Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2743, pp. 1-13, figs. 1-4, table 1 August 4, 1982 Redefinition, Revised Phylogeny, and Relationships of Pseudopsinae (Coleoptera, Staphylinidae) ALFRED F. NEWTON, JR.'

ABSTRACT New evidence is presented to support transfer vised cladistic analysis of the four genera is pre- of the genera Nanobius, Asemobius, and Zalobius sented with the conclusion that Pseudopsis is the from the Piestinae to Pseudopsinae, currently in- sister group of the remaining genera. The rela- cluding only Pseudopsis. The subfamily is rede- tionships of the subfamily are discussed; a close scribed, a key to genera given, and the natural relationship to the Phloeocharinae rather than to history and distribution briefly reviewed. A re- the Oxytelinae is suggested. INTRODUCTION The subfamily Pseudopsinae, with the sin- paper is intended as a supplement to, rather gle included genus Pseudopsis, has recently than a replacement for, Herman's revisions been thoroughly revised by Herman (1975). (1975, 1977). Thus the treatment is largely The same author shortly thereafter revised limited to making changes in the subfamily the three piestine genera Nanobius, Asemo- description, generic keys, etc. that are re- bius, and Zalobius (Herman, 1977). In the quired by combining all four genera in one latter work Herman cited many similarities subfamily. A cladistic analysis is presented, among these four genera but concluded that combining data from Herman's analyses the available evidence did not adequately (1975, 1977) with new data. support the hypothesis that the four genera constituted a monophyletic unit; Nanobius and its allies were tentatively retained in the MATERIALS AND METHODS Piestinae. In the course ofthis study I have examined The purpose of the present work is to pro- KOH-cleared, chlorazyl black-stained spec- vide new evidence that, in my view, firmly imens of the Pseudopsinae with both dis- establishes a close relationship of the four secting and compound microscopes. All genera and justifies placing them in a single species ofthe subfamily have been examined subfamily, Pseudopsinae sensu novo. This with the exception of most species of the

I Research Associate, Department of Entomology, American Museum of Natural History; Curatorial Associate, Entomology Department, Museum of Comparative Zoology, Harvard University.

Pseudopsis sulcata complex, which according tum, elytra, and sometimes head with lon- to Herman (1975) are distinguishable only gitudinal carinae or costae. Vestiture sparse, by details of the aedeagus. Several cleared of simple tapered setae and/or modified and partially dissected specimens were crit- scalelike or clubbed setae. Color reddish ical-point-dried, coated with gold-palladium brown. Body with or without tendency to mixture and examined with a scanning elec- accumulate mud and debris. tron microscope. Labrum with anterior margin truncate or The cladistic methods used here are essen- sinuate, with exposed epipharyngeal lobes, tially those of Herman (1970, 1975, 1977). dorsal surface with long setae. Epipharynx Other subfamilies of the Staphylinidae and with globosetae near anterior margin. Ante- related families such as the Silphidae and clypeus large and apparently permitting ex- Leiodidae have been used as out-groups to trusion of labrum. Supra-antennal ridge car- determine character polarities. inate or not. Epistomal suture present, posteriorly convex, evident externally as ACKNOWLEDGMENTS ridge or groove. Neck more or less distinct, at least laterally. Antenna slender to rela- I thank Dr. Lee H. Herman, Jr., and Dr. tively stout, 11-segmented, without distinct J. Milton Campbell for the gift of specimens club. Mandibles prominent, usually asym- relevant to this study. Dr. Herman and my metrical, apices acute, each with one to four wife, Margaret K. Thayer, have reviewed and teeth along mesial margin; molar lobe absent provided useful comments on the manuscript or thin, with microtrichia on mesial surface; for which I am grateful. The SEM photo- prostheca present. Maxilla with lacinia setose graphs were made with the support of Na- and bearing small apical spine; galea with tional Science Foundation grants BMS- dense brush of setae at apex; and palp four- 7412494 and BMS-7502606, and with the segmented, fourth segment about as long as assistance of Mr. Edward Seling. or longer than third and wide or narrow. La- bium with three-segmented palp; with one to PSEUDOPSINAE, SENSU NOVO three premental sclerites. Hypopharynx (including ligula) with one to three pairs of api- Pseudopsini Ganglbauer, 1895. cal lobes, the median pair spatulate. Gular Pseudopsinae ofauthors plus Piestinae: Nanobius, sutures separated, divergent anteriorly and Asemobius, and Zalobius (see Herman, 1975, posteriorly; submentum and gula not sepa- 1977 and references therein). rated by suture. Subocular carina present or DIAGNOSIS: The structure ofthe genital seg- absent. ment, with ninth terga dorsally fused in both Pronotal lateral margin rounded, serrate, sexes and with a fine presumed stridulatory or spinose; dorsal surface broadly elevated. file on each side (figs. 1, 2), will distinguish Protergosternal suture present. Procoxal fis- the Pseudopsinae from all other staphylinid sure broadly or narrowly open with exposed subfamilies. The Pseudopsinae are more eas- trochantin, or fissure closed and trochantin ily recognizable by a combination of the concealed. Postprocoxal lobe present. Pro- coarse punctation ofthe head and pronotum; hypomeron with or without microporous longitudinal carinae or costae of the prono- depression. Procoxae with or without shal- tum, elytra and sometimes of the head; 11- low, carina-delimited groove on mesial sur- segmented unclubbed antennae; and either face. Prosternal process carinate or spini- a comb at the apex of the eighth abdominal form. tergum or a deep incision of the posterior Scutellum with apex exposed behind margin of each elytron. pronotum. Elytra with or without posterior DESCRIPTION: Length 1.8 to 7.0 mm. Body emargination; elytral epipleural ridge pres- moderately broad, moderately flattened dor- ent. One or pair of sclerites present anterior soventrally. Head and pronotum weakly to to mesosternum but posterior to and not as- strongly sculptured with reticulate (large, ir- sociated with mesothoracic spiracular peri- regular, subcontiguous) punctures; prono- tremes (fig. 3). Mesosternum carinate or not. 1982 NEWTON: PSEUDOPSINAE 3

FIGS. 1, 2. 1. Pseudopsis obtusa, female, right dorsolateral view oftergum IX. 2. Nanobius serricollis, female, left dorsolateral view of tergum IX. Scale line = 0.1 mm.

Mesocoxae contiguous or narrowly separated minuta only) 3-3-3. Tarsal empodium bise- by meso- and meta-stemal processes. Hind tose. coxae transverse, contiguous. Abdominal segments II or III to VII each Tibiae with or without longitudinal rows with one or two pairs oflaterosclerites. Terga ofspines. Tarsal formula 5-5-5 or (Pseudopsis II (Pseudopsis) or III to VII with transverse 4 AMERICAN MUSEUM NOVITATES NO. 2743

FIG. 3. Pseudopsis subulata, pro- and meso-thorax, ventral view. subbasal ridge; terga with or without baso- or without transverse subbasal ridge. Ab- lateral ridges or basal, longitudinal carinae; dominal intersegmental membranes with or terga without wing-folding setae or spicules; without pattern; attached to apex or anterior tergum VIII with apical cuticular comb or to apex of preceding segment. median process, or with apex rounded. Spi- Tergum IX ofboth sexes consisting of sin- racles placed in edges of terga II (Pseudopsis) gle large sclerite with fine file on each side or III to VIII. Sternum I represented by strap- (figs. 1, 2), the ridges of the file becoming like sclerite, medially divided or not. Sternite more closely spaced posteriorly. Anterior, II short and fused to sternite III. Sternites II dorsolateral margins of tergum IX with sin- and III with or without median longitudinal uation or shallow emargination which, in carina; sternites III to VII with (Pseudopsis) Pseudopsis at least, marks the opening of a 1982 NEWTON: PSEUDOPSINAE 5 large glandular reservoir. Tergum X ovoid 3(2). Subocular carina present; lateral margin of or triangular, placed in large apical emargi- pronotum coarsely serrate; abdominal nation of tergum IX. terga without longitudinal basal carinae Male. Aedeagus symmetrical or asymmet- ...... Asemobius rical, with or without parameres, with or Subocular carina absent; lateral margin of pronotum spinose; abdominal terga III to without basal piece. Aedeagus in repose with VI each with one basal midlongitudinal parameres facing dorsal surface of beetle. carina ...... Zalobius Female. Coxites circular to oval in cross- section and free, or flattened and fused to one another. Styli present and small, or absent. NATURAL HISTORY Intergonopodal sclerite present or absent. All species, as far as known, occupy cool TAXA INCLUDED: The subfamily as rede- temperate montane forested regions (Her- fined here includes four genera: man, 1975, 1977). Pseudopsis and Nanobius Asemobius Horn (one species, A. caelatus species are most commonly found in moist Horn, from western North America) forest floor litter; some Pseudopsis species Nanobius Herman (one species, N. serri- have also been associated with dung, fungi, collis (LeConte), from western North or mammal nests in the same areas. Zalobius America) and, from the single known record, Asemo- Pseudopsis Newman (30 species as listed bius species are found in wet moss or flood by Herman (1975) from the Holarctic debris along small mountain streams. and Neotropical regions and New Zea- No direct feeding observations have been land) made on pseudopsines. Nanobius gut con- Zalobius LeConte (two species as listed by tents indicate predation on other animals Herman (1977) from western North (Herman, 1977). Cleared specimens of all America) examined species lack the masses of fungal hyphae, spores, algae, or decomposing or- ganic detritus in the gut that characterize fun- KEY TO GENERA OF THE givorous, algophagous or saprophagous PSEUDOPSINAE staphylinid species such as those of the 1. Last segment of maxillary palp less than a subfamilies Piestinae, Osoriinae, and Oxy- third as wide as penultimate segment; pro- telinae (personal observ.). These character- coxal fissure widely open, trochantin well istics and the structure of the mouthparts, exposed; abdominal terga with basolateral especially the mandibles and hypopharynx, ridges, tergum VIII with comb on poste- suggest that pseudopsine adults are carniv- rior margin; abdominal sterna II and III without intercoxal carina . Pseudopsis orous as are other staphylinids with similar Last segment of maxillary palp subequal in mouthparts and known feeding habits (e.g., width to penultimate segment; procoxal the phloeocharine Charhyphus picipennis, fissure narrowly open or closed, trochan- based on personal observ.). tin barely or not visible; abdominal terga Immature stages and life histories are un- without basolateral ridges, tergum VIII known for all Pseudopsinae. Although other without comb on posterior margin; ab- workers and I have made numerous collec- dominal sterna II and III with midlongi- tions ofadults ofall genera except Asemobius tudinal intercoxal carina ...... 2 and have simultaneously searched for larvae, 2(1). Antenna densely pubescent from seventh none that could be attributed to the pseu- segment to apex; procoxal fissure closed, trochantin concealed; mesocoxae sepa- dopsine genera have yet been found. rated by meso- and meta-sternal processes; head with dorsal basal carinae BIOGEOGRAPHY ...... Nanobius Antenna densely pubescent from eighth seg- Herman (1975) has analyzed the distri- ment to apex; procoxal fissure narrowly bution of the widespread genus Pseudopsis. open, trochantin not concealed; meso- He concluded, from the number and diver- coxae contiguous; head without dorsal sity ofspecies and from his proposed cladistic basal carinae ...... 3 history of the genus, that Pseudopsis origi- 6 AMERICAN MUSEUM NOVITATES NO. 2743 nated in the New World, possibly in western in an increased number ofindependently de- North America, with subsequent dispersal by rived apomorphic character states among the some taxa to Eurasia and to South America four genera and would, I believe, represent and New Zealand. The addition of the three less probable cladistic histories ofthe genera. genera Nanobius, Asemobius, and Zalobius A number ofcharacters used in the analysis to the Pseudopsinae as the sister group of that are problematic or ofspecial interest will Pseudopsis reinforces Herman's proposal of be discussed individually. The numbers in a North American origin of the subfamily parentheses refer to character transforma- since these three genera are confined to west- tions of table 1. ern North America. Modified setae (4) vary in structure and in placement, and were subdivided by Herman (1975, 1977) into clubbed and scalelike setae. CLADISTIC ANALYSIS It is not clear whether these setal types are Herman has provided separate cladistic derivable from one another or whether scale- analyses of the species of Pseudopsis (Her- like setae in Pseudopsis and Zalobius are ho- man, 1975) and ofNanobius, Asemobius and mologous or not; for these reasons, modified Zalobius (Herman, 1977). The hypothesis setae of either type are here lumped and here that these four genera together form a treated as a single synapomorphy. monophyletic unit requires a new cladistic A longitudinally carinate or costate head analysis to determine the synapomorphies (6) occurs in Nanobius and in some Pseu- shared by all four genera and to determine dopsis species. The configuration ofcostae or the cladistic relationships among the genera. carinae differs in the two genera and has Characters and character polarities are probably been independently derived. adopted from Herman (1975, 1977) with a A transverse nuchal impression (8) is few changes noted below, and a number of found in all genera except Asemobius which new characters are added (see table 1). The is here assumed to have lost it. Herman most parsimonious branching sequence of (1977) chose the opposite polarity, which the four genera (based on these characters) would require an independent derivation of is given in figure 4. the nuchal impression in the other three gen- This cladistic analysis suggests that all four era. genera share at least 12 synapomorphies. Of The hypopharynx (21) ofmost other staph- these, the single most significant one is the ylinids is broadly bilobed. A hypopharynx newly discovered file located on each side of very similar to that of Pseudopsis and Ase- the ninth tergum in all species of Pseudop- mobius (21A) is found in some Phloeochar- sinae (character 43; figs. 1, 2). This file has inae (sensu lato) and is here considered to be been found in no other staphylinids. Virtually independently derived or, possibly, synapo- all other synapomorphies shown for the four morphic for these two subfamilies. pseudopsine genera in figure 4 are known to A mesial procoxal groove (32) of the type occur elsewhere in the Staphylinidae, al- found in some Pseudopsinae occurs in some though many are uncommon and some, such genera of many staphylinid subfamilies, as as character 33, are probably not homologous noted by Herman (1977). The polarity ofthis with apparently similar character transfor- character transformation is thus open to mations found elsewhere. In combination doubt. with the unique file, I think these synapo- Sclerites in the connecting membrane be- morphies provide strong support for the tween pro- and meso-sterna (33) occur in sev- monophyly of the Pseudopsinae as defined eral staphylinid subfamilies and in the Sil- here. phidae-Silphinae. The location of these The relationships ofthe genera ofthe Pseu- sclerites immediately anterior to the meso- dopsinae shown in figure 4 represent the most sternum and posterior to, and not associated parsimonious arrangement possible using the with, the mesothoracic spiracular sclerites is characters and character polarities oftable 1. apparently found only in the Pseudopsinae, Alternative branching patterns would result in Apateticini of Piestinae and in the Silphi- 1982 NEWTON: PSEUDOPSINAE 7

TABLE 1 Relative Plesiomorphy and Apomorphy of Characters Used for Cladistic Analysis of Pseudopsinaea Plesiomorphic (P) Apomorphic (A) 1. Pronotum and elytra without carinae or costae Pronotum and elytra longitudinally carinate or costate 2. Punctation of head and pronotum punctiform Punctation of head and pronotum reticulate 3. Dorsum of head, pronotum and elytra clean, Dorsum of head, pronotum, and elytra with without accumulations of mud and debris accumulations of mud and debris 4. Setae unmodified Some body setae modified 5. Setae large, visible with dissecting microscope Setae small, visible only with compound microscope at ca. 400X 6. Head not longitudinally carinate or costate Head longitudinally carinate or costate *7. Neck absent Neck present as lateral constriction and dorsal transverse impression 8. Neck distinguished dorsally by transverse Neck not distinguished dorsally impression *9. Antenna densely pubescent from fourth, fifth, or Antenna densely pubescent from seventh segment to sixth segment to apex apex 10. (=9A) Antenna densely pubescent from eighth segment to apex

11. Subocular carina absent Subocular carina present 12. Epistomal suture evident externally as groove Epistomal suture evident externally as ridge 13. Labrum without peglike setae on anterior margin Labrum with row of peglike setae on anterior margin 14. Epipharynx without globosetae Epipharynx with globosetae present, forming median patch 15. (=14A) Epipharyngeal globosetae present, forming transverse row on anterior edge *16. Mandibles without subapical teeth Mandibles with one or more large subapical teeth on mesial edge *17. Mandibles without serrate edges Right mandible with dorsal, subapical serrate edge *18. Mandibles with large molar lobes bearing mesial Mandibles with thin molar lobes bearing mesial grinding surfaces microtrichia *19. (=18A) Mandibles without interacting molar lobes *20. Maxillary palp with fourth segment subequal in Maxillary palp with fourth segment less than a third width to third as wide as third segment *21. Hypopharynx broadly bilobed at apex Hypopharynx with two pairs of lobes, the median pair spatulate and subcontiguous at base, the lateral pair acute *22. (=21A) Hypopharynx with six lobes, both lateral pairs acute *23. (=21A) Hypopharynx with median pair of lobes largely fused to lateral pair *24. Labium with palpigers free Labium with palpigers completely fused to one another 25. Mentum not carinate Mentum with midlongitudinal carina 26. Gula not carinate Gula carinate 8 AMERICAN MUSEUM NOVITATES NO. 2743

TABLE 1-(Continued) Plesiomorphic (P) Apomorphic (A) 27. Pronotum without deep pits Pronotum with deep pits near anterior and posterior margins 28. Prohypomeron unmodified Prohypomeron with microporous depression *29. Procoxal fissure widely open, trochantin well Procoxal fissure narrowly open, trochantin barely exposed exposed *30. (=29A) Procoxal fissure closed for most of its length, trochantin concealed *31. Prosternal process cariniform Prosternal process spiniform 32. Procoxa without carina-delimited mesial groove Procoxa with shallow, carina-delimited mesial groove *33. Antemesosternal sclerites absent One or pair of antemesostemal sclerites present (fig. 3) *34. Mesosternum not carinate Mesosternum midlongitudinally carinate *35. Mesocoxae narrowly separated by mesosternal or Mesocoxae contiguous meso- and metasternal processes 36. Elytron with posterior margin entire Elytron with lateral portion of posterior margin deeply incised 37. Abdominal terga without basolateral ridges Abdominal terga with basolateral ridges 38. Abdominal terga without longitudinal carinae Abdominal terga each with partial median longitudinal carina 39. (=38P) Abdominal terga each with several partial longitudinal carinae 40. Abdominal tergum VIII with unmodified posterior Abdominal tergum VIII with comb on posterior margin margin 41. (=40P) Abdominal tergum VIII with posterior margin biemarginate 42. Abdominal tergum IX without openings of large Abdominal tergum IX with openings of pair of large gland reservoirs gland reservoirs anteriorly at dorsolateral margin *43. Abdominal tergum IX without "stridulatory" file Abdominal tergum IX with fine "stridulatory" file on each side (figs. 1, 2) *44. Some of abdominal terga with patches of wing- Abdominal terga without wing-folding setae or folding setae or spicules spicules *45. Abdominal segments III-VI with a single pair of Abdominal segments III-VI with two pairs of laterosclerites per segment laterosclerites per segment *46. Abdominal intersegmental membranes between Abdominal intersegmental membranes attached to segments III-VII attached to apex of preceding inner surface of preceding segment anterior to its segment apex *47. Abdominal intersegmental membranes of segments Abdominal intersegmental membranes without III-VII with pattern of minute sclerites pattern *48. Abdominal sternites II and III with midlongitu- Abdominal sternites II and III not midlongitudinally dinal intercoxal carina carinate 49. Tergum IX of male with lateral portion compressed Tergum IX of male with lateral portion cylindrical and triangular in lateral view, not reaching and produced well beyond apex of tergum X beyond apex of tergum X 1982 NEWTON: PSEUDOPSINAE 9

TABLE 1-(Continued) Plesiomorphic (P) Apomorphic (A) 50. Aedeagus with U-shaped basal piece Aedeagus without basal piece *51. Tergum IX of female divided dorsally by tergum X Tergum IX of female broadly fused dorsally anterior to tergum X 52. Female with intergonopodal sclerite Female without intergonopodal sclerite 53. Coxites separate Coxites fused basally 54. Coxites cylindrical or slightly flattened Coxites broadly flattened 55. Stylus elongate Stylus reduced, knoblike 56. Stylus present Stylus absent a Those marked with an asterisk (*) were not used by Herman (1975, 1977). dae-Silphinae. Tentatively, I am assuming and it is perhaps possible that such reservoirs that these sclerites have been independently exist in the other genera of Pseudopsinae but derived in the three taxa cited, and that these are not detectable without analysis of better sclerites are not homologous with the more preserved specimens. It is also possible that anterior sclerites associated with mesotho- gland reservoirs have been lost in the other racic spiracular sclerites in the Tachyporini pseudopsine genera. Based on the evidence and other Staphylinidae. available at present, I conclude that the gland The presence of openings of large gland openings are probably independently derived reservoirs in the ninth tergum (42) of Pseu- in Pseudopsis and in the Oxytelinae. dopsis is unusual and is apparently found The fine file (43) at each side ofabdominal elsewhere only in the subfamily Oxytelinae tergum IX was overlooked by Herman (1975, (Herman, 1970). This character was consid- 1977), but is found in all examined pseudop- ered synapomorphic for the two taxa by Her- sine species. A careful search among other man (1975). It should be mentioned, how- staphylinids has produced no comparable ever, that the openings in Pseudopsis are at structure. The file resembles known stridu- the anterior edge of tergum IX, whereas in latory files, except for the great change in the Oxytelinae the openings are in the terga spacing between individual ridges from the or in a deep invagination ofthe anterior mar- widely spaced anterior ridges to the ex- gin of the terga. Furthermore, the configu- tremely close and fine posterior ridges (see ration of the ninth and tenth terga is quite figs. 1, 2). The file is in such a position that different in the Oxytelinae (ninth terga sep- it would rub against the apex of the inflexed arated dorsally by tenth tergum and "strid- sides of tergum VIII as the ninth segment is ulatory" files absent) and in Pseudopsis (ninth in- or evaginated. An alternative function of terga broadly fused dorsally anterior to tenth the file, suggested by the close proximity of tergum, and "stridulatory" file present on the gland opening of Pseudopsis (42) to the each side). The configuration of ninth and anterior end of the file, is that the file might tenth terga is identical in Pseudopsis and in somehow aid in the dispersal or application the other three genera ofPseudopsinae which ofthe gland secretion. Since the file is as well apparently lack the glandular reservoirs. developed in the other pseudopsine genera These genera do, however, have an inflection (which apparently lack glands) as in Pseu- in the anterior margin of the ninth tergum dopsis, this possible function seems unlikely. similar in placement and appearance to the Those workers with access to living material point at which each gland reservoir of Pseu- of this unfortunately rare and spottily dis- dopsis opens. The gland reservoirs of some tributed subfamily should attempt to deter- species of Pseudopsis are quite difficult to mine the functional significance of both the detect (Herman, in litt., and personal observ.) files and glands. Pseudopsis Nanobius AsemobiuJs Za/obius 44 44 8 44 64 5 11 14 13 1114 64 l147 15 17 22 19 20 264 23 244 27 244 324 30 25 354 39 264 37 41 38 40 454 454 42 504 49 48 524 564 504 524 Si 53 54 10 564 12 35-4 46

I 3 I 9 I 28 I 29 I 31 I 324 I 34 I 36

I I I I 2 I I 7 I 14 I 16 I 18 I 21 I 33 I 43 I 44 I 51 4 I 55

FIG. 4. Cladistic analysis of the genera of Pseudopsinae. Numbers refer to character transformations listed in table 1. Half-filled squares indicate that only some species of the genus show the apomorphic condition of that character. A triangle following a number indicates a character transformation that occurs independently on this cladogram. 1982 NEWTON: PSEUDOPSINAE 11

Many subfamilies of staphylinids have convergence in dorsal appearance of male members with either one or two pairs of lat- and female terminalia is not uncommon in erosclerites per segment (45). I have followed the Staphylinidae and has occurred in a few Herman (1970) in assuming that one pair is other families such as the Hydraenidae and plesiomorphic, because it is prevalent among Ptiliidae. primitive staphylinids and allied families and An intergonopodal sclerite (52) ofthe ovi- is most readily derivable from the condition positor is found in many staphylinids as (no laterosclerites) found in other staphyli- noted by Herman (1977), and in many other noid families such as the Silphidae and Coleoptera (Tanner, 1927). I consider such Leiodidae. Although coded as derived for sclerites homologous and so reverse Her- Zalobius, only two segments have two pairs man's polarity assignment. of laterosclerites while the remainder have It should be noted that in spite of the ad- one pair. dition of numerous new characters and in Abdominal intersegmental membranes be- spite ofa few reversals ofcharacter polarities tween segments III and VIII (46) are usually selected by Herman (1977), the branching attached at the apex of each preceding seg- sequence among the genera Nanobius, Ase- ment in Coleoptera and in many Staphylin- mobius, and Zalobius in figure 4 is the same idae, but are attached preapically in many as that in Herman's most-parsimonious staphylinid subfamilies. The same mem- cladogram (Herman, 1977, fig. 2) ofthe same branes in staphylinids usually have a distinct genera. The cladistic analysis of the species pattern of minute sclerites (47); this pattern or species groups ofPseudopsis given by Her- is reduced or absent in many subfamilies and man (1975) would be virtually unaffected by genera, especially those in which the attach- the changes or additions made here. One ex- ment is preapical. ception would be the addition of an aut- The U-shaped sclerite (50) surrounding the apotypic trait (tarsal formula 3-3-3 rather basal orifice ofthe aedeagus was assumed by than 5-5-5) to the Pseudopsis minuta branch; Herman (1975, 1977) to be independently this tarsal reduction was overlooked by Her- derived wherever it occurs. A similar sclerite man (1975). occurs in the same position in many staphylinids (e.g., some Omaliinae, Proteininae, and Paederinae as well as some Pseudopsi- SUBFAMILY POSITION nae) and in members of other staphylinoid Herman presented analyses of possible families such as the Silphidae, Leiodidae, and subfamily relationships for Pseudopsis (1975) Ptiliidae. I know ofno reason not to consider and for the three genera Nanobius, Asemo- this sclerite as a homologue ofthe basal piece bius, and Zalobius (1977). He concluded that ofother Coleoptera, which has been reduced Pseudopsis was the sister group of all the in all the Staphylinoidea and lost in many Oxytelinae, based exclusively on the presence taxa ofthis superfamily. Thus I have reversed and position of exit of a pair of abdominal Herman's polarity of this character change. gland reservoirs. Similarities to other In most primitive Staphylinidae, in closely subfamilies, especially the Phloeocharinae allied families such as the Silphidae and and Piestinae, were noted. A more detailed Leiodidae, and in most other Coleoptera the analysis of characters shared by Nanobius, sclerites ofabdominal segments IX and X are Asemobius, and Zalobius with various other strongly sexually dimorphic. In females, staphylinid taxa was given. These three gen- these sclerites form the ovipositor (Tanner, era were found to share more derived char- 1927). Tergum IX of females is usually rep- acters (carinate body, epipharyngeal globo- resented by a pair of sclerites (paraprocts) setae, dorsally fused ninth tergum) with that are separated by tergum X (proctiger). Pseudopsis than with any other staphylinids. The dorsal fusion of these sclerites to form However, these synapomorphies are absent a single tergum IX anterior to tergum X (5 1) in the Oxytelinae, which share with Pseu- results in a configuration similar to that of dopsis the glandular openings in the ninth males, which in most Coleoptera have a sin- tergum. Herman concluded that this contra- gle tergum IX anterior to tergum X. Such a diction prevented a satisfactory resolution of 12 AMERICAN MUSEUM NOVITATES NO. 2743 relationships of these taxa; he tentatively re- gut. The sister group of the Oxytelinae is tained Pseudopsis as sole representative of probably best sought among a portion or all the Pseudopsinae and Pseudopsinae as the of the Piestinae and Osoriinae. Since the sister group ofthe Oxytelinae, while retaining Piestinae and Osoriinae have no trace of Nanobius, Asemobius, and Zalobius in the ninth tergal glands, I would conclude that subfamily Piestinae even though no synapo- ninth tergal glands have been independently morphies that would justify such placement evolved in the Oxytelinae and in some or all had been found. the Pseudopsinae. In the present work I have pointed out, in Some characteristics of many Pseudopsi- the cladistic analysis of figure 4, a number of nae such as pronotal and elytral carinae, additional synapomorphies shared by Pseu- scalelike setae and detritus accumulation oc- dopsis, Nanobius, Asemobius, and Zalobius. cur in a variety of staphylinids as noted by Most important among these is the presumed Herman (1977), and I agree with Herman stridulatory file on each side of abdominal that such occurrences have probably arisen tergum IX (figs. 1, 2), unique in the Staphy- independently in or within the subfamilies linidae. In my view these synapomorphies in in which they are found. All of these char- combination firmly establish that the four acteristics are found in a newly discovered genera together form a monophyletic unit, genus from Australia which is described and here called the Pseudopsinae (sensu novo). discussed in detail elsewhere (Newton, 1982). It follows that either the ninth tergal glan- The piestine genus Euphanias also has elytral dular openings of Pseudopsis and the Oxy- carinae and scalelike setae and has occasion- telinae are a convergent development, or that ally been considered to be allied to Pseudopsis the glandular openings have had a wider dis- (e.g., by Mulsant and Rey, 1856). Evidence tribution in the past and have been lost in that both Euphanias and the new Australian Nanobius, Asemobius, and Zalobius, and genus belong in the Oxytelinae and are closely possibly in other taxa. Ifthe glands have been related to Syntomium is presented elsewhere lost in Nanobius, Asemobius, and Zalobius, (Newton, op. cit.), and the similarities to the or if they are present in those genera but not Pseudopsinae are considered to be a result of detected with available material, then a sis- convergence. ter-group relationship of the Pseudopsinae If the Pseudopsinae are excluded from the (sensu novo) and Oxytelinae would still be group of subfamilies including the Piestinae, possible. Oxytelinae, and Osoriinae, then where does Evidence other than the presence and po- the subfamily belong in relation to other sition of ninth tergal gland openings that staphylinids? Herman (1975) pointed out the would support a sister-group relationship of near identity of the hypopharynx in Pseu- the Pseudopsinae and Oxytelinae is very dopsis and in some Phloeocharinae, and the scarce. Of the 12 synapomorphies of the presence ofglobosetae in Rimulincola which Pseudopsinae shown in figure 4, only two is usually placed in the Phloeocharinae. Un- (nos. 44 and probably 55) also characterize fortunately, this small subfamily of fewer all the Oxytelinae. These two apomorphies than a dozen genera is poorly known. Neither are known to have been independently de- the subfamily as a whole nor many of the rived in many other staphylinid subfamilies included genera have been adequately char- and are perhaps the weakest of all detected acterized, and the limits ofthe subfamily are synapomorphies ofthe Pseudopsinae. In con- unstable with some included genera (Rimu- trast, the Oxytelinae share a number of less lincola, Olisthaerus, Giulianium) probably common synapomorphies with the subfam- belonging elsewhere. Any detailed consider- ilies Piestinae and Osoriinae: divided ninth ation ofthe relationship ofthe Pseudopsinae terga in both sexes; similar mouthparts in- to Phloeocharinae is, therefore, beyond the cluding numerous multifid or plumose setae scope of the present work. of the labrum, mandibular prostheca and Immature stages have not yet been de- galea; and saprophagous adult (and larval) scribed for any member ofthe Pseudopsinae. feeding habits with resultant masses of de- [Ac ording to comments ofP. M. Hammond composed plant material, fungi, etc. in the (in litt.), larvae of Pseudopsis were reared by 1982 NEWTON: PSEUDOPSINAE 13

W. 0. Steel and are in the British Museum LITERATURE CITED (Natural History), but these cannot now be Ganglbauer, Ludwig located.] Discovery (or rediscovery) of the 1895. Die KAfer von Mitteleuropa. Band II. larva of any pseudopsine should provide a Familienreihe Staphylinoidea. 1. Theil: test of the alternative hypotheses of subfam- Staphylinidae, Pselaphidae. Vienna, 880 ily relationships just discussed. If Herman's PP. (1975) hypothesis that Pseudopsis is the sister Hammond, Peter group of the Oxytelinae is correct, and/or if 1975. The phylogeny of a remarkable new ge- the placement of Nanobius, Asemobius, and nus and species of gymnusine staphylinid (Coleoptera) from the Auckland Is- Zalobius in the Piestinae is correct, then we lands. Jour. Ent. (B), vol. 44, no. 2, pp. would expect pseudopsine larvae to possess 153-173. several synapomorphies shared by larvae of Herman, Lee H., Jr. the Piestinae, Oxytelinae and Osoriinae. 1970. Phylogeny and reclassification of the These include saprophagous feeding habits, genera of the rove-beetle subfamily Ox- a transverse truncate ligula (except the pies- ytelinae ofthe world (Coleoptera, Staph- tine genus Trigonurus), bifid, multifid, or ylinidae). Bull. Amer. Mus. Nat. Hist., more highly modified mandibular apices, vol. 142, pp. 343-454. and a cervicosternum of a single sclerite (see 1975. Revision and phylogeny of the mono- Newton, 1982). In contrast, if pseudopsine generic subfamily Pseudopsinae for the world (Staphylinidae, Coleoptera). Ibid., larvae are predatory as suggested by adult vol. 155, pp. 241-318. evidence plus the usual correlation of adult 1977. Revision and phylogeny of Zalobius, and larval feeding habits in Staphylinidae, Asemobius and Nanobius, new genus then the larval mouthparts would be ex- (Coleoptera, Staphylinidae, Piestinae). pected to resemble those of relatively gen- Ibid., vol. 159, pp. 45-86. eralized predatory staphylinid larvae such as Mulsant, E., and Claudius Rey occur in the Phloeocharinae and Omaliinae. 1856. Description d'un col6optere inedit con- We might then find an acute mandible with stituant un genre nouveau voisin du G. serrate inner edge and at most a single sub- Pseudopsis (famille des brachelytres). apical tooth, a digitiform ligula, and perhaps Ann. Soc. Linn. Lyon, ser. 2, vol. 3, pp. 7-11. a three-segmented cervicosternum as is usual Newton, Alfred F., Jr. among staphylinid larvae. 1982. New genus and species of Oxytelinae At present I tentatively leave the Pseudop- from Australia, with description of its sinae as a subfamily of uncertain position larva, systematic position and phylo- within the Staphylinidae, and suggest that the genetic relationships (Coleoptera, Sta- sister group of the subfamily be sought phylinidae). Amer. Mus. Novitates no. among the Phloeocharinae or allied subfam- 2744, pp. 1-24. ilies (see Hammond, 1975) rather than Tanner, Vasco among the Piestinae, Oxytelinae, and Oso- 1927. A preliminary study of the genitalia of riinae. female Coleoptera. Trans. Amer. Ent. Soc., vol. 53, pp. 5-50, pls. 2-15.