owadoÂw Polski. Kenntnis der Staphyliniden). Reichen- CzeÎsÂc XIX. ChrzaÎszczeÐColeoptera. bachia 5: 25±46. Zeszyt 24 e. KusakowateÐStaphylini- 1967. Beitrag zur Kenntnis einiger Staphyli- dae. KusakiÐStaphylininae. PanÂ- nus- und Ocypus-Arten Spaniens (Col., stwowe Warszawa: Wydawnictwo Nau- Staphylinidae). (77. Beitrag zur Kennt- kowe, 164 pp. nis der Staphyliniden). Acta Faun. En- Thomson, C. G. 1858. FoÈrsoÈk till uppstaÈllning af Sveriges Sta- tomol. Mus. Nat. Pragae 12: 197±200. phyliner. OÈ fvers. K. VetensAkad. 15: 1971. Zur Kenntnis der Staphylinus- und 27±40. Ocypus-Arten Irans (Coleoptera, Staph- Tottenham, C. E. ylinidae). (88. Beitrag zur Kenntnis der 1939. Some notes on the nomenclature of the Staphyliniden). Acta Ent. Bohemoslov. Staphylinidae (Coleoptera). Proc. R. 68: 263±269. Entomol. Soc. London (B) 8: 224±226. Stephens, J. F. Westwood, J. O. 1829. The nomenclature of British insects; 1827. Observations upon Siagonium quadri- being a compendious list of such spe- corne of Kirby, and the other portions cies as are contained in the Systematic of the Brachelytra. Zool. J. 3: 56±66, 1 Catalogue of British Insects, and form- pl.

INDEX OF SPECIES AND GENERIC SYNONYMS Original combinations, when included, are in parentheses. Current combinations are in brackets. Valid genera and subgenera of Staphylinina studied in detail in this paper are in boldface type. Valid species and generic names are in italic type and synonyms are in roman type.

abbreviatus (Motschulsky, 1858) [ϭ Ocypus af®nis Wollaston, 1864 [Ocypus (Ocypus)], 35, 42 (Pseudocypus) fuscatus (Gravenhorst, Agelosus Sharp, 1889, 3, 7±11, 15±17, 20, 35,45 1802)], 44 ahrendti Wendeler, 1921 [Physetops], 42 abbruzzensis (J. MuÈller, 1926) [ϭ Ocypus (Pseu- Alapsodus Tottenham, 1939 [ϭ Tasgius (Rayach- docypus) picipennis picipennis (Fabricius, eila)], 2, 3, 36, 37, 46 1792)], 44 albanicus (J. MuÈller, 1943) [Ocypus (Matidus)], Abemus Mulsant and Rey, 1876, 3, 6, 8±11, 16, 42 17, 20, 23,39 al®erii (Bernhauer, 1925) [Ocypus (Ocypus)], 42 Acylophorus Nordmann, 1837, 10, 14, 15 algericus (Coiffait, 1964) [Tasgius (Rayacheila)], addendus (Lindberg, 1953) [Ocypus (Pseudocy- 46 pus)], 43 algiricus (J. MuÈller, 1932) [ϭ Ocypus (Pseudo- adonis (Coiffait, 1956) [subsp. of Dinothenarus cypus) obscuroaeneus obscuroaeneus Fair- (Dinothenarus) ¯avocephalus (Goeze, maire, 1852], 44 1777)], 41 Algon Sharp, 1874, 17, 18 aeneicollis (Bernhauer, 1911) [ϭ Platydracus ase- mus (Kraatz, 1859)], 39 aliquoi (Bordoni, 1976) [subsp. of Tasgius (Ray- aeneocephalus (DeGeer, 1774) [Ocypus (Pseudo- acheila) falcifer (Nordmann, 1837)], 46 cypus)], 43 Allocypus Coiffait, 1964 [ϭ Tasgius (Rayachei- aeneoniger (Bernhauer, 1933) [Platydracus], 39 la)], 36, 37, 46 aenescens Eppelsheim, 1889 (Ocypus (Goerius)) Allostenopsis Bernhauer, 1921, 17, 18 [incertae sedis], 47 almensis Coiffait, 1967 [Ocypus (Pseudocypus)], aereus Cameron, 1928 [Ocypus (Pseudocypus)], 43 43 almorensis (Cameron, 1932) [ϭ Ocypus (Pseu- aethiops (Waltl, 1835) [Ocypus (Pseudocypus)], docypus) helleni (J. MuÈller, 1926)], 32, 33, 32, 43 44 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_52 Allen Press • DTPro System File # 01cc

52 AMERICAN MUSEUM NOVITATES NO. 3287

alpestris (Erichson, 1840) [Ocypus (Matidus)], 42 ater (Gravenhorst, 1802) (Staphylinus)[Tasgius alsaticus Coiffait, 1964 [ϭ Ocypus (Matidus) (Tasgius)], 36, 46 chevrolatii (Baudi, 1848)], 43 Atlantogoerius Coiffait, 1956 [ϭ Ocypus (Pseu- altaiensis (J. MuÈller, 1926) [subsp. of Ocypus docypus)], 3, 31, 43 (Pseudocypus) picipennis (Fabricius, 1792), atlasicus (Coiffait, 1974) [ϭ Ocypus (Pseudocy- 45 pus) obscuroaeneus obscuroaeneus Fair- altissimus Coiffait, 1977 (Ocypus) [incertae sed- maire, 1852], 44 is], 47 atripes Bernhauer, 1939 [Naddia], 38 amamiensis Ito, 1982 [Platydracus], 39 atrocyaneus (Fairmaire, 1860) [subsp. of Ocypus ambiguus Coiffait, 1964 (Ocypus) [incertae sedis], (Ocypus) ophthalmicus (Scopoli, 1763)], 42 47 atronitidus (Reitter, 1909) [Tasgius (Tasgius)], 46 Amichrotus Sharp, 1889, 11, 15, 16 Aulacocypus J. MuÈller, 1925 [subgenus of Ocy- amiculus (J. MuÈller, 1925) [Tasgius (Tasgius)], 45 pus], 21, 32, 34, 45 amoenanus, nom. nov. [for Tasgius (Rayacheila) aureofasciatus (Motschulsky, 1862) [Platydra- amoenus (Reitter, 1909)], 37, 46 cus], 39 amoenus Olivier, 1795 (Staphylinus), 37 auricomus (BrulleÂ, 1842) (Staphylinus)[ϭ Glenus amoenus (Reitter, 1909) (Staphylinus)[ϭ Tasgius chrysis (Gravenhorst, 1806)], 33 (Rayacheila) amoenanus, nom. nov.], 37, 46 auricomus Cameron, 1929: 65 [Staphylinus], 33 Anaquedius Casey, 1915, 14, 15 auricomus (Lindberg, 1953) (Staphylinus)[ϭ anatolicus (Coiffait, 1964) [ϭ Ocypus (Pseudo- Ocypus (Pseudocypus) mateui (Coiffait, cypus) picipennis picipennis (Fabricius, 1954)], 33, 44 1792)], 44 auripennis (Kraatz, 1859) [ϭ Platydracus goryi Anchocerus Fauvel, 1905, 10, 14, 15 (Laporte, 1835)], 39 andorranus (Coiffait, 1964) [ϭ Ocypus (Pseudo- auroguttatus (Cameron, 1932) [Ocypus (Ocy- cypus) picipennis picipennis (Fabricius, pus)], 31, 42 1792)], 44 aurosericans (Fairmaire, 1891) [Platydracus], 39 angustatus Stephens, 1832 [ϭ Ocypus (Pseudo- aurosparsus (Fauvel, 1895) [Ontholestes], 38 cypus) sericeicollis (MeÂneÂtrieÂs, 1832)], 35, azoricus (MeÂquignon, 1942) [subsp. of Ocypus 45 (Ocypus) olens (O. MuÈller, 1764)], 42 angustulus Eppelsheim, 1888 [Ocypus (Pseudo- baderlei (Scheerpeltz, 1966) [ϭ Dinothenarus cypus)], 43 (Parabemus) fossor (Scopoli, 1772)], 41 Anisolinina , 7±15,16 badipes (LeConte, 1863) [Dinothenarus (Para- Anisolinus Sharp, 1889, 7, 15, 16 bemus)], 41 Anodus Nordmann, 1837 [ϭ Tasgius (Rayachei- badius (Mannerheim, 1830) [ϭ Platydracus zon- la)], 2, 3, 36, 37, 46 atus (Gravenhorst, 1802)], 41 anophthalmicus (HernandeÂz and Aguiar, 1988) baicalensis (Eppelsheim, 1887) [ϭ Ocypus (Ocy- [Ocypus (Pseudocypus)], 43 pus) ophthalmicus ophthalmicus (Scopoli, Apecholinus Bernhauer, 1933 [ϭ Agelosus], 35, 1763)], 42 45 balcanicus (J. MuÈller, 1923) [subsp. of Ocypus Apostenolinus Bernhauer, 1934, 20, 29, 30, 42 (Ocypus) ophthalmicus (Scopoli, 1763)], 42 aprutianus (J. MuÈller, 1926) [ϭ Ocypus (Pseudo- balearicus (J. MuÈller, 1926) [subsp. of Ocypus cypus) picipennis picipennis (Fabricius, (Ocypus) ophthalmicus (Scopoli, 1763)], 42 1792)], 44 banghaasi (J. MuÈller, 1932) [ϭ Platydracus muel- apterus (Scheerpeltz, 1976) [Ocypus (Pseudocy- lerianus (Scheerpeltz, 1933)], 40 pus)], 43 barbarus (J. MuÈller, 1926) [subsp. of Ocypus arrosus (Eppelsheim, 1890) [Dinothenarus (Par- (Pseudocypus) picipennis (Fabricius, 1792)], abemus)], 41 45 arrowi (J. MuÈller, 1932) [Tasgius (Rayacheila)], baronii (Coiffait, 1982) [Ocypus (Ocypus)], 42 46 beckeri Bernhauer, 1933 [Ascialinus], 29, 42 Ascialinus Bernhauer, 1933, 11, 16, 17, 20, 29, becquarti (Bernhauer, 1938) [Platydracus], 39 42 Beeria Hatch, 1957, 14, 15 asemus (Kraatz, 1859) [Platydracus], 39 bellicosus (Fairmaire, 1855) [Tasgius (Rayachei- assamensis Cameron, 1932 [Naddia], 38 la)], 46 Astrapaeus Gravenhorst, 1802, 15, 36 benoiti Drugmand, 1998 [subsp. of Ocypus (Ocy- asturicus Coiffait, 1956 [ϭ Ocypus (Pseudocypus) pus) ophthalmicus (Scopoli, 1763)], 42 picipennis nevadensis (J. MuÈller, 1926)], 45 bernhaueri (J. MuÈller, 1925) [Ocypus (Matidus)], Atanygnathus Jakobson, 1909, 13 42 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_53 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 53

bhutanicus Coiffait, 1977 (Ocypus) [incertae se- bureschi Coiffait, 1971 (Ocypus) [incertae sedis], dis], 47 47 bicolor (Cameron, 1944) (Staphylinus (Ocypus)) [Tasgius (Tasgius)], 37, 46 caeruleipennis (Mannerheim, 1831) (Philonthus), biharicus (J. MuÈller, 1926) [Ocypus (Matidus)], 8, 13 42 caeruleoides Scheerpeltz, 1976 (Staphylinus bimaculatus (Cameron, 1932) (Staphylinus)[ϭ (Goerius)) [incertae sedis], 47 Ocypus (Ocypus) cameroni, nom. nov.], 33, caerulescens (Coiffait, 1982) (Nudabemus)[Ocy- 42 pus (Ocypus)], 31, 42 bimaculatus (Schrank, 1798) (Staphylinus)[ϭ caesareus Cederhjelm, 1798 [Staphylinus, and as Lordithon trinotatus (Erichson, 1839)], 33 subsp.], 26, 41 birmanus (Fauvel, 1895) [Thoracostrongylus], 39 Ca®us Stephens, 1829, 7, 14 bisetosus (J. MuÈller, 1943) [ϭ Ocypus (Matidus caliginosus (Erichson, 1839) [Platydracus], 39 brenskei (Reitter, 1884)], 42 callistus (Hochhuth, 1849) [Ontholestes], 38 Bisnius Stephens, 1829, 10 cameroni, nom. nov. [for Ocypus (Ocypus) bi- Bolitobiini, 9, 10 maculatus (Cameron, 1932)], 33, 42 Bolitogyrus Chevrolat, 1842, 14 campestris Coiffait, 1977 [Platydracus], 39 bosphoranus (J. MuÈller, 1926) [subsp. of Tasgius canariensis Gemminger and Harold, 1868 [Ocy- pus (Pseudocypus)], 43 (Rayacheila) solskyi (Fauvel, 1875)], 46 cantabricus (J. MuÈller, 1926) [subsp. of Ocypus brachycerus, nom. nov. [for Platydracus (Platyd- (Matidus) pedemontanus (J. MuÈller, 1924)], racus) brevicornis (Weise, 1877)], 24, 39 43 brachypterus (BrulleÂ, 1839) (Staphylinus)[ϭ Pro- capitatus (Bland, 1864) [Dinothenarus (Dinoth- togoerius brullei, nom. nov.], 25, 30, 42 enarus)], 41 brachypterus (Geoffroy, 1785) (Staphylinus) Caranistes Erichson, 1840 [ϭ Naddia], 3, 22 [Aleochara], 25, 30 carinatus (Sharp, 1874) (Goerius)[Agelosus], 35, brachypterus (Kraatz, 1859) (Staphylinus)[ϭ Pla- 45 tydracus (Platydracus) brevipennis, nom. cariniceps Bernhauer, 1934 [Apostenolinus], 29, nov.], 25, 39 30, 42 brachypterus (Marsham, 1802) (Staphylinus) caroli (Jarrige, 1943) (Staphylinus (Goerius)) [Aploderus], 25, 30 [Ocypus (Matidus)], 43 brenskei (Reitter, 1884) [Ocypus (Matidus)], 35, caseyi (Scheerpeltz, 1933) [ϭ Platydracus zona- 42 tus (Gravenhorst, 1802)], 41 brevicornis (Motschulsky, 1862) [Platydracus caspius (Bernhauer, 1906) [Tasgius (Rayacheila)], (Platydracus)], 18, 39 46 brevicornis Weise, 1877 [ϭ Platydracus brachy- catalonicus Coiffait, 1967 [Platydracus], 39 cerus, nom. nov.], 24. 39 caucasicus (J. MuÈller, 1926) [subsp. of Ocypus brevipennis (Heer, 1839) [Ocypus (Matidus), and (Pseudocypus) picipennis (Fabricius, 1792)], as subsp.], 42 45 brevipennis, nom. nov. [for Platydracus brachyp- centralis (Sharp, 1884) [Platydracus], 39 terus Kraatz, 1859], 25, 39 cerceticus Coiffait, 1964 [Ocypus (Matidus)], 43 brevipennis Sharp, 1889 [Miobdelus], 28, 42 cerdanicus Coiffait, 1956 [ϭ Ocypus (Pseudocy- brigitteae Drugmand, 1998 [subsp. of Ocypus pus) picipennis nevadensis (J. MuÈller, (Ocypus) ophthalmicus (Scopoli, 1763)], 42 1926)], 45 brullei, nom. nov. [for Protogoerius brachypterus Chaetodracus J. MuÈller, 1926 [subgenus of Pla- (BrulleÂ, 1839)], 6, 30, 42 tydracus], 11, 18, 24, 25,41 brunneopubens (J. MuÈller, 1926) [subsp. of Ocy- chalcescens (Sharp, 1889) [Platydracus], 39 pus (Pseudocypus) graeseri Eppelsheim, chalcocephalus (Fabricius, 1801) [Platydracus], 1887], 44 39 brunnipes (Fabricius, 1781) [Ocypus (Matidus), chalcopygus (Hochhuth, 1849) [Ontholestes], 38 and as subsp.], 43 chapmani (Bernhauer, 1933) [Platydracus], 39 bucharicus (Bernhauer, 1912) [Tasgius (Rayach- chevrolatii (Baudi, 1848) [Ocypus (Matidus)], 43 eila)], 46 chinensis (Bernhauer, 1914) [Platydracus], 39 bulgaricus (Coiffait, 1970) [ϭ Ocypus (Pseudo- chinensis Bernhauer, 1929 [Naddia], 38 cypus) picipennis picipennis (Fabricius, chinensis Li, 1992 [Agelosus], 45 1792)], 45 chloropterus (Panzer, 1796) [Abemus], 23, 39 bulgaricus (Coiffait, 1971) [Tasgius (Rayachei- choui Smetana, 1992 [Dinothenarus (Dinothena- la)], 46 rus)], 41 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_54 Allen Press • DTPro System File # 01cc

54 AMERICAN MUSEUM NOVITATES NO. 3287

chrysocomus (Mannerheim, 1830) (Staphylinus) depressus (Hochhuth, 1849) [Tasgius (Rayachei- [Dinothenarus (Parabemus)], 26, 41 la)], 46 Cileoporus Campbell, 1995, 10 deuvei Coiffait, 1978 [Ocypus (Matidus)], 43 cingulatus (Gravenhorst, 1802) [Ontholestes], 38 deuvei Coiffait, 1984 [Tasgius (Tasgius)], 46 cinnamopterus (Gravenhorst, 1802) [Platydra- dieckmanni Smetana, 1958 [Ontholestes], 38 cus], 39 dimidiaticornis Gemminger, 1851 [Staphylinus], coeruleipennis Coiffait, 1983 [Platydracus], 39 9, 41 coeruleus Cameron, 1932 (Staphylinus (Goerius)) Dinothenarus Thomson, 1858 [and subgenus], 2, [incertae sedis], 47 6, 8±11, 16, 17, 20, 26,41 coiffaiti Bordoni, 1977 [Dinothenarus (Parabe- doriae (J. MuÈller, 1932) [Tasgius (Rayacheila)], mus)], 41 46 collaris Coiffait, 1977 [Platydracus], 39 dorsalis Sharp, 1889 [Ocypus (Pseudocypus)], 43 comes (LeConte, 1863) [Platydracus], 39 dudgeoni (Cameron, 1932) [Platydracus], 39 compressus (Marsham, 1802) [ϭ Tasgius (Ray- duplicatus J. MuÈller, 1943 (Staphylinus (Goerius)) acheila) morsitans (Rossi, 1790)], 46 [incertae sedis], 47 con®nis (Stephens, 1832) (Goerius)[ϭ Ocypus Dysanellus Bernhauer, 1911, 18 (Pseudocypus) sericeicollis (MeÂneÂtrieÂs, ejulans (Tottenham, 1939) [ϭ Platydracus cali- 1832)], 35, 45 ginosus (Erichson, 1839)], 39 consularis (Bernhauer, 1915) [Platydracus], 39 Elmas Blackwelder, 1952, 8, 17, 18 coreanus (J. MuÈller, 1925) [Ocypus (Matidus)], 43 Emus Leach, 1819, 2, 7±11, 16±18, 22,38 corporaali Sainte Claire Deville, 1927 [subsp. of ensifer (J. MuÈller, 1932) [ϭ Tasgius (Rayacheila) Staphylinus caesareus Cederhjelm, 1798], 41 gracilicornis (Hochhuth, 1849)], 46 costatus (Fauvel, 1895) [Thoracostrongylus], 39 eppelsheimi Reitter, 1887 [Ocypus (Pseudocy- Craspedomerina, 12, 14 pus)], 43 Craspedomerus Bernhauer, 1911, 13, 14 eppelsheimianus (Jakobson, 1909) [Tasgius (Ray- Creophilus Leach, 1819, 2, 6, 8±12, 16±18, 21, acheila)], 37, 46 38 Erichsonius Fauvel, 1874, 7, 8, 10, 12±14 cubae (Jacquelin du Val, 1857) [ϭ Platydracus erythropterus LinneÂ, 1758 [Staphylinus and as tomentosus (Gravenhorst, 1802)], 41 subsp.], 25, 26, 41 cupreus Fourcroy, 1785 (Staphylinus), 35 Eucibdelus Kraatz, 1859, 10, 11, 17, 18 cupreus (Rossi, 1790) [ϭ Ocypus (Pseudocypus) Eugastus Sharp, 1876, 8, 17, 18 sericeicollis (MeÂneÂtrieÂs, 1832)], 35, 45 Euryporus Erichson, 1839, 15 cupripennis (Melsheimer, 1845) [Platydracus], excisus (J. MuÈller, 1950) [Ocypus (Pseudocypus)], 25, 39 43 curticollis (Bernhauer, 1917) [ϭ Platydracus tar- exulans (Erichson, 1839) [Platydracus], 39 salis (Mannerheim, 1843)], 41 curtipennis Motschulsky, 1849 [Ocypus (Ocy- falcifer (Nordmann, 1837) [Tasgius (Rayacheila), pus)], 42 and as subsp.], 46 cyaneopubens (Reitter, 1913) [Ocypus (Ocypus)], fallaciosus (J. MuÈller, 1926) [subsp. of Ocypus 42 (Pseudocypus) picipennis (Fabricius, 1792)], cyaneus (Paykull, 1789) (Staphylinus)[ϭ Ocypus 45 ophthalmicus (Scopoli, 1763)], 31 femoratus (Fabricius, 1801) [Platydracus], 39 cyanochloris Hochhuth, 1849 [ϭ Ocypus (Ocy- fenyesi (Bernhauer, 1917) [ϭ Platydracus cupri- pus) ophthalmicus ophthalmicus (Scopoli, pennis (Melsheimer, 1845)], 39 1763)], 42 festae (J. MuÈller, 1926) [Ocypus (Ocypus)], 42 cyclopus Peyron, 1858 (Ocypus) [incertae sedis], ¯avocephalus (Goeze, 1777) [Dinothenarus (Din- 47 othenarus), and as subsp.], 41 ¯avopunctatus (Latreille, 1804) [Platydracus], 39 daimio Sharp, 1889 [Staphylinus], 41 ¯uviaticus (Casey, 1915) [ϭ Platydracus mysticus dauricus (Mannerheim, 1830) [Platydracus], 39 (Erichson, 1840)], 40 decipiens Cameron, 1932 [Naddia], 38 fokiensis (Bernhauer, 1933) (Staphylinus)[ϭ Din- dehradunensis (Bernhauer, 1915) (Staphylinus) othenarus (Parabemus) insignis (J. MuÈller, [Dinothenarus (Parabemus)], 26, 27, 41 1926)], 27, 41 demissus (J. MuÈller, 1925) [Platydracus], 39 for®cularius (Motschulsky, 1860) (Matidus)[Ocy- densissimus (Bernhauer, 1933) [Ocypus (Pseudo- pus (Matidus)], 31, 43 cypus)], 43 formosae (Bernhauer, 1933) [Platydracus], 39 depolii (J. MuÈller, 1924) [ϭ Ocypus (Matidus) te- formosanus Shibata, 1982 [Thoracostrongylus], nebricosus (Gravenhorst, 1846)], 43 39 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_55 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 55

fortunatarum Wollaston, 1871 [Ocypus (Pseudo- graecus (Scheerpeltz, 1958) [ϭ Ocypus (Pseudo- cypus)], 31, 44 cypus) picipennis picipennis (Fabricius, Fortunocypus Coiffait, 1964 [ϭ Ocypus (Pseu- 1792)], 44 docypus)], 31, 43 graeseri Eppelsheim, 1887 [Ocypus (Pseudocy- fossator (Gravenhorst, 1802) [Platydracus], 39 pus), and as subsp.], 44 fossor (Fabricius, 1792) (Staphylinus)[ϭ Dinoth- gridellii (J. MuÈller, 1924) [ϭ Ocypus (Ocypus) enarus (Parabemus) fossor Scopoli, 1772], curtipennis Motschulsky, 1849], 42 26, 27, 41 grigiensis (Reitter, 1918) [subsp. of Ocypus (Ma- fossor (Scopoli, 1772) (Staphylinus)[Dinothena- tidus) nitens (Schrank, 1781)], 43 rus (Parabemus)], 26, 27, 41 griseipennis Fairmaire, 1889 (Staphylinus) [incer- frater Smetana, 1965 [Ocypus (Matidus)], 43 tae sedis], 47 fraternus (Fairmaire, 1891) [Agelosus], 45 griseosericans Fairmaire, 1894 [Emus], 38 fujiensis Naomi, 1992 [Ocypus (Pseudocypus)], 44 Hadropinus Sharp, 1889, 18 fulvicupreus Coiffait, 1956 [ϭ Ocypus (Pseudo- Hadrotes MaÈklin 1852, 8±11, 16±18, 22 cypus) sericeicollis (MeÂneÂtrieÂs, 1832)], 35, Haematodes Laporte, 1835, 10, 15 45 hagai Naomi, 1992 [Ocypus (Pseudocypus)], 44 fulvipennis Erichson, 1840 [Ocypus (Pseudocy- hairaerensis Li and Chen, 1993 [Ontholestes], 38 pus)], 32, 44 hakusanus Naomi, 1992 [Ocypus (Pseudocypus)], fulvipes (Motschulsky, 1858) [ϭ Tasgius (Ray- 44 acheila) morsitans (Rossi, 1790)], 46 harbinensis Li and Chen, 1993 (Ocypus (Neotas- fulvipes (Scopoli, 1763) [Platydracus], 39 gius)) [incertae sedis], 47 fulvotomentosus Eppelsheim, 1889 [Ocypus haroldi (Eppelsheim, 1884) [Ontholestes], 38 (Pseudocypus)], 31, 33, 44 hauseri (Bernhauer, 1933) [ϭ Ocypus (Pseudocy- fuscatoides (Coiffait, 1964) [Ocypus (Pseudocy- pus) fulvotomentosus Eppelsheim, 1889], 33, pus)], 44 44 fuscatus (Gravenhorst, 1802) [Ocypus (Pseudo- hauserianus (Bernhauer, 1933) [Platydracus], 39 cypus)], 44 hayashii Li, 1992 [Ontholestes], 38 fuscoaeneus Solsky, 1871 [Ocypus (Pseudocy- hebraeus (Smetana, 1978) [Abemus], 39 pus)], 44 heinzi Smetana, 1965 [Ocypus (Matidus)], 43 fuscofemoratus (J. MuÈller, 1923) [subsp. of Pla- Heinzia Korge, 1972, 14, 15 tydracus stercorarius (Olivier, 1795)], 40 helleni (J. MuÈller, 1926) [Ocypus (Pseudocypus)], fuscolineatus (Bernhauer, 1934) [Platydracus], 39 33, 44 fusiformis (Casey, 1915) [ϭ Platydracus centralis Hemiquedius Casey, 1915, 14 (Sharp, 1884)], 39 herculeanus (Coiffait, 1964) [Tasgius (Rayachei- Gabronthus Tottenham, 1955, 12±14 la)], 46 gallicus Coiffait, 1956 [ϭ Ocypus (Pseudocypus) herculeanus Semenov, 1906 [Physetops], 42 picipennis nevadensis (J. MuÈller, 1926)], 45 herzegovinensis (J. MuÈller, 1932) [ϭ Ocypus ganglbauerianus Bernhauer, 1939 (Staphylinus (Pseudocypus) picipennis picipennis (Fabri- (Xanthocypus)) [Dinothenarus (Parabe- cius, 1792)], 44 mus)], 27, 28, 41 Hesperus Fauvel, 1874, 14 garganicus Fiori, 1894 [ϭ Ocypus (Matidus) it- Heterothops Stephens, 1829, 10, 14, 15 alicus (Aragona, 1830)], 43 hidanus Watanabe, 1987 [Ocypus (Pseudocypus)], Gastrisus Sharp, 1876, 8, 18 44 giganteus Semenov, 1906 [Physetops], 42 himalaycus (Coiffait, 1982) [Ocypus (Pseudocy- Glenus Kraatz, 1857, 8, 18 pus)], 44 globulifer (Geoffroy, 1785) (Staphylinus)[Tas- himalayicus (Cameron, 1935) [ϭ Ocypus (Pseu- gius (Rayacheila)], 36, 46 docypus) helleni (J. MuÈller, 1926)], 33, 44 gloriosus Sharp, 1874 [Ocypus (Aulacocypus)], hirtus (LinneÂ, 1758) [Emus], 22, 38 32, 45 hissaricus DvorÏaÂk, 1984 [Ocypus (Matidus)], 43 Glyphesthus Kraatz, 1858, 15 hochhuthi Eppelsheim, 1878 [Ocypus (Matidus)], Goerius Westwood, 1827 [ϭ Ocypus (Ocypus)], 43 2, 31, 42 holosetosus (J. MuÈller, 1943) [ϭ Ocypus (Mati- goryi (Laporte, 1835) [Platydracus], 39 dus) brenskei (Reitter, 1884)], 42 gracilicornis (Hochhuth, 1849) [Tasgius (Rayach- horsti (Coiffait, 1974) [ϭ Ocypus (Pseudocypus) eila)], 46 obscuroaeneus obscuroaeneus Fairmaire, gracilis (Sharp, 1874) [Ontholestes], 38 1852], 44 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_56 Allen Press • DTPro System File # 01cc

56 AMERICAN MUSEUM NOVITATES NO. 3287

Hypabemus Scheerpeltz, 1966 [ϭ Dinothenarus kiushiuensis (Bernhauer, 1939) [Platydracus], 40 (Parabemus)], 26, 27, 41 kobensis Cameron, 1930 [ϭ Ocypus lewisius hypocrita (J. MuÈller, 1925) [Platydracus], 39 Sharp, 1874], 34, 44 hypsibatus (Bernhauer, 1899) [subsp. of Ocypus korgei Smetana, 1965 [ϭ Ocypus (Matidus) for- (Ocypus) ophthalmicus (Scopoli, 1763)], 42 ®cularius (Motschulsky, 1860)], 43 ibericus Coiffait, 1964 [ϭ Ocypus (Ocypus) krejcii Coiffait, 1984 (Parocypus) [incertae sedis], ophthalmicus atrocyaneus (Fairmaire, 47 1860)], 42 kuntzeni (J. MuÈller, 1926) [Ocypus (Matidus)], 43 immaculatus (Mannerheim, 1830) (Staphylinus) kyproticus Coiffait, 1964 [ϭ Ocypus (Ocypus) [Platydracus (Platydracus)], 25, 39 curtipennis Motschulsky, 1849], 42 impennis (Fauvel, 1882) [Ocypus (Pseudocypus)], latebricola Gravenhorst, 1806) [Platydracus], 40 44 Leistotrophus Perty, 1830, 2, 3, 11 imperatorius (Bernhauer, 1916) [Platydracus], 40 Leptopeltus Bernhauer, 1906, 10 impotens (Eppelsheim, 1889) [Platydracus], 40 Leucotachinus Coiffait and Saiz, 1968, 9 inauratus (Mannerheim, 1830) [Ontholestes], 38 lewisius Sharp, 1874 [Ocypus (Pseudocyphus), inderiensis (Motschulsky, 1845) (Rayacheila) 34, 44 [Tasgius (Rayacheila)], 36, 37, 46 liaoningensis Li and Chen, 1993 (Ocypus) [incer- indicus (Kraatz, 1859) [Platydracus], 40 tae sedis], 47 Indoquedius Cameron, 1932, 14, 15 libanicus Coiffait, 1956 [ϭ Ocypus (Pseudocypus) inexspectatus Eppelsheim, 1887 [Ocypus (Pseu- excisus (J. MuÈller, 1950)], 44 docypus)], 44 libanoticus (J. MuÈller, 1950) [ϭ Ocypus (Ocypus) inornatus (Sharp, 1874) [Platydracus], 40 curtipennis Motschulsky, 1849], 42 insignis (J. MuÈller, 1926) [Dinothenarus (Para- limbifrons (Hochhuth, 1849) [Tasgius (Rayachei- bemus)], 27, 28, 41 la)], 46 integer Abeille, 1900 [ϭ Ocypus (Ocypus) curti- lineatus (Walker, 1859) [ pennis Motschulsky, 1849], 42 ϭ Platydracus indicus intermedius (J. MuÈller, 1911) [subsp. of Ocypus (Kraatz, 1859)], 40 (Matidus) brunnipes (Fabricius, 1781)], 43 Liusus Sharp, 1889, 18 iranicus (Smetana, 1971) [ϭ Ocypus (Matidus) lomii (Cerruti, 1951) [Platydracus], 40 matilei Jarrige, 1971], 43 longicornis Stephens, 1832 (Philonthus), 9 ishiharai Shibata, 1994 [Naddia], 38 longimanus Smetana, 1965 [Ocypus (Matidus)], italicus (Aragona, 1830) (Staphylinus)[Ocypus 43 (Matidus)], 33, 43 lucanus (Horn, 1894) [ϭ Platydracus tarsalis italicus Erichson, 1840 [ϭ Ocypus (Matidus) it- (Mannerheim, 1843)], 40 alicus (Aragona, 1830)], 33, 43 luigionii (J. MuÈller, 1926) [ϭ Ocypus (Pseudo- iyonus Naomi, 1992 [Ocypus (Pseudocypus)], 44 cypus) aethiops (Waltl, 1835)], 43 izayoi Naomi,1992 [Ocypus (Pseudocypus)], 44 lusitanicus (J. MuÈller, 1943) [Tasgius (Rayachei- la)], 46 japonicus Sawada, 1965 (Ocypus (Ocypus)) [in- luteipes (LeConte, 1861) [Dinothenarus (Para- certae sedis], 47 bemus)], 41 javanus (Bernhauer, 1915) (Ontholestes)[Thora- costrongylus], 23 macgregori (Cooper, 1933) [ϭ Platydracus jeannei Coiffait, 1980 [Ocypus (Pseudocypus)], phoenicurus (Nordmann, 1837)], 40 44 macrocephalus (Gravenhorst, 1802) [Ocypus kaiseri (Bernhauer, 1933) (Apecholinus)[Agelo- (Matidus)], 43 sus], 35, 45 maculipennis (Kraatz, 1859) [Platydracus], 40 kansuensis (Bernhauer, 1933) (Staphylinus (Xan- maculosus (Gravenhorst, 1802) [ϭ Platydracus thocypus)) [Ocypus (Aulacocypus)], 33, 45 viduatus (Fabricius, 1801)], 41 kasyi (Scheerpeltz, 1962) [Platydracus], 40 maderae (Jarrige, 1943) [Tasgius (Rayacheila)], kendappa Naomi, 1992 [Ocypus (Pseudocypus)], 46 44 malaisei Scheerpeltz, 1965 [Naddia], 38 khnzoriani Coiffait, 1967 [Ocypus (Matidus)], 43 malaisei Scheerpeltz, 1965 [Thoracostrongylus], kiautschauensis (Bernhauer, 1916) [Dinothenarus 39 (Parabemus)], 41 manceps Smetana, 1965 [Ocypus (Matidus)], 43 kiimontanus Naomi, 1992 [Ocypus (Pseudocy- marginalis (GeneÂ, 1836) [Ontholestes], 38 pus)], 44 marginatus Cameron, 1944 (Staphylinus (Abe- kisonus Naomi, 1992 [Ocypus (Pseudocypus)], 44 mus)) [incertae sedis], 47 kiulungensis (Bernhauer, 1933) [Platydracus], 40 maritimus Coiffait, 1956 [ϭ Ocypus (Pseudocy- AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_57 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 57

pus) picipennis fallaciosus (J. MuÈller, 1926)], murgulensis Coiffait, 1978 (Ocypus) [incertae 45 sedis], 47 marocanus (Coiffait, 1964) [ϭ Ocypus (Pseudo- murinus (LinneÂ, 1758) [Ontholestes], 22, 38 cypus) picipennis barbarus (J. MuÈller, mus (BrulleÂ, 1832) (Staphylinus)[Ocypus (Pseu- 1926)], 45 docypus)], 31, 32, 44 martensi (Coiffait, 1982) [ϭ Abemus olivaceus myops (J. MuÈller, 1925) [Tasgius (Rayacheila)], (Cameron, 1928)], 23, 39 46 masaoi Hayashi, 1991 [subsp. of Agelosus uni- mysticus (Erichson, 1840) [Platydracus], 40 color Naomi, 1983], 45 mateui (Coiffait, 1954) [Ocypus (Pseudocypus)], Naddia Fauvel, 1867, 3, 6±11, 16, 17, 19, 22,38 33, 44 Nausicotus Sharp, 1884, 18 Matidus Motschulsky, 1860 [subgenus of Ocy- nemotoi Naomi, 1992 [Ocypus (Pseudocypus)], pus], 2, 7±9, 11, 15±17, 21, 29, 31, 32, 42 44 matilei Jarrige, 1971 [Ocypus (Matidus)], 43 Neobisnius Ganglbauer, 1895, 14 maxillosus (LinneÂ, 1758) [Creophilus, and as neomexicanus (Bernhauer and Schubert, 1914) [ϭ subsp.], 21, 38 Platydracus sepulchralis (Erichson, 1839)], medioximus Fairmaire, 1852 [Staphylinus], 41 40 megalocephalus (Nordmann, 1837) [Ocypus (Ma- Neotasgius J. MuÈller, 1925 [ϭ Platydracus (Pla- tidus)], 43 tydracus)], 24, 39 Megaquedius Casey, 1915 (subgen. of Quedius), nepalensis (Scheerpeltz, 1976) [Platydracus], 40 15 nepalicus (Coiffait, 1981) [Ocypus (Pseudocy- melanarius (Heer, 1839) [Tasgius (Rayacheila), pus)], 44 and as subsp.], 46 nepalicus (Coiffait, 1982) (Parontholestes)[Thor- meridionalis (Rosenhauer, 1847) [Platydracus], acostrongylus], 23, 39 40 nero (Faldermann, 1835) [ϭ Ocypus (Matidus) ni- messor (Nordmann, 1837) [Tasgius (Rayacheila)], tens Schrank, 1781], 34, 43 46 nevadensis (J. MuÈller, 1926) [subsp. of Ocypus Metocypus Coiffait, 1964 [ϭ Tasgius (Rayachei- (Pseudocypus) picipennis (Fabricius, 1792)], la)], 3, 36, 46 45 Microsaurus Dejean, 1833 (subgen. of Quedius), nigrellus (Horn, 1879) [Dinothenarus (Parabe- 15 mus)], 41 milleri Quedenfeldt, 1882 [Ocypus (Matidus), and nigrinus (Lucas, 1849) [Tasgius (Rayacheila)], 46 as subsp.], 43 nigroaeneus Sharp, 1889 [Ocypus (Pseudocy- minax (Mulsant and Rey, 1861) [Tasgius (Ray- pus)], 44 acheila)], 46 nitens Schrank, 1781 [Ocypus (Matidus), and as miniata Fauvel, 1895 [Naddia], 9, 38 subsp.], 34, 43 Miobdelus Sharp, 1889, 3, 8, 9, 11, 16, 17, 20, Nordus Blackwelder, 1952, 17 28,42 nubigena Smetana, 1965 [Ocypus (Matidus)], 43 Nudabemus Coiffait, 1982 [ϭ Ocypus (Ocypus)], mirtensis J. MuÈller, 1943 [ϭ Ocypus (Matidus) it- 31, 32, 43 alicus (Aragona, 1830)], 43 nudicollis Coiffait, 1977 (Ocypus) [incertae se- miwai (Bernhauer, 1943) [Ocypus (Ocypus)], 42 dis], 47 miyakei Bernhauer, 1943 [Thoracostrongylus], 39 modestus (Fall, 1907) [ϭ Platydracus sepulchral- obscuripes (Bernhauer, 1900) (Ocypus, ru®pes is (Erichson, 1839)], 40 var.) [ϭ Tasgius (Rayacheila) eppelsheimi- modestus (Sharp, 1884) [ϭ Platydracus centralis anus (Jakobson, 1909)], 37 (Sharp, 1884)], 39 obscuroaeneus Fairmaire, 1852 [Ocypus (Pseu- Moeocerus Fauvel, 1899, 15 docypus), and as subsp.], 44 montanus (Cameron, 1942) [Ocypus (Pseudocy- ochropus (J. MuÈller, 1950) [subsp. of Ocypus pus)], 44 (Matidus) nitens (Schrank, 1781)], 43 montanus Coiffait, 1977 [Platydracus], 40 oculatus (Bernhauer, 1929) (Staphylinus)[ϭ Pla- monticola Shibata, 1994 [Naddia], 38 tydracus (Platydracus) oculosus, nom. nov.], morio (Gravenhorst, 1802) (Staphylinus)[Tasgius 25, 40 (Rayacheila)], 36 oculatus (Fabricius, 1775) (Staphylinus)[Creo- morsitans (Rossi, 1790) [Tasgius (Rayacheila)], philus], 25 46 oculatus O. F. MuÈller, 1776 [Staphylinus], 25 mortuorum (Bernhauer, 1912) [Platydracus], 40 oculatus (Sharp, 1874) [Ontholestes], 38 muellerianus (Scheerpeltz, 1933) [Platydracus], oculosus, nom. nov. [for Platydracus oculatus 40 (Bernhauer, 1929)], 25, 40 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_58 Allen Press • DTPro System File # 01cc

58 AMERICAN MUSEUM NOVITATES NO. 3287

Ocypus Leach, 1819 [and subgenus], 2, 3, 6±11, pedemontanus (J. MuÈller, 1924) [Ocypus (Mati- 15±17, 21, 28, 31, 35, 42 dus), and as subsp.], 43 ohkurai Hayashi, 1978 [Agelosus], 45 pendjabensis (Bernhauer, 1915) [Tasgius (Ray- ohtsukaorum Naomi, 1992 [Ocypus (Pseudocy- acheila)], 46 pus)], 44 perniger (Scheerpeltz, 1976) [Platydracus], 40 olens (O. MuÈller, 1764) (Staphylinus)[Ocypus pernigrus Coiffait, 1964 [ϭ Ocypus (Ocypus) (Ocypus), and as subsp.], 31, 42 ophthalmicus atrocyaneus (Fairmaire, olivaceus (Cameron, 1928) [Abemus], 23, 39 1860)], 42 olympicus (Baudi, 1857) [Tasgius (Rayacheila)], perreaui Coiffait, 1984 [Platydracus], 40 46 peyerimhof® (J. MuÈller, 1926) [Tasgius (Tasgius)], Ontholestes Ganglbauer, 1895, 2, 3, 7±11, 16, 17, 46 19, 22,38 Philonthina, 6±12, 13, 14±17 opaciceps (Scheerpeltz, 1976) [Platydracus], 40 Philonthus Stephens, 1829, 7-10, 13 opacus Roth, 1851 (Ocypus) [incertae sedis], 47 Philothalpus Kraatz, 1857, 17, 18 ophthalmicus (Scopoli, 1763) (Staphylinus)[Ocy- phoenicurus (Nordmann, 1837) [Platydracus], 40 pus (Ocypus), and as subsp.], 31, 38, 42 Physetops Mannerheim, 1830, 2, 7±11, 16, 17, 19, orientalis Bernhauer, 1906 [Ontholestes], 38 28,42 orientalis (Bernhauer and Schubert, 1914) (Sta- picipennis (Fabricius, 1792) [Ocypus (Pseudocy- phylinus)[ϭ Ocypus (Pseudocypus) orientis, pus), and as subsp.], 3, 38, 44, 45 nom. nov.], 34, 44 pindensis (Coiffait, 1964) [ϭ Ocypus (Pseudocy- orientalis (Motschulsky, 1858) (Staphylinus)[ϭ pus) picipennis picipennis (Fabricius, 1792)], Creophilus maxillosus (LinneÂ)], 34 45 orientis, nom. nov. [for Ocypus (Pseudocypus) or- pinorum (Casey, 1915) [Platydracus], 40 ientalis (Bernhauer and Schubert, 1914)], 34, plagiicollis (Fairmaire, 1891) [Ocypus (Pseudo- 44 cypus)], 45 ormayi (Reitter, 1887) [Ocypus (Matidus)], 43 planipennis (AubeÂ, 1842) [Tasgius (Tasgius)], 46 ornaticauda (LeConte, 1863) [Staphylinus], 41 Platydracus Thomson, 1858 [and subgenus], 2, 6, ottomanus Fauvel, 1900 [Ocypus (Matidus)], 43 8±11, 15, 16, 18, 22, 24, 25, 38, 39 plebejus (Bernhauer, 1915) [Platydracus], 40 Paederinae, 7, 8 pleuralis (LeConte, 1861) [Dinothenarus (Para- paganus Sharp, 1874 [ϭ Platydracus brevicornis bemus)], 27, 41 (Motschulsky, 1862)], 39 pliginskii (Bernhauer, 1915) [Tasgius (Rayachei- Parabemus Reitter, 1909 [subgenus of Dinothen- la)], 46 arus], 10, 11, 16, 20, 26, 27, 41 Plociopterus Kraatz, 1857, 7, 17, 18 Paralapsodus Coiffait, 1974 [ϭ Tasgius (Rayach- Polyphematiana Strand, 1914, 18 eila)], 36, 37, 46 ponticus (Coiffait, 1978) [ϭ Ocypus (Pseudocy- paramurinus Li and Chen, 1993 [Ontholestes], 38 pus) picipennis picipennis (Fabricius, 1792)], Paraquedius Casey, 1915 (subgen. of Quedius), 45 14, 15 ponticus Smetana, 1968 [Ocypus (Matidus)], 43 Paratasgius Jarrige, 1952 [ϭ Tasgius (Tasgius)], praelongus (Mannerheim, 1830) [Platydracus], 36, 45 40 Paraxenopygus Bernhauer, 1911, 18 praetorius (Bernhauer, 1915) [Tasgius (Tasgius)], Parocypus Bernhauer, 1915 [ϭ Dinothenarus 46 (Parabemus)], 26±28, 41 prainae (Eppelsheim, 1895) [Dinothenarus (Par- Parontholestes Coiffait, 1982 [ϭ Thoracostron- abemus)], 41 gylus], 23, 39 pratti (Scheerpeltz, 1962) [Platydracus], 40 parvulus Sharp, 1874 [Ocypus (Aulacocypus)], 45 primigenius (J. MuÈller, 1923) [subsp. of Ocypus parvulus, nom. nov. [for Platydracus parvus (Matidus) syriacus Baudi, 1848], 43 (Cameron, 1932)], 40 Protabemus Scheerpeltz, 1966 [ϭ Dinothenarus parvus (Cameron, 1932) (Staphylinus)[ϭ Platyd- (Dinothenarus)], 26, 41 racus parvulus, nom. nov.], 40 Protocypus J. MuÈller, 1923 [ϭ Ocypus (Pseudo- patricius (Bernhauer, 1915) [Platydracus (Chae- cypus)], 31, 32, 43 todracus)], 25, 41 Protogoerius Coiffait, 1956, 6±11, 16, 17, 21, 30, pecoudi Jarrige, 1954 (Staphylinus (Goerius)) [in- 42 certae sedis], 47 proximus Kirschenblat, 1936 [Ontholestes], 38 pedator (Gravenhorst, 1802) (Staphylinus)[Tas- pseudoalpestris (J. MuÈller, 1926) [subsp. of Ocy- gius (Tasgius)], 7, 36, 46 pus (Matidus) brevipennis (Heer, 1839)], 42 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_59 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 59

pseudobrevipennis Coiffait, 1964 [ϭ Ocypus (Ma- ru®ceps (Cameron, 1932) [Dinothenarus (Dinoth- tidus) chevrolatii (Baudi, 1848)], 43 enarus)], 41 Pseudocypus Mulsant and Rey, 1876 [subgenus ru®cornis Bernhauer, 1913 [ϭ Staphylinus rubri- of Ocypus], 2, 3, 6±8, 11, 16, 21, 31, 32, 43 cornis (AdaÂm, 1987)], 41 pseudolens Coiffait, 1964 [ϭ Ocypus (Ocypus) ru®pennis Bernhauer, 1915 [Naddia], 38 curtipennis Motschulsky, 1849], 42 ru®pes Eppelsheim, 1884 (Ocypus)[ϭ Tasgius pseudopaganus (Bernhauer, 1914) [Platydracus, (Rayacheila) eppelsheimianus (Jakobson, and as subsp.], 40 1909)], 37 pseudopatricius (J. MuÈller, 1926) [subsp. of Pla- ru®pes Latreille, 1806 (Astrapaeus)[ϭ Tasgius tydracus pseudopaganus (Bernhauer, 1914)], (Tasgius) pedator (Gravenhorst, 1802)], 36, 40 37 Pseudotasgius Seidlitz, 1891 [ϭ Tasgius (Tas- rutilicauda (Horn, 1879) [Platydracus], 40 gius)], 36, 45 pubescens (DeGeer, 1774) (Staphylinus)[Dinoth- sabi Naomi, 1992 [Ocypus (Pseudocypus)], 45 enarus (Dinothenarus)], 26, 41 sachalinensis (Matsumura, 1911) [Platydracus], pullus Hochhuth, 1849 [Ocypus (Matidus)], 34, 40 43 sagaris Smetana, 1992 [Dinothenarus (Dinoth- pyrenaeus (J. MuÈller, 1924) [subsp. of Ocypus enarus)], 41 (Matidus) pedemontanus (J. MuÈller, 1924)], sahlbergi (J. MuÈller, 1926) [subsp. of Tasgius 43 (Rayacheila) melanarius (Heer, 1839)], 46 saphyrinus (LeConte, 1861) [Dinothenarus (Par- quadraticeps (Casey, 1924) [ϭ Platydracus zon- abemus)], 42 atus (Gravenhorst, 1802)], 41 Scariphaeus Erichson, 1839, 15 quadraticeps (MeÂneÂtrieÂs, 1832) [Ocypus (Mati- schatzmayri (J. MuÈller, 1923) [subsp. of Ocypus dus)], 43 (Pseudocypus) obscuroaeneus Fairmaire, quadrimaculatus (Cameron, 1932) [Ocypus (Ocy- 1852], 44 pus)], 31, 32, 42 scutiger Sharp, 1889 [Ocypus (Pseudocypus)], Quediina, 6±10, 12±14, 15, 17 45 Quedionuchus Sharp, 1884 (subgen. of Que- semenowi Reitter, 1887 [Ocypus (Pseudocypus)], dius),8, 14, 15 45 Quedius Stephens, 1829, 14, 15 semialatus (J. MuÈller, 1904) [subsp. of Ocypus queinneci Coiffait, 1984 (Ocypus?) [incertae sed- (Matidus) nitens (Schrank, 1781)], 43 is], 47 semipurpureus (Kraatz, 1859) [Platydracus], 40 septentrionalis Watanabe, 1984 [Ocypus (Pseu- Rabigus Mulsant and Rey, 1876, 10 docypus)], 45 rambouseki (J. MuÈller, 1925) [ϭ Ocypus (Pseu- sepulchralis (Erichson, 1839) [Platydracus], 40 docypus) nigroaeneus Sharp, 1889], 44 sericeicollis (MeÂneÂtrieÂs, 1832) (Pseudocypus) Raphirus Stephens, 1829 (subgen. of Quedius), 14 [Ocypus (Pseudocypus)], 35, 45 Rayacheila Motschulsky, 1845 [subgenus of Tas- sericeomicans (Bernhauer, 1931) [Ocypus (Pseu- gius], 2, 6±8, 11, 16, 20, 36, 37, 46 docypus)], 45 reimoseri (Bernhauer, 1906) [Ocypus (Matidus)], serotinus (AdaÂm, 1992) [Ocypus (Pseudocypus)], 43 45 reitterianus (Bernhauer, 1933) [Platydracus], 40 sharpi (Fauvel, 1901) [Platydracus], 40 rhaeticus Eppelsheim, 1873 [Ocypus (Matidus)], sibiricus (Gebler, 1830) [Dinothenarus (Parabe- 42 mus)], 42 riojanus Hozman, 1977 [Platydracus], 40 sikkimensis (Bernhauer, 1920) [Tasgius (Tas- robustus Coiffait, 1964 [subsp. of Ocypus (Mati- gius)], 46 dus) milleri Quedenfeldt, 1882], 43 sikkimensis Coiffait, 1985 (Ocypus) [incertae se- rodopensis Coiffait, 1971 [subsp. of Ocypus (Ocy- dis], 47 pus) ophthalmicus (Scopoli, 1763)], 42 sikkimensis Wendeler, 1927 [Creophilus], 38 rodopensis (Coiffait, 1971) [ϭ Ocypus (Pseudo- silensis Fiori, 1894 [ϭ Ocypus (Matidus) italicus cypus) picipennis picipennis (Fabricius, (Aragona, 1830)], 43 1792)], 45 similis: (Fabricius, 1792) [ϭ Ocypus (Matidus) ni- rossii Jarrige, 1954 [ϭ Ocypus (Pseudocypus) ser- tens (Schrank, 1781)], 34, 43 iceicollis (MeÂneÂtrieÂs, 1832)], 35, 45 similis (Paykull, 1789) (Staphylinus)[Tasgius rubricornis (AdaÂm, 1987) [Staphylinus], 41 (Rayacheila)], 34 rubripennis (Reiche and Saulcy, 1856) [Tasgius simulator Eppelsheim, 1878 [Ocypus (Matidus)], (Rayacheila)], 46 34, 35, 43 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_60 Allen Press • DTPro System File # 01cc

60 AMERICAN MUSEUM NOVITATES NO. 3287

simulator Kirschenblat, 1936 [Ontholestes], 38 tenuicornis (Kraatz, 1859) [Ontholestes], 38 smetanai Naomi, 1992 [Ocypus (Pseudocypus)], teruelensis (Coiffait, 1976) [ϭ Ocypus (Pseudo- 45 cypus) picipennis nevadensis (J. MuÈller, Smilax Laporte, 1835, 10, 14 1926)], 45 solarii (J. MuÈller, 1923) [Ocypus (Matidus)], 43 tessellatus (Geoffroy, 1785) [Ontholestes], 38 solskyi (Fauvel, 1875) (Staphylinus)[Tasgius testaceipes Fairmaire, 1887 (Ocypus) [incertae (Rayacheila), and as subsp.], 36, 46 sedis], 47 sparsus (Cameron, 1932) [Platydracus], 40 teter (Bondroit, 1913) [ϭ Platydracus aureofas- speculifrons (Bernhauer, 1939) [Platydracus], 40 ciatus (Motschulsky, 1862)], 39 spilleri Bernhauer, 1923 [Physetops], 42 Thinopinus LeConte, 1852, 8, 9, 11, 12, 16, 17, springeri J. MuÈller, 1923 [subsp. of Staphylinus 18, 22 erythropterus LinneÂ, 1758], 41 Thoracostrongylus Bernhauer, 1915, 8±11, 16, Staphylinina, 6±15, 16±18, 22 17, 19, 23,39 Staphylinus LinneÂ, 1758, 2, 3, 6±8, 10, 11, 16± tomentosus (Baudi, 1869) (Staphylinus)[ϭ Ocy- 18, 20, 25, 27, 41 pus (Pseudocypus) orientis, nom. nov.], stercorarius (Olivier, 1795) [Platydracus and as tomentosus (Gravenhorst, 1802) (Staphylinus) subsp.], 9, 24, 40 [Platydracus], 34, 41 stevensi (Cameron, 1932) [Wasmannellus], 45 torvus Smetana, 1965 [Ocypus (Matidus)], 43 Strouhalium Scheerpeltz, 1962, 15 transadriaticus (J. MuÈller, 1926) [ϭ Ocypus (Pseu- Styngetus Sharp, 1884, 17, 18 docypus) mus (BrulleÂ, 1832)], 44 subaeneipennis (Scheerpeltz, 1976) [Platydra- transversiceps (Luze, 1904) [Tasgius (Tasgius)], cus], 40 46 subaenescens Wollaston, 1864 [Ocypus (Pseudo- trapezensis Coiffait, 1964 [Ocypus (Matidus)], 43 cypus)], 45 Triacrina, 12 subaeneus (Roth, 1851) [Platydracus], 40 Triacrus Nordmann, 1837, 18 subauronotatus Coiffait, 1977 [Platydracus], 40 Trichocosmetes Kraatz, 1859, 18 submarmorellus (Schubert, 1908) [Platydracus], Trichoderma Stephens, 1829 [ϭ Dinothenarus], 2 40 tricinctus (Aragona, 1830) [Tasgius (Rayachei- submetallicus (LeConte, 1861) [ϭ Platydracus la)], 46 tarsalis (Mannerheim, 1843)], 40 tricolor (Coiffait, 1974) [Tasgius (Rayacheila)], subtilis Tikhomirova, 1973 (Ocypus (Pseudocy- 46 pus)) [Ocypus (Aulacocypus)], 45 Trigonopselaphus Gemminger and Harold, 1868, subviridis (Bernhauer, 1933) [Platydracus], 40 18 sumakowi (Bernhauer, 1911) [Platydracus], 40 triplicans (Casey, 1924) [ϭ Platydracus viridanus suturalis Matsumura, 1911 (Staphylinus (Ocy- (Horn, 1879)], 41 pus)) [incertae sedis], 47 tristis Bernhauer, 1920 [Wasmannellus], 35, 45 svozili M. DvorÏaÂk, 1984 [Ocypus (Pseudocypus)], turcicus (Bernhauer, 1923) [Ocypus (Matidus)], 45 43 sylvaticus Wollaston, 1865 [Ocypus (Pseudocy- Tympanophorus Nordmann, 1837, 17, 18 pus)], 31, 45 syriacus Baudi, 1848 [Ocypus (Matidus), and as umbricola Wollaston, 1864 [Ocypus (Pseudocy- subsp.], 43 pus)], 45 szechuanensis (Bernhauer, 1935) [Dinothenarus unicolor Naomi, 1983 [Agelosus, and as subsp.], (Parabemus)], 42 45 ussuriensis (Solsky, 1871) [Platydracus], 41 Tachinoporus Cameron, 1928, 10 Tachyporini, 9, 10, 14 Valdiviodes Smetana, 1981, 14, 15 taiwanensis Shibata, 1979 [Naddia], 38 varipes (Sachse, 1852) [ϭ Platydracus femoratus tarsalis (Mannerheim, 1843) [Platydracus], 40 (Fabricius, 1801)], 39 Tasgius Stephens, 1829 [and as subgenus], 2, 3, Velleiopsis Fairmaire, 1882, 14 6, 7±11, 16, 17, 20, 36,45 Velleius Leach, 1819, 14 tataricus (Pallas, 1773) (Staphylinus)[Physetops], velutinus Scheerpeltz, 1965 [Thoracostrongylus], 28, 42 39 tauricus (J. MuÈller, 1932) [ϭ Ocypus (Pseudocy- vesubiensis Coiffait, 1964 [ϭ Ocypus (Matidus) pus) mus (BrulleÂ, 1832)], 44 chevrolatii (Baudi, 1848)], 43 temporalis (Casey, 1915) [Platydracus], 41 vicarius (Sharp, 1889) [Platydracus], 41 tenebricosus (Gravenhorst, 1846) [Ocypus (Mati- viduatus (Fabricius, 1801) [Platydracus], 41 dus)], 10, 43 villipennis Kraatz, 1859 [Creophilus], 38 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_61 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 61

villosus (Gravenhorst, 1802) [subsp. of Creophi- xanthocephalus (Kraatz, 1859) (Staphylinus) lus maxillosus (LinneÂ, 1758)], 38 [Dinothenarus (Dinothenarus)], 26, 41 violaceus (Gravenhorst, 1802) [ϭ Platydracus Xanthocypus J. MuÈller, 1925 [ϭ Ocypus (Ocy- (Platydracus) cupripennis (Melsheimer, pus)], 27, 31, 33, 42 1844)], 25, 39 Xanthopygina, 6±10, 12, 13, 15±17,21 violaceus (Olivier, 1795) (Staphylinus)[Plochion- Xanthopygus Kraatz, 1857, 9, 18 ocerus], 25 Xenopygus Bernhauer, 1906, 18, viridanus (Horn, 1879) [Platydracus], 41 vulpinus (Nordmann, 1837) (Staphylinus)[Platyd- yamato Naomi, 1992 [Ocypus (Pseudocypus)], 45 racus], 25, 41 yolensis (Cerruti, 1951) [Platydracus], 41 yoroi Naomi, 1992 [Ocypus (Pseudocypus)], 45 walkeri (Fauvel, 1898) [Tasgius (Rayacheila)], 46 yuinus Naomi, 1992 [Ocypus (Pseudocypus)], 45 Wasmannellus Bernhauer, 1920, 8±11, 17, 21, 35, yunnanensis (Bernhauer, 1933) (Staphylinus) 45 [Platydracus (Platydracus)], 25, 41 wasmanni (Bernhauer, 1920) [Ocypus (Ocypus)], yunnanensis (Bernhauer, 1943) (Staphylinus)[ϭ 31, 42 Platydracus (Platydracus) yunnanicus, nom. weisei Harold, 1877 [Ocypus (Ocypus)], 11, 31, nov.], 25, 41 42 yunnanicus, nom. nov. [for Platydracus (Platyd- westermanni (Erichson, 1840) (Caranistes)[Nad- racus) yunnanensis (Bernhauer, 1943)], 25, dia], 22 41 winkleri (Bernhauer, 1906) (Staphylinus)[Tasgius (Rayacheila)], 36, 46 zimmermanni Korge, 1965 [ϭ Ocypus (Matidus) wittmeri Coiffait, 1977 [Platydracus], 41 chevrolatii (Baudi, 1848)], 43 wittmeri Coiffait, 1982 [Naddia], 38 zonatus (Gravenhorst, 1802) [Platydracus], 41 AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_62 Allen Press • DTPro System File # 01cc

62 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 4±10. 4. Tasgius melanarius, left mandible, ventral view. 5. Tasgius ater, left mandible, ventral view. 6, 7. Platydracus stercorarius: left and right mandibles (both in ventral view). 8. Staphylinus erythropterus, both mandibles, ventral view. 9. Naddia sp., right mandible, dorsal view. 10. Ocypus ophthalmicus, both mandibles, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_63 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 63

Figs. 11±16. 11. Platydracus stercorarius, right mandibular prostheca, ventral view. 12. Abemus chloropterus, left mandibular prostheca, ventral view. 13. Ocypus picipennis, left mandibular prostheca, ventral view. 14. Tasgius ater, left mandibular prostheca, ventral view. 15. Staphylinus dimidiaticornis, left mandibular prostheca, ventral view. 16. Tasgius melanarius, left mandibular prostheca, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_64 Allen Press • DTPro System File # 01cc

64 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 17±21. 17. Anisolinus elegans, left maxilla, ventral view. 18. Craspedomerus sp., left maxilla, ventral view. 19. Plociopterus comptus, right maxilla and mentum, ventral view. 20. Philonthus caeru- leipennis, labium, mentum, and left maxilla, ventral view. 21. Quedius laticollis, right maxilla, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_65 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 65

Figs. 22±27. 22. Dinothenarus pubescens, last segment of maxillary palpus. 23. Ocypus sylvaticus, last segment of maxillary palpus. 24. Staphylinus dimidiaticornis, last segment of maxillary palpus. 25. Ocypus tenebricosus, last segment of maxillary palpus. 26. Staphylinus erythropterus, last segment of maxillary palpus. 27. Staphylinus erythropterus, peglike setae in glandular openings on last segment of maxillary palpus. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_66 Allen Press • DTPro System File # 01cc

66 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 28±32. 28. Dinothenarus fossor, last segment of maxillary palpus. 29. Tasgius ater, last seg- ment of maxillary palpus. 30. Tasgius melanarius, last segment of maxillary palpus. 31. Ontholestes murinus, longitudinal furrows at base of last segment of maxillary palpus. 32. Philonthus politus, lon- gitudinal furrows at base of last segment of maxillary palpus. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_67 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 67

Figs. 33±38. 33. Philonthus politus, labium, mentum, submentum, and left maxilla, ventral view. 34. Amichrotus sp., anterolateral setae on mentum. 35. Ocypus picipennis, labium, mentum, and sub- mentum. 36. Ontholestes cingulatus, labium, mentum, and submentum. 37. Ocypus tenebricosus, labium, mentum, and submentum. 38. Dinothenarus pubescens, labium, labial palpus, and mentum. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_68 Allen Press • DTPro System File # 01cc

68 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 39±44. 39. Philonthus politus, dorsal face of neck. 40. Abemus chloropterus, dorsal face of neck. 41. Naddia sp., left posterior corner of head, and part of dorsal face of neck. 42. Dinothenarus fossor, joining of superior and inferior marginal lines of pronotal hypomeron, ventral view. 43. Tasgius ater, joining of superior and inferior marginal lines of pronotal hypomeron, ventral view. 44. Ontholestes murinus, joined marginal lines of pronotal hypomeron, narrowly rounded at anterolateral margin of prosternum, and angulately extended anterior angles of pronotum, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_69 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 69

Figs. 45±52. 45. Philonthus caeruleipennis, superior and inferior marginal lines of pronotal hypo- meron not joined, ventral view. 46. Agelosus carinatus, superior and inferior marginal lines of pronotal hypomeron distinctly not joined, ventral view. 47. Ocypus weisei, superior and inferior marginal lines of pronotal hypomeron not quite joined, ventral view. 48. Ocypus tenebricosus, superior and inferior marginal lines of pronotal hypomeron not quite joined, ventral view. 49. Creophilus maxillosus, superior marginal line of pronotal hypomeron becoming obsolete near anterior angles, ventral view. 50. Staphy- linus dimidiaticornis, joining of superior and inferior marginal lines of pronotal hypomeron, ventral view. 51. Miobdelus sp., joining of superior and inferior marginal lines of pronotal hypomeron, ventral view. 52. Platydracus stercorarius, superior and inferior marginal lines of pronotal hypomeron not quite joined, appearing as a grooved carina anteriorly, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_70 Allen Press • DTPro System File # 01cc

70 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 53±58. 53. Xanthopygus analis, disc of pronotum virtually invisible in ventral view. 54. Din- othenarus pubescens, joining of superior and inferior marginal lines of pronotal hypomeron and pro- sternum with strongly arched sternacostal ridge (arrow), ventral view. 55. Quedius laticollis, proepi- meron, ventral view. 56. Abemus chloropterus, proepimeron, ventral view. 57. Staphylinus erythropterus, proepimeron absent. 58. Ocypus ophthalmicus, medial margin of sternacostal ridge forming a prominent ®nlike carina. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_71 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 71

Figs. 59±64. 59. Dinothenarus pubescens, series of long setae (some represented only by their sockets) on mesosternum. 60. Miobdelus sp., series of long setae on mesosternum. 61. Craspedomerus sp., series of long setae on mesosternum without transverse ridge. 62. Tasgius melanarius, mesocoxal acetabulum. 63. Ocypus weisei, mesocoxal acetabulum. 64. Staphylinus dimidiaticornis, mesocoxal cav- ities, not margined posteromedially. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_72 Allen Press • DTPro System File # 01cc

72 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 65±71. 65. Dinothenarus pubescens, dorsal apicolateral lobe of hind coxa, with a series of spinelike setae. 66. Ontholestes cingulatus, dorsal apicolateral lobe of hind coxa with serial setae almost indistinguishable from other setae. 67. Anisolinus elegans, empodial setae of middle tarsus, ventral view. 68. Miobdelus sp., empodial setae of middle tarsus, ventral view. 69. Quedius laevicollis, empodial setae of middle tarsus, ventral view. 70. Philonthus splendens, apical portion of last segment of middle tarsus, empodial setae absent, ventral view. 71. Craspedomerus sp., apical portion of last segment of middle tarsus, empodial setae absent, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_73 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 73

Figs. 72±77. 72. Philonthus politus, anterior portions of superior and inferior marginal lines of pronotal hypomeron, ventral view. 73. Craspedomerus sp., anterior portions of superior and inferior marginal lines of pronotal hypomeron, ventral view. 74. Philonthus politus, ventral view of head, no infraorbital ridge. 75. Quedius laticollis, superior marginal line of pronotal hypomeron not de¯ected under anterior angle of pronotum, ventrolateral view. 76. Quedius laticollis, pronotal hypomeron and prosternum, notosternal suture present. 77. Quedius rainieri, head with infraorbital ridge (arrow), oblique ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_74 Allen Press • DTPro System File # 01cc

74 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 78±83. 78. Quedius rainieri, dorsal face of neck, dorsal basal ridge absent. 79. Xenopygus analis, superior marginal line of pronotal hypomeron not de¯ected under anterior angle of pronotum, ventral view. 80. Xenopygus analis, dorsal face of neck, dorsal basal ridge present (arrow). 81. Xeno- pygus analis, head with postmandibular ridge, ventrolateral view. 82. Ocypus weisei, head with rudi- mentary postmandibular ridge, ventral view. 83. Anisolinus elegans, superior marginal line of pronotal hypomeron not distinctly de¯ected under anterior angle of pronotum, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_75 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 75

Figs. 84±87. 84. Amichrotus sp., maxillary palpus. 85. Ocypus picipennis, maxillary palpus. 86. Philonthus caeruleipennis, prosternum. 87. Philonthus caeruleipennis, mesosternum, showing transverse ridge, and anterior metasternal projection. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_76 Allen Press • DTPro System File # 01cc

76 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 88±91. 88. Craspedomerus sp., narrowly, somewhat irregularly swollen lateral margin of pro- notum, imitating the superior marginal line of pronotum, lateral view. 89. Quedius rainieri, ligula not emarginate. 90. Quedius laticollis, mentum with two anterolateral setae. 91. Quedius rainieri, right mandible with dorsolateral ridge (arrow), dorsal view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_77 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 77

Figs. 92±98. 92. Amichrotus sp., superior and inferior marginal lines of pronotal hypomeron not joined, ventral view. 93. Amichrotus sp., ligula minutely emarginate. 94. Anisolinus elegans, left man- dible, ventral view. 95. Amichrotus sp., palpifer with two apical setae. 96. Amichrotus sp., mandibles, ventral view. 97. Amichrotus sp., prosternum. 98. Amichrotus sp., mesosternum with longitudinal carina, and anterior metasternal projection. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_78 Allen Press • DTPro System File # 01cc

78 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 99±104. 99. Dinothenarus badipes, mentum with two setae. 100. Xenopygus analis, right dorsal face of neck, with dorsal basal ridge (arrow). 101. Emus hirtus, intercoxal process of mesosternum widely separating middle coxae. 102. Platydracus fossator, left posterolateral area of head, showing postocular seta (arrow), dorsal view. 103. Staphylinus erythropterus, right posterolateral area of head, showing postocular seta (arrow), dorsal view. 104. Abemus chloropterus, left posterolateral area of head, showing postocular seta (arrow), dorsal view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_79 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 79

Figs. 105±110. 105. Naddia sp., head, dorsal view. 106. Naddia sp., left anterior portion of pro- notum. 107. Ontholestes cingulatus, left posterior area of head, showing eye and tempus, dorsal view. 108. Ontholestes cingulatus, left mandible, ventral view. 109. Ocypus nitens semialatus, pronotum, dorsal view. 110. Ontholestes cingulatus, pronotum, dorsal view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_80 Allen Press • DTPro System File # 01cc

80 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 111±116. 111. Dinothenarus pubescens, pronotum showing both marginal lines of pronotal hypomeron, and two lateral macrosetae, lateral view. 112. Staphylinus erythropterus, pronotum showing both marginal lines of pronotal hypomeron, and two lateral macrosetae, lateral view. 113. Ocypus ophthalmicus ophthalmicus, pronotum showing both marginal lines of pronotal hypomeron, and two lateral macrosetae, ventrolateral view. 114. Ocypus nitens semialatus, pronotum showing both marginal lines of pronotal hypomeron, and two lateral macrosetae, lateral view. 115. Physetops sp., dorsal face of neck, with dorsal portion of nuchal ridge missing. 116. Ontholestes cingulatus, dorsal face of neck, with nuchal ridge. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_81 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 81

Figs. 117±122. 117. Ontholestes murinus, mesosternum with fully developed medial carina. 118. Thoracostrongylus sp., mesosternum with short medial carina at base. 119. Dinothenarus fossor, labium with emarginate ligula, ventral view. 120. Dinothenarus pubescens, labium and mentum. 121. Dinoth- enarus pubescens, right mandible and right maxilla, ventral view. 122. Abemus chloropterus, mesoster- num, showing sockets of serial setae originating behind small elevations. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_82 Allen Press • DTPro System File # 01cc

82 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 123±128. 123. Abemus chloropterus, labium and mentum. 124. Abemus chloropterus, left mandible, dorsal view. 125. Miobdelus sp., posterolateral portion of head, showing pitlike depressions. 126. Miobdelus sp., ®rst three visible abdominal tergites, showing bisinuate posterior basal line. 127. Ocypus picipennis picipennis, left mandible, ventral view. 128. Ocypus picipennis picipennis, right mandible, ventral view. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_83 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 83

Figs. 129±135. 129. Protogoerius brullei, last segment of maxillary palpus. 130. Ocypus picipennis picipennis, hind tibia with spines (arrow) on dorsolateral face. 131. Ocypus mus, last segment of labial palpus. 132. Ocypus weisei, last segment of labial palpus. 133. Ocypus olens, last segment of labial palpus. 134. Platydracus stercorarius, last segment of maxillary palpus. 135. Platydracus stercorarius, last segment of labial palpus. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_84 Allen Press • DTPro System File # 01cc

84 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 136±141. 136. Platydracus stercorarius, pronotal hypomeron, ventral view. 137. Platydracus stercorarius, right coxal cavity, ventral view. 138. Staphylinus dimidiaticornis, left mandible, ventral view. 139. Staphylinus erythropterus, last segment of labial palpus. 140. Dinothenarus pubescens, pos- teromedial portion of head and neck with pubescence, dorsal view. 141. Dinothenarus pubescens, pro- notum with pubescence. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_85 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 85

Figs. 142±145. 142. Dinothenarus fossor, mesosternum. 143. Miobdelus sp., left mandibular pros- theca, ventral view. 144. Miobdelus sp., maxillary palpus. 145. Miobdelus sp., labial palpus. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_86 Allen Press • DTPro System File # 01cc

86 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 146±151. 146. Protogoerius brullei, underside of head, showing gula. 147. Protogoerius brul- lei, left mandible, dorsal view. 148. Protogoerius brullei, maxillary (left) and labial palpus (right). 149. Ocypus tenebricosus, maxilla, showing setae on palpifer. 150. Ocypus tenebricosus, maxillary palpus. 151. Ocypus tenebricosus, labial palpus. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_87 Allen Press • DTPro System File # 01cc

2000 SMETANA AND DAVIES: RECLASSIFICATION OF STAPHYLINUS 87

Figs. 152±155. 152. Agelosus carinatus, underside of head, showing abbreviated infraorbital ridge (arrow). 153. Agelosus carinatus, last segment of labial palpus. 154. Tasgius melanarius, mesosternal intercoxal process. 155. Tasgius ater, pronotal hypomeron and prosternum. AMNH NOVITATES Tuesday Dec 11 2001 10:11 AM 2000 novi 99163 Mp_88 Allen Press • DTPro System File # 01cc

88 AMERICAN MUSEUM NOVITATES NO. 3287

Figs. 156±159. 156. Tasgius melanarius, mentum and submentum. 157. Tasgius melanarius, pro- sternum. 158. Tasgius melanarius, metasternum, showing anterior metasternal projection. 159. Platyd- racus stercorarius, pronotum.

a This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).