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 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.

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. 2) but several interesting comparisons can be made. near the Chott Djerid, a large salt marsh in Bledius has 420 species found in nearly all Tunisia, 141 of the 217 specimens studied parts of the world. They burrow into the soil were collected. Chott Djerid would be an ap- adjacent to fresh and saline water of rivers, propriate place to try to answer the preceding lakes, ponds, and the ocean; some live in tem- questions and perhaps others. porarily wet soil that is separated from stand- ing water. They feed on algae. Bledius is high- ACKNOWLEDGMENTS AND ly variable anatomically; the degree of ABBREVIATIONS variation permits recognition of many species groups based on numerous characters. These Specimens used in this study were bor- groups often have more characters separating rowed from the individuals and institutions 1983 HERMAN: EPPELSHEIMIUS 5

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c) 6 AMERICAN MUSEUM NOVITATES NO. 2764 listed below. Abbreviations preceding the presence of five tarsomeres and the submen- name of each institution are used in the text tal processes the diagnostic features cited in to indicate the location of specimens. The the preceding paragraph are found in all or name of the person who lent the material some of the species of Bledius. The species follows the name of the institution. I grate- of Eppelsheimius are more "loosely con- fully thank them for their assistance. I es- structed" than are those of Bledius. The me- pecially thank Dr. Heinrich Schonmann, who sotibia of Bledius has two rows of spinelike lent types, Miss Beatrice Brewster, who trans- setae; that of Eppelsheimius has one row of lated articles from French and German, and elongate setae among the scattered ones. The Miss Joan Whelan, who took the scanning postprocoxal lobe is larger in Bledius than in electron micrographs. Eppelsheimius. The genital appendages ofthe AMNH, American Museum of Natural History. females of Bledius consist of a pair of elon- BMNH, British Museum (Natural History), Lon- gate, flattened sclerites; those ofEppelsheim- don; Mr. Peter Hammond. ius are paired, elongate, transversely divided FMNH, Field Museum of Natural History, Chi- (into a coxite and a valvifer) and compressed cago; Dr. Larry Watrous. (fig. 33). HCC, Dr. H. Coiffait, Toulouse. DESCRIPTION: Length 3.1 to 6.8 mm. IRSN, Institut Royal des Sciences Naturelles, Color pale to dark reddish brown with yel- Brussels; Dr. Leon Baert. lowish brown elytra. Form slender and sub- MGF, Museum G. Frey, via. Dr. Gerhard Scherer. cylindrical. Body sparsely pubescent. MHMV, Naturhistorisches Museum Wien, Vi- and enna; Dr. Heinrich Sch6nmann. Head (figs. 14, 41) sparsely pubescent MNHN, Museum National d'Histoire Naturelle, punctate; lateral margins gradually conver- Paris; Miss Nicole Berti. gent from eyes toward base; neck absent, MSNM, Museo Civico di Storia Naturale, Milano; postocular transverse groove absent. Clypeal Dr. Carlo Leonardi. length variable. Epistomal suture straight; su- MTC, Marc Tronquet, Paris. ture approximately even with anterior mar- NHMB, Naturhistorisches Museum Basel, Basel; gin ofsupra-antennal ridge. Eyes (figs. 14, 41) Dr. M. Brancucci. slightly to moderately protruding from sides of head, not extending onto ventral surface. EPPELSHEIMIUS BERNHAUER Supra-antennal ridge small. Anterior and Figures 1-49 dorsal tentorial arms present. Antenna (fig. Eppelsheimius Bernhauer, 1915, p. 270. Scheer- 26) with first segment elongate and genicu- peltz, 1958, pp. 14-17. Herman, 1970, p. 369. late; segments 9, 10, and 11 expanded to form TYPE SPECIES: Eppelsheimius pirazzolii (Ep- loose, well-defined club. Gular sutures (fig. pelsheim). 30) confluent for most of length, sutures Oncophorus Eppelsheim, 1885, p. 46; preoccu- sharply divergent at base. Submentum (figs. pied. TYPE SPECIES: Oncophorus pirazzolii Ep- 13, 16, 42, 45) with short to long, slender to pelsheim. stout spiniform process on anterior lateral Oncogenys Champion, 1919, p. 154. TYPE SPECIES: edge. Labium (fig. 21) with trapezoidal men- Oncogenys pirazzolii (Eppelsheim). tum; palpal segments subequal. Hypophar- DIAGNOSIS: Eppelsheimius is separated ynx as in figures 3 and 4. Labrum (figs. 23, from other genera of the Oxytelinae by the 36) fused, without midlongitudinal fissure; presence of an epistomal suture (fig. 14), ge- surface polished and with long, stout setae niculate antennae (fig. 26), open procoxal fis- near anterior margin; posterior margin with sure (figs. 12, 46), spiniform submental pro- long, posteriorly directed, internal strut on cesses (figs. 13, 16, 42, 45), confluent gular lateral edge; ventroposterior median surface sutures (fig. 30), dehiscent elytra suture (fig. with triangular internal strut. Epipharynx 1), absence of elytral epipleural ridges, pres- (figs. 23, 36) with short to long membranous, ence of five tarsomeres, and expanded, spi- anteriorly directed lobe; lobe with long, slen- nous protibiae (fig. 6). der, apically bifurcate process on mesial mar- The genus is similar to Bledius in form and gin; epipharyngeal lobe with long, stout setae shares with it many characters. Except for the near base of dorsal surface. Maxilla as in fig- 1983 HERMAN: EPPELSHEIMIUS 7

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FIGS. 3-7. Eppelsheimius pirazzolii. 3. Hypopharynx, 270X. 4. Hypopharynx, 674X. 5. Scutellum, 220X. 6. Protibia and tarsus, 105X. 7. Metatibia and tarsus, 121 X. ure 8; fourth segment ofmaxillary palpus stout 22, 38) with short to long spine on anterior (fig. 9); galea (figs. 27, 35) with row of stout margin ofdorsal surface. Mandibles (figs. 25, spinelike setae on dorsal surface; stipes (figs. 47) edentate, long, curved mesially, and 8 AMERICAN MUSEUM NOVITATES NO. 2764

FIGS. 8-1 1. Eppelsheimiuspirazzolii. 8. Maxilla, 206X. 9. Maxillary palpus, fourth segment, partially collapsed, 479X. 10. Epipharynx, ventral surface, 1 56X. 1 1. Epipharynx, ventral surface, 491 X. touching at apices when closed, but not cross- cess (figs. 12, 46) elongate, carinate and ven- ing. troposteriorly directed; ventral edge straight. Pronotum (figs. 17, 44) longer than wide; Scutellum with apex slightly exposed to con- lateral margins gradually curved to nearly cealed under pronotum; surface with elongate straight or strongly sinuous with bulge near oval impression (fig. 5). Elytra elongate; su- basal third; surface sparsely pubescent and ture dehiscent (fig. 1); posterior margin with- punctate with shallow midlongitudinal out membranous lobe; epipleural ridge ab- groove. Pronotal lateral marginal bead pres- sent; surface sparsely punctate and pubescent ent (fig. 12) or absent (fig. 46). Prohypomeron and without longitudinal striae. Mesostemal impressed anterior to coxae; postprocoxal (fig. 15) process short to moderately long and portion present as small (fig. 46) to large (fig. extending between coxae; process broad and 12) lobe. Protergostemal (figs. 12, 46) suture tapered to acute apex. Mesocoxae (fig. 15) evident as weak ridge. Procoxal fissure pres- separated by mesostemal process and broad, ent and widely open (figs. 12, 46). Protro- rounded metastemal ridge. Mesendosternite chantin exposed. Prostemum (fig. 18) with (fig. 28) with expanded blunt apex; latero- long, stout setae anterior to procoxae; setig- posterior arm absent. Metendosternite (fig. erous pit absent. Prostemal intercoxal pro- 29) with long, narrow stalk and furcal arms; 1983 HERMAN: EPPELSHEIMIUS 9

FIGS. 12-20. Eppelsheimius pirazzol/i. 12. Prothorax, lateral view. 13. Submentum and gular region, male. 14. Head. 15. Pterothorax, ventral view.- 16. Submentum and gular region, female. 17. Pronotum. 18. Prothorax, ventral view, left coxa removed. 19. Sternum IX, male. 20. Protibia, apex. furcal arms sclerotized and anterolater-ally di- spinelike setae (figs. 20, 37). Mesotibiae with rected; anterior tendons close to one another one row of, and many scattered, elongate se- and arising from elongate median stalk. tae. Tarsal formula 5-5-5. Procoxae broad and strongly expanded. Abdomen with pair of elongate narrow Protibiae (fig. 6) strongly expanded; posterior sclerites (stemite I?) anterior to sternite II. surface with numerous spinelike setae and Sternite II well developed; midlongitudinal other stout setae; apex with pair of thick, carina absent. Terga without basolateral 10 AMERICAN MUSEUM NOVITATES NO. 2764

FIGS. 21-30. Eppelsheimius pirazzolii. 21. Labium. 22. Maxilla, palps, galea, and lacinia removed. 23. Labrum, epipharyngeal lobe removed from right side and setae from left. 24. Mandible, lateroapical view. 25. Mandible, dorsal view, left. 26. Antenna. 27. Galea, dorsal surface, apical setigerous brush removed. 28. Mesendostemite. 29. Metendosternite. 30. Head, ventral view. ridges. Tergum VII with fringe on posterior vided dorsally by tergum X, each half con- margin. Tergum VIII with posterior margin nected by narrow, sclerotized strap anterior broadly and feebly emarginate. Segments II to tergum X; strap fused to middle ofanterior to VII each with one or two pairs of latero- margin of tergum X; opening for abdominal sternites. Tergum X (figs. 31, 34) nearly di- glands with membranous dorsal surface. Ter- 1 983 HERMAN: EPPELSHEIMIUS I1I

FIGS. 31-34. Eppelsheimius pirazzolii. 31. Segments IX and X, dorsal view, female. 32. Aedeagus, dorsal view. 33. Segments IX and X, ventral view, female. 34. Segments IX and X, dorsal view, male. FIGS. 35-39. Eppelsheimius miricollis. 35. Galea, dorsal surface, apical setigerous brush removed. 36. Labrum, epipharyngeal lobe removed from left and setae removed from right. 37. Protibia, apex. 38. Maxilla, palps, galea, and lacinia removed. 39. Protibia, spinelike setae of posterior apex. gum IX of male (fig. 34) with long struts on female with struts on anterior margin ofven- anterior margin of ventromedial edges. Ster- tromedial edge. Coxites ovoid and flattened num IX of male flattened and of variable (fig. 33). Valvifers flattened and of irregular shape (figs. 19, 43). Tergum IX (fig. 31) of shape (fig. 33). Stylus absent (fig. 33). 12 AMERICAN MUSEUM NOVITATES NO. 2764

TABLE 2 Measurements (in Millimeters) for Eppelsheimius pirazzolii Pronotal width at Head width Head width Width Interocular anterior Pronotal Interocular Pronotal of head width margin length width width pirazzolii (All specimens, excluding 0.76a 0.04b 0.56 0.03 0.56 0.04 0.68 0.04 1.36 0.03 1.16 0.02 type of persicus) (N = 40) (0.65-0.84)c (0.48-0.63) (0.55-0.76) (0.57-0.77) (1.30-1.41) (1.10-1.21) (Type) 0.81 0.61 0.69 0.74 1.33 1.17 (Iranian specimen) 0.78 0.57 0.65 0.68 1.37 1.20 persicus (Type) 0.72 0.50 0.58 0.60 1.44 1.24 a Mean; b standard deviation; c range.

Aedeagus (fig. 32) trilobed. Parameres stout, ture. Abdomen dark reddish brown to pale elongate and unmodified. Median lobe broad reddish brown; segments VII to X usually basally and with acute apex. darker than anterior segments. Legs and an- Spermatheca (fig. 40) membranous; sper- tennae reddish brown. mathecal gland present. Dorsum ofhead with moderately strong to DISTRIBUTION: Eppelsheimius is known weak ground sculpturing and surface shining from Morocco eastward across northern Af- dully to polished; pubescence (fig. 14) mod- rica to Buchara (fig. 2). erately dense. Head (fig. 14) moderately Eppelsheimius pirazzolii (Eppelsheim) strongly convergent toward base. Clypeus less than 1.5 times wider than long (fig. 14); sur- Figures 1-34; Table 2 face sparsely pubescent. Eyes protruding Oncophorus pirazzolii Eppelsheim, 1885, p. 47. slightly from head (fig. 14). Submentum (figs. Fauvel, 1902, p. 73. (Type locality: Tunis, Tu- nisia.) 13, 16) with anteriorly directed, straight pro- Eppelsheimius pirazzolii (Eppelsheim): Bemhauer, cess on each side; males (fig. 13) with long 1915, p. 270. Koch, 1934, p. 35; 1936, pp. 119, processes, female (fig. 16) with short pro- 134; 1937, p. 229. Scheerpeltz, 1958, pp. 14- cesses. Maxilla with short anteriorly directed 16. process on dorsal surface of stipes (fig. 22); Eppelsheimius persicus Scheerpeltz, 1958, p. 14. galea with short row of five spiniform setae (Type locality: Iran, Sultanabad. Type in Natur- (fig. 27). Mandible (figs. 24, 25) tapered to historisches Museum, Wien.) NEW SYNONYM. apical point. First segment of antenna mod- DIAGNOSIS: Eppelsheimius pirazzolii can erately sinuate in lateral view; segments 9, be separated from miricollis by the pointed 10, 1 1 nearly cylindrical in cross section. mandibular apex (figs. 24, 25), long clypeus Pronotum with feeble ground sculpturing (fig. 14), presence of a pronotal lateral mar- and moderately shining or without ground ginal bead (fig. 12), long postprocoxal lobe sculpturing and polished; surface (fig. 17) with (fig. 12), and presence of two pairs of para- moderately dense pubescence. Pronotum (fig. tergites per abdominal segment. Numerous 17) with lateral margins gradually convergent other characters that differentiate the two from anterior angles to basal angles; lateral species are given in the description. margin nearly straight to subsinuate; prono- DESCRIPTION: Length 3.6 to 5.1 mm. tum widest near anterior margin. Pronotal Color pale to dark reddish brown with yel- lateral marginal bead present (fig. 12); post- lowish brown elytra. Head black to pale red- procoxal lobe present and moderately long dish brown. Prothorax dark reddish brown (fig. 12). Protergosternal suture evident as to pale reddish brown. Elytra yellowish brown feeble ridge. Prosternal intercoxal process with pale reddish brown infusions along su- short and extending between coxae (fig. 12). 1 983 HERMAN: EPPELSHEIMIUS 13

Protibia (fig. 20) with short apical process 2 and 3, pronotal, cephalic and elytral width, anterior to thick curved apical spinelike seta. pronotal length and cephalic punctation and Mesosternal intercoxal process (fig. 15) mod- sculpturing), other specimens have these erately long and tapered apically; apex acute. characters in a variety of combinations. Elytra with moderately dense short pubes- Scheerpeltz indicates that the eyes ofpersicus cence; surface without long setae. are larger and bulge (laterally) more than the Abdomen moderately densely pubescent. eyes ofpirazzolii. To evaluate this I measured Abdominal segments II to VII each with two the width of the head across the eyes and the pairs of paratergites. interocular width and made a ratio ofthe two, Male. Tergum IX (fig. 34) with slender an- dividing the head width by the interocular teriorly directed struts on dorsomedian edge; width (table 2). The higher the resulting num- dorsal surface with pair of moderately long ber, the more bulging the eyes. The head and slender, anteriorly directed struts on an- width/interocular width ofpersicus is 1.44. I terior margin. Sternum IX (fig. 19) broad, measured 40 specimens of pirazzolii and in- with pair of anteriorly directed lobes on an- cluded specimens from all localities. The terior margin. range for head width is 0.65 to 0.84 mm., for Female. Tergum IX without struts (fig. 31) interocular width 0.48 to 0.63 mm., and the on dorsomedian edge. Tergum X with mod- ratio is 1.30-1.41. The head-interocular ratio erately long and slender, anteriorly directed is 1.40 to 1.41 for five specimens from five strut on anterior margin. localities. The only other specimen known SEXUAL DIMORPHISM: The males have from Iran has a ratio of 1.37; the specimen longer submental processes (fig. 13) than the from Iraq has a ratio of 1.38. The type of females (fig. 16). persicus does in fact have more bulging eyes VARIATION: In spite of the description of but only barely. an Iranian specimen of pirazzolli as a new Another character that is easily analyzed species (Scheerpeltz, 1958), there is relatively is the width of the head compared with the little notable variation, certainly less than for width of the anterior pronotal margin. Di- miricollis. The characters used by Scheerpeltz viding the width of the head by the width of are found widely in the species and in differ- the pronotum gives a figure of 1.24 for per- ent combinations. Some of the variation of sicus and a range of 1. 10 to 1 .21 for 40 spec- size is shown in table 2. imens ofpirazzolii. Four specimens from four SYNONYMY: Eppelsheimius persicus, de- localities (including the second Iranian lo- scribed by Scheerpeltz (1958) from one spec- cality) have a ratio larger than 1.20; many imen, was the second report ofthe genus from others are near 1.20. The head of persicus is the region of the southern border of Iran and relatively wider and the pronotum relatively Iraq (Koch, 1937). Until Koch's citation the narrower than specimens of pirazzolii, but genus was known from Algeria, Tunisia, again only slightly. In absolute measure- Egypt, and French Somaliland (specimens ments, persicus falls near the upper part of from the last locality are E. miricollis). the range of variation seen for pirazzolii. Scheerpeltz, possibly impressed by the mag- The characters used by Scheerpeltz are nitude ofthe geographical disjunction, sought variable in pirazzolii and not greatly different characters that would distinguish the Iranian in persicus. Further collecting, particularly specimen from other species of the genus. from Iran and Iraq, probably will reveal suf- With only one Iranian specimen and perhaps ficient variation to bridge the gap more de- few ofpirazzolii (only six were seen from the finitively. I therefore regard persicus and pi- Scheerpeltz collection), it was inevitable that razzolii to be conspecific. little consideration would be given to varia- Parenthetically, the holotype ofpersicus is tion. All the characters that purport to dis- a female, not a male as indicated by Scheer- tinguish persicus and pirazzolii can be bridged peltz. Although I did not dissect the holotype, by specimens ofpirazzoli. Although the types the coxites ofthe genital segment can be seen of persicus and pirazzolii can be distin- without dissection. The submental processes guished by characters cited by Scheerpeltz are short, as they are in other females of pi- (relative size of the eyes, antennal segments razzolli. 14 AMERICAN MUSEUM NOVITATES NO. 2764

TABLE 3 Measurements (in Millimeters) for Eppelsheimius miricollis Head width Pronotal Pronotal width at Pronotal width at Width anterior width at Pronotal anterior of head margin widest length margin Large form 1.17a 0.04b 0.87 0.04 0.95 0.04 1.02 0.04 1.35 0.02 (N = 10; Tunisia) (I.I1-1.23)c (0.82-0.94) (0.90-1.01) (0.98-1.07) (1.29-1.38) Intermediate form 0.96 0.02 0.71 0.02 0.80 0.03 0.87 0.03 1.36 0.03 (N = 5; Iran) (0.93-0.98) (0.67-0.72) (0.75-0.84) (0.84-0.91) (1.33-1.42) Small form 0.81 0.06 0.62 0.04 0.68 0.03 1.77 0.04 1.32 0.04 (N = 4; throughout range) (0.71-0.91) (0.57-0.68) (0.63-0.72) (0.69-0.82) (1.26-1.41) a Mean; b standard deviation; c range. The numbers are in the same sequence throughout the table.

HABITAT AND DISTRIBUTION: The species MNHN). Iraq: Basra, April 1, 1936, Frey (1 is known from Morocco across north Africa MGF). Morocco: Tenouchan, November 3, to Egypt then again in southeastern Iraq and 1971, H. Coiffait (10 HCC). Saudi Arabia: southwestern and south-central Iran (fig. 2, Dammam [Ad Dammam], May 18,1976, W. see Material Examined). Buettiker (1 NMMB). Tunisia: Bel Assel: (1 None of the specimens have habitat data MNHN); Chott de Tozeur (1 MNHN); To- associated with them; a few specimens were zeur, G. C. Champion (1 FMNH; 1 MHMV; collected at light. All the localities from which 73 BMNH), May 1954, R. Demoflys (5 MTC; the species was collected were at or near sa- 5 AMNH); Kebili (3 IRSN), April 1887 (1 line habitats. MNHN), R. Demoflys (10 MTC; 5 AMNH); MATERIAL EXAMINED: 170 specimens. Gabes (1 IRSN); Matmata, May 1939, R. Africa: (Type of pirazzolii; 1 female Demoflys (1 HCC); Tunis (5 BMNH); El MHMV.) Algeria: Touggourt (1 MNHN), Hamma, May 1939, R. Demoflys (8 MTC; May 1898, L. Vareilles (1 BMNH), 7/2/29, 2 AMNH); Kriz, April 1949, R. Demoflys, A. Schatzmayr (1 BMNH; 2 MSNM), Le- (1 MTC); DJ [Djebel]; Tebaga, May 1939, R. vasseur (1 BMNH); Tidikelt, In Salah, central Demoflys (6 MTC); Zarzis, March 1950, R. Sahara, April 24-30, 1912, Hartert and Hilg Demoflys (2 MTC). (1 BMNH); Massif du Hoggar, Peyerimhoff (4 MHMV); Oasis Beni-Abbes, sur Orane- sado, J. Mateu (3 HCC); Beni Abbes, 4/5/65, Eppelsheimius miricollis (Fauvel) R. Gauthier, at light (1 MNHN), April 13, Figures 2, 35-49; Table 3 1963, J. Mateu, at light (1 MNHN), July 23, Oncophorus miricollis Fauvel, 1898, p. 96. Fair- 1969 (1 MNHN); Zerhamra, northwest Sa- maire, 1892, p. 79 (cited as pirazzolii). Fauvel, hara, 60 Km de [word illegible] Atlas, May 1902, p. 73. (Type locality: I have not examined 1961, F. Pierre (2 MNHN); M'Guebra, April the type. In the original description two locali- 1887 (1 MNHN); El Golea (2 MNHN). Egypt: ties, "Province de Constantine" and "Tunisie," Mariut [Maryut], April 26, 1932, Priesner (1 are cited.) MSNM; specimen missing head and pro- Eppelsheimius miricollis (Fauvel): Koch, 1934, p. thorax). Iran: Sultanabad [=Soltanabad], 35; 1936, pp. 119, 134. Scheerpeltz, 1958, pp. Bodemeyer (Type ofpersicus; MHMV); Bal- 15-16. out Chab lut [=Baluch Ab, about 100 km. DIAGNOSIS: Eppelsheimius miricollis can NE Bam],2 March 9, 1969, F. Pierre (1 be separated from pirazzolii by the truncate, flattened mandibular apex (figs. 47,48), short clypeus (fig. 41), absence of a pronotal lateral 2 The location of Baluch Ab was provided by Ms. N. marginal bead (fig. 46), short postprocoxal Berti who contacted F. Pierre, the collector. lobe (fig. 46), and presence of one pair of 1 983 HERMAN: EPPELSHEIMIUS 15

FIGS. 40-49. Eppelsheimius miricollis. 40. Spermatheca. 41. Head. 42. Submentum and gular region, large form. 43. Sternum IX, male. 44. Prothorax. 45. Submentum and gular region, small form. 46. Prothorax, lateral view. 47. Mandible, dorsal view, left. 48. Mandible, lateroapical view. 49. Segments IX and X, dorsal view, male. paratergites per abdominal segment. Many Color pale to dark reddish brown with yel- other characters given in the description dif- lowish brown elytra. Head and prothorax pale ferentiate the species. reddish brown, head paler than prothorax. DESCRIPTION: Length 3.1 to 6.8 mm. Elytra yellowish brown with pale reddish 16 AMERICAN MUSEUM NOVITATES NO. 2764 brown infusions along suture. Abdomen red- VARIATION: The size and several structures dish yellow to reddish brown to dark reddish are notably variable. Table 3 describes some brown. Legs and antennae pale reddish brown. of the variation of size. Dorsum of head with feeble ground sculp- The species varies from 3.1 to 6.8 mm. turing and surface strongly shining to pol- long. The head width is 0.71 to 1.23 mm., ished on central portion; lateral region ad- and the pronotal length 0.69 to 1.07 mm. The jacent to eye with strong ground sculpturing largest individuals (5.5 to 6.8 mm. long) are and dully shining; pubescence sparse (fig. 41). from Zarzis and Kebili (Tunisia), interme- Head strongly convergent toward base (fig. diate ones are from Iran. The smallest indi- 41). Clypeus more than 2 times wider than viduals (3.1 to 4.9 mm. long) are found at long (fig. 41); surface sparsely pubescent. Eyes many localities including Zarzis, Tunisia. At (fig. 41) protruding moderately strongly from Zarzis four specimens, two of the large form head. Submentum (figs. 42, 45) with ante- and two of the small, were collected in Oc- riorly directed, sinuate (fig. 42) to straight (fig. tober 1946; attached to them is the same 45) process on each side. Maxilla with long, handwritten label: "B-3380." All these spec- anteriorly directed process on dorsal surface imens were presumably collected together. of stipes (fig. 38); galea with long row of 10 Other than size and the form of the submen- spiniform setae (fig. 35). Mandible (figs. 47, tal processes, nothing separates them. 48) tapered to blunt, emarginate apex. First The form of the submental process varies segment ofantenna strongly sinuate in lateral from strongly sinuous (fig. 42) to straight (fig. view; segments 9, 10, 1 1 compressed in cross 45). The large form has the sinuous processes, section. the small ones the straight ones. However, Pronotum with feeble ground sculpturing, some of the small and intermediate speci- surface polished; surface sparsely pubescent mens have weakly sinuous processes. (fig. 44). Pronotum with lateral margins sin- The mesosternal intercoxal process has an uous and widest just behind middle (fig. 44). acute apex in most specimens of the large Pronotal lateral marginal bead absent (fig. 46); form, one has a blunt apex. The intercoxal postprocoxal lobe present and small (fig. 46). process of the small form has a blunt apex, Protergosternal suture evident as feeble ridge. a few have an acute apex. Prosternal intercoxal process long and ex- Specimens from most localities have a dark tending between coxae (fig. 46). Protibia with reddish brown abdomen; specimens from long apical process (fig. 37) anterior to thick Obock, French Somaliland and Al Qunfud- curved apical spinelike seta (fig. 39). Mesos- hah, Saudi Arabia have a reddish yellow ab- ternal intercoxal process short and tapered domen. apically; apex acute to rounded. Elytra with HABITAT AND DISTRIBUTION: Eppelsheim- sparse short pubescence; surface with a few, ius miricollis is known from 10 localities scat- scattered long setae. tered from Algeria to Iran, the Soviet Union, Abdomen sparsely pubescent. Abdominal and French Somaliland (fig. 2; see Material segments II to VII each with one pair ofpara- Examined). A few specimens were collected tergites. at light. Eppelsheimius miricollis and E. pi- Male. Tergum IX (fig. 49) with broad an- razzolii probably live in similar saline habi- teriorly directed struts on dorsomedian edge; tats; both species were collected at three lo- dorsal surface with pair of short, broad, an- calities in Tunisia and one in Iran. teriorly directed struts on anterior margin; DISCUSSION: The variation is sufficiently sternum IX (fig. 43) broad posteriorly, ta- great as to suggest the possibility that E. mi- pered anteriorly, then expanded near anterior ricollis is two species. At Zarzis and Kebili, margin; anterior margin with pair of antero- Tunisia, seven specimens ofa large form with laterally directed lobes. sinuous submental processes were collected Female. Tergum IX without strut on dor- (see Variation). Two other specimens, much somedian edge. Tergum X with short, broad, smaller and with straight submental process- anteriorly directed strut on anterior margin. es were collected from Zarzis with two of the SEXUAL DIMORPHISM: None. larger form. At all of the other localities the 1983 HERMAN: EPPELSHEIMIUS 17 specimens are small or intermediate between with notes on certain allied forms. Ent. the two forms. The occurrence of a few spec- Mon. Mag., vol. 55, pp. 154-156. imens that seem to bridge the gaps between Eppelsheim, Eduard the large and small forms supports the hy- 1885. Eine neue Oxytelinen-Gattung der pothesis that E. miricollis is one variable Meditteran-Fauna. Deutsche Ent. Zeitschr., vol. 29, pp. 46-48. species, not two. Further collecting with an Fairmaire, Leon. emphasis on long series will permit testing of 1892. Coleopteres d'Obock. Troisieme partie. this idea. As ofthis writing only 45 specimens Rev. d'Ent. vol. 11, pp. 77-127. have been examined. Fauvel, Albert A specimen from Repetek, Buchara, al- 1898. Catalogue des Staphylinides de Barba- though lacking a head and pronotum, can be rie, et des Iles Acores, Maderes, Sal- identified by the presence of only one pair of vages et Canaries. Rev. d'Ent., vol. 17, paratergites per abdominal segment which is pp. 93-113. characteristic of E. miricollis. 1902. Catalogue des staphylinides de la Bar- MATERIAL EXAMINED: 45 specimens. barie de la Basse-Egypte et des iles Acores, Maderes, Salvages et Canaries. Algeria: Chott Melrhir, May 1891 (6 (ed. 5). Rev. d'Ent. 21, pp. 45-189. MHMV; 1 FMNH; small form). French So- Hammond, Peter M. maliland: Obock, G. Hardy (6 MNHN; small 1975. Reports of the Lund University Expe- form). Iran: Balout Chab lut [=Baluch Ab, dition in 1962. Report no. 34 Coleop- about 100 km. NE Bam],2 March 9, 1969, F. tera: Staphylinidae: Oxytelini from Pierre (5 MNHN; intermediate form). Saudi Ceylon. Ent. Scand., suppl. 4, pp. 141- Arabia: Hofuf, April 19, 1977, W. Buettiker 178. (1 BMNH; 3 NHMB; small form); near El 1976. A review of the genus Anotylus C. G. Gumfuda [=Al Qunfudhah], March 1936, R. Thomson (Coleoptera: Staphylinidae). C. M. Darling, at light (1 BMNH; small form). Bull. Brit. Mus. (Nat. Hist.). Ent. vol. 33, no. 2, pp. 137-187,3 plates, 18 figs. Sudan: Mersa Halaib, January 1, 1933, Pries- Herman, Lee H., Jr. ner (1 MSNM). Tunisia: Kebili, May 1952, 1970. Phylogeny and reclassification of the R. Demoflys (4 MTC; 3 AMNH; large form); genera ofthe rove-beetle subfamily Ox- Tozeur, May 1954, R. Demoflys (1 MTC; ytelinae of the world (Coleoptera, small form); Zarzis, October 1946 (4 MTC; Staphylinidae). Bull. Amer. Mus. Nat. large and small forms), December 1950 (1 Hist., vol. 142, art. 5, pp. 343-454, figs. MTC; large form), R. Demoflys. U.S.S.R.: 1-73, table 1. Buchara: Repetek, April 1900, F. Hauser (1 1972. Revision of Bledius and related genera. FMNH; small form; head and prothorax Part 1. The aequatorialis, mandibularis, missing). and semiferrugineus groups and two new In to I genera (Coleoptera, Staphylinidae, Ox- addition the preceding specimens, ytelinae). Bull. Amer. Mus. Nat. Hist. have studied one with the following data: Gr. vol. 149, art. 2, pp. 111-254, figs. 1- Balachan, Dschebell, 1898, F. Hauser 451, maps 1-17, tables 1-5. (FMNH). Dschebell is most probably Djebel 1976. Revision of Bledius and related genera. for Mountain, the remainder is obscure. Two Part II. The armatus, basalis, and me- other specimens are labeled: Dschebell lanocephalus groups (Coleoptera, (MHMV). Staphylinidae, Oxytelinae). Bull. Amer. Mus. Nat. Hist. vol. 157, art. 2, pp. 71- 172, figs. 1-391, tables 1-2. LITERATURE CITED [In press] Revision of Bledius, Part III. The an- nularis and emarginatus groups, with Bemhauer, Max data for species of previous revised 1915. Beitrag zur Kenntnis der Paliiarktischen groups. Bull. Amer. Mus. Nat. Hist. Staphyliniden-Fauna IV. Miinchener Koch, Carl Kol. Zeitschr., vol. 4, pp. 262-270. 1934. Wissenschaftliche Ergebnisse der En- Champion, George C. tomologischen Expeditionen seiner 1919. On some new species of the staphylinid Durchlaucht des Fuirsten Alessandro C. genus Planeustomus Duv. from India, della Torre e Tasso nach Aegypten und 18 AMERICAN MUSEUM NOVITATES NO. 2764

auf die Halbinsel Sinae. IV. Staphyli- Newton, Alfred nidae: Coleoptera. Bull. Soc. Roy. Ent. 1982. A new genus and species of Oxytelinae Egypte. vol. 18, pp. 33-91. from Australia, with a description of its 1936. Wissenschaftliche Ergebnisse der En- larva, systematic position, and phylo- tomologischen Expeditionen seiner genetic relationships (Coleoptera, Durchlaucht des Fiirsten Alessandro C. Staphylinidae). Amer. Mus. Novitates, della Torre e Tasso nach Aegypten und no. 2744, pp. 1-24, figs. 1-41, table 1. auf die Halbinsel Sinae. XIII. Staphy- Scheerpeltz, Otto linidae. Pubbl. Mus. Ent. Pietro Rossi, 1958. Eine neue Art der Gattung Eppelsheim- vol. 1, pp. 115-232. ius Bemh. (Col., Staphylinidae) nebst einer Bestimmungstabelle der bis heute 1937. Uber einige Staphylinidae aus dem Ost- lichen Mediterriingebiet. Pubbl. Mus. bekannt gawordenen Arten dieser Gat- tung. Kol. Rundsch., vol. 35 (1957), pp. Ent. Pietro Rossi, vol. 2, pp. 229-264. 14-17.