AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2764, pp. 1-18, figs. 1-49, tables 1-3 June 23, 1983 Eppelsheimius: Revision, Distribution, Sister Group Relationship (Staphylinidae, Oxytelinae) LEE H. HERMAN' ABSTRACT Eppelsheimius is a small genus of beetles that pirazzolii and E. miricollis, that are distinguished occurs in arid regions from northern Africa to by many characters. Both species are variable. The southwestern Asia. The species share characters genus and species are described and illustrated and with Planeustomus, Manda, and Bledius. Evi- their distributions described. One species, E. per- dence is presented that Bledius and Eppelsheimius sicus, is newly synonymized with E. pirazzolii. are sister groups. The genus has two species, E. INTRODUCTION The present paper was stimulated by a in a forthcoming paper on Bledius (Herman, search for the sister group ofBledius. Earlier, in prep.). Ultimately, several rearrangements but without supporting characters, Herman in the classification of the Oxytelinae will be (1970, p. 354) presented two groups ofgenera required. as the sister group of Bledius. One of these Eppelsheim (1885) described pirazzolii in groups, the Carpelimus lineage, includes Oncophorus. A second species, miricollis, was Carpelimus, Apocellagria, Trogactus, Thi- added by Fauvel (1898); both were from Tu- nodromus, Xerophygus, Ochthephilus, nisia. In 1915, Oncophorus was discovered Mimopaederus, Teropalpus, Pareiobledius to be a homonym of a genus of Mal- and Blediotrogus; the other, the Thinobius lophaga and a genus of "worms" of indeter- lineage, includes Thinobius, Sciotrogus, and minate placement. Bernhauer (1915) pub- Neoxus. My own subsequent studies and those lished a replacement name, Eppelsheimius, of others (Hammond, 1975, 1976; Newton, and a few years later Champion (1919), un- 1982) have pointed to a number of inade- aware of Bernhauer's change, proposed On- quacies in my phylogeny of the Oxytelinae cogenys to supersede Oncophorus. Koch in a (Herman, 1970, p. 354). One problem, the series of papers (1934, 1936, 1937) estab- relationship of Bledius to Epplesheimius, is lished the occurrence of the genus in many addressed herein; some others are discussed parts of northern Africa east to Iraq. Eppel- ' Curator, Department of Entomology, American Museum of Natural History. Copyright © American Museum of Natural History 1983 ISSN 0003-0082 / Price $1.95 2 AMERICAN MUSEUM NOVITATES NO. 2764 /I I FIG. 1. Eppelsheimius pirazzolii. sheimius was reported in southwested tIran species, persicus. The genus was redescribed when Scheerpeltz (1958) described a third by Herman (1970, p. 369). 1983 HERMAN: EPPELSHEIMIUS 3 TABLE 1 Polarity of Some Character States of the Oxytelinae Character Plesiomorphic Apomorphic Labral setae Short, slender Long, stout (figs. 23, 36) Hypopharynx Apical lobe entire Apical lobe bifid Mandibular mola Well developed Reduced, nearly absent Antennae Not geniculate Geniculate (fig. 26) Gular sutures Separated Confluent (fig. 30) Neck Present, broad Absent (figs. 1, 14, 41) Clypeus Large, at same level as dorsum Narrow, depressed below level of dorsum Eyes Restricted to side of head Extend onto ventral surface of head Protibiae Slender Expanded (fig. 6) Dorsal seta of epipharyngeal lobe Absent Present (figs. 23, 36) Elytral suture Straight Dehiscent (fig. 1) When Eppelsheim described the genus he geniculate antenna (fig. 26; first segment elon- placed it near Manda (cited as Acrognathus) gate and the second segment posteriorly flex- and Planeustomus (cited as Compsochilus) ile on first). Some ofthese characters are found because of unspecified similarities. Herman in other oxyteline genera but not in the com- (1970, p. 355) found support for this hy- bination found for Bledius and Eppelsheim- pothesis in the presence in the three genera ius. The polarity ofthe character states given ofelongate, membranous epipharyngeal lobes in table 1 was derived by their relative dis- (fig. 23). tribution in other subfamilies, particularly the Although I said (Herman, 1970, p. 355) Omaliinae, Piestinae, and Osoriinae. The that the species of Eppelsheimius share the apical lobe of the hypopharynx is entire, the enlarged, membranous, epipharyngeal lobes mandibular mola well developed, the clypeus with Manda and Planeustomus, dissection of large and at the same level as the dorsum, the species ofEppelsheimius reveals that only the eyes restricted to the side ofthe head, the E. pirazzolii has enlarged lobes (figs. 10, 1.1, antennae normal (not geniculate), the gular 23); miricollis has shorter but well-developed sutures separated, the neck broad and weakly lobes (fig. 36). However, the epipharyngeal developed, the protibiae slender, the dorsal lobes of the species of the three genera do seta ofthe epipharyngeal lobe absent, and the have apically bifurcate or multifurcate cutic- elytral suture straight in most of the genera ular processes and the labrum of each has of the Osoriinae, Piestinae, and Omaliinae. long stout labral setae (figs. 23, 36). Both of Variation from these conditions in the Oxy- these features seem to be derived but I find telinae are derived (table 1). no others that the three genera share. No other genera ofthe Oxytelinae have the Manda and Planeustomus share a narrow expanded protibiae. The configuration of the clypeus that is depressed below the level of confluence of the gular sutures as found in the dorsum, eyes that extend onto the ventral Bledius and Eppelsheimius (fig. 30) is also surface ofthe head, reduction to near absence found in the genera of the Carpelimus and of the mandibular mola, and the bifid apical Thinobius lineages (see Herman, 1970, p. lobes of the hypopharynx. The first two of 354). Dehiscent elytra are found in Thinobius these four states are unique in the family. and in some species of Platystethus. Species Manda and Planeustomus are a well-defined ofCoprophilus and one ofManda lack a neck. and supported group. Geniculate antennae are found only in Ble- Eppelsheimius and Bledius share six de- dius and Eppelsheimius. The dorsal epipha- rived features: confluent gular sutures (fig. 30), ryngeal seta is found in one species ofManda. absence of a neck (fig. 1), expanded protibiae The hypothesis that Eppelsheimius is the (fig. 6), a dorsal seta on the epipharyngeal lobe sister taxon of Bledius rather than of Manda (figs. 23, 36), dehiscent elytra (fig. 1), and and Planeustomus is supported by the fact 4 AMERICAN MUSEUM NOVITATES NO. 2764 that it shares six apomorphic features with them than do some genera ofthe Oxytelinae. Bledius but only two with the two other gen- The species within each group may be ex- era. Except as noted in the preceding para- tremely difficult to distinguish (Herman, graph, Eppelsheimius shares no other appar- 1972, 1976, in press, and in prep.). ently derived features with other genera that By contrast, although Eppelsheimius might are not shared by most genera ofthe subfam- share the saline habitat with Bledius, it has ily. This hypothesis along with the relation- a more restricted geographical and ecological ship of Eppelsheimius and Bledius to other distribution and is a relatively minuscule ge- oxyteline genera is discussed further in a pa- nus. However, the two species of Eppel- per dealing with the phylogeny of Bledius sheimius are separated by as many characters (Herman, in prep.). The present paper is a as are some species groups of Bledius. This first step toward understanding the relation- divergence in Eppelsheimius coupled with the ships of the species of Bledius and of the ge- intraspecific variation leads me to wonder nus to other genera of the subfamily. Many whether the two species actually represent questions remain. What is the sister group to complexes of subtly distinguishable species. Bledius and Eppelsheimius? How are other I was unable to detect discrete variation with- genera, such as Aploderus, Pareiobledius, in the two recognized species with samples Blediotrogus, and Xerophygus, that share available to me. some derived characters with Bledius and Information on the habitat of species of Eppelsheimius related to them? What are the Eppelsheimius has not been published or in- relationships ofthe genera ofthe Carpelimus cluded on the specimen labels. Since all the and Thinobius lineages to Bledius and Ep- localities from which the species have been pelsheimius and to other genera of the taken are near salt water-either near the sea subfamily? These and similar problems are or near inland salt lakes, flats, and marshes beyond the scope of the present paper. it is probable that they live in saline habitats. Based on similarities to Bledius, that is the Although the species have been collected only expanded protibiae, the absence of a neck, near salt marshes of northern Africa, along and the subcylindrical body, I expect that the coasts ofthe Mediterranean and Red seas, species of Eppelsheimius make burrows in and at a few probably saline habitats in Saudi the soil. Eppelsheimius, as does Bledius, may Arabia, Iraq, Iran, and the USSR (fig. 2), they use their elongate mandibles to excavate bur- are likely to occur near the salt lakes and salt rows. The absence ofa neck permits the great- marshes that are found across arid parts of er range of movement needed to pick up and Asia to China. The species are attracted to deposit sand. The presence of a neck restricts light. the degree of rotation of the head. The ex- In addition to querying the extent of the panded protibiae are not used by Bledius to geographical range, further collection can de- construct a burrow, and it is unlikely that termine where the species live, whether they Eppelsheimius uses them in that way. Species make burrows, and if so how, what they eat, ofboth genera may use them to facilitate agile where the larvae live, and what their anatom- movement in the burrow. ical features are. The collection sites of Ep- IfEppelsheimius and Bledius are sister taxa, pelsheimius are widely scattered (fig.