Eppelsheimius : revision, distribution, sister group relationship (Staphylinidae, Oxytelinae)

No AMERICAN MUSEUM

vitates

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY

CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y.
Number 2764, pp. 1-18, figs. 1-49, tables 1-3

10024
June 23, 1983

Eppelsheimius: Revision, Distribution,
Sister Group Relationship
(Staphylinidae, Oxytelinae)

LEE H. HERMAN!

ABSTRACT

Eppelsheimius is a small genus of beetles that
occurs in arid regions from northern Africa to
southwestern Asia. The species share characters
with Planeustomus, Manda, and Bledius. Evi-
dence is presented that Bledius and Eppelsheimius
are sister groups. The genus has two species, E.

pirazzolii and E. miricollis, that are distinguished
by many characters. Both species are variable. The
genus and species are described and illustrated and
their distributions described. One species, E. per-
sicus, is newly synonymized with E. pirazzolii.

INTRODUCTION

The present paper was stimulated by a
search for the sister group of Bledius. Earlier,
but without supporting characters, Herman
(1970, p. 354) presented two groups of genera
as the sister group of Bledius. One of these
groups, the Carpelimus lineage, includes
Carpelimus, Apocellagria, Trogactus, Thi-
nodromus, Xerophygus, Ochthephilus,
Mimopaederus, Teropalpus, Pareiobledius
and Blediotrogus;, the other, the Thinobius
lineage, includes Thinobius, Sciotrogus, and
Neoxus. My own subsequent studies and those
of others (Hammond, 1975, 1976; Newton,
1982) have pointed to a number of inade-
quacies in my phylogeny of the Oxytelinae
(Herman, 1970, p. 354). One problem, the
relationship of Bledius to Epplesheimius, is
addressed herein; some others are discussed

in a forthcoming paper on Bledius (Herman,
in prep.). Ultimately, several rearrangements
in the classification of the Oxytelinae will be
required.

Eppelsheim (1885) described pirazzolii in
Oncophorus. A second species, miricollis, was
added by Fauvel (1898); both were from Tu-
nisia. In 1915, Oncophorus was discovered
to be a homonym of a genus of Mal-
lophaga and a genus of ““worms”’ of indeter-
minate placement. Bernhauer (1915) pub-
lished a replacement name, Eppelsheimius,
and a few years later Champion (1919), un-
aware of Bernhauer’s change, proposed On-
cogenys to supersede Oncophorus. Koch in a
series of papers (1934, 1936, 1937) estab-
lished the occurrence of the genus in many
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

NO. 2764

AMERICAN MUSEUM NOVITATES

lti.

imius pirazzo

Eppelshe

Fic. 1.

. The genus was redescribed

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sheimius was reported in southwestern Iran
when Scheerpeltz (1958) described a third

1983

HERMAN: EPPELSHEIMIUS 3

TABLE 1
Polarity of Some Character States of the Oxytelinae

Character Plesiomorphic

Labral setae Short, slender

Hypopharynx Apical lobe entire
Mandibular mola Well developed
Antennae Not geniculate
Gular sutures Separated
Neck Present, broad
Clypeus Large, at same level as dorsum
Eyes Restricted to side of head
Protibiae Slender
Dorsal seta of epipharyngeal
lobe Absent
Elytral suture Straight

When Eppelsheim described the genus he
placed it near Manda (cited as Acrognathus)
and Planeustomus (cited as Compsochilus)
because of unspecified similarities. Herman
(1970, p. 355) found support for this hy-
pothesis in the presence in the three genera
of elongate, membranous epipharyngeal lobes
(fig. 23).

Although I said (Herman, 1970, p. 355)
that the species of Eppelsheimius share the
enlarged, membranous, epipharyngeal lobes
with Manda and Planeustomus, dissection of
the species of Eppelsheimius reveals that only
E. pirazzolii has enlarged lobes (figs. 10, 11,
23); miricollis has shorter but well-developed
lobes (fig. 36). However, the epipharyngeal
lobes of the species of the three genera do
have apically bifurcate or multifurcate cutic-
ular processes and the labrum of each has
long stout labral setae (figs. 23, 36). Both of
these features seem to be derived but I find
no others that the three genera share.

Manda and Planeustomus share a narrow
clypeus that is depressed below the level of
the dorsum, eyes that extend onto the ventral
surface of the head, reduction to near absence
of the mandibular mola, and the bifid apical
lobes of the hypopharynx. The first two of
these four states are unique in the family.
Manda and Planeustomus are a well-defined
and supported group.

Eppelsheimius and Bledius share six de-
rived features: confluent gular sutures (fig. 30),
absence of a neck (fig. 1), expanded protibiae
(fig. 6), a dorsal seta on the epipharyngeal lobe
(figs. 23, 36), dehiscent elytra (fig. 1), and

Apomorphic

Long, stout (figs. 23, 36)

Apical lobe bifid

Reduced, nearly absent

Geniculate (fig. 26)

Confluent (fig. 30)

Absent (figs. 1, 14, 41)

Narrow, depressed below level of dorsum
Extend onto ventral surface of head
Expanded (fig. 6)

Present (figs. 23, 36)
Dehiscent (fig. 1)

geniculate antenna (fig. 26; first segment elon-
gate and the second segment posteriorly flex-
ile on first). Some of these characters are found
in other oxyteline genera but not in the com-
bination found for Bledius and Eppelsheim-
ius. The polarity of the character states given
in table 1 was derived by their relative dis-
tribution in other subfamilies, particularly the
Omaliinae, Piestinae, and Osoriinae. The
apical lobe of the hypopharynx is entire, the
mandibular mola well developed, the clypeus
large and at the same level as the dorsum,
the eyes restricted to the side of the head, the
antennae normal (not geniculate), the gular
sutures separated, the neck broad and weakly
developed, the protibiae slender, the dorsal
seta of the epipharyngeal lobe absent, and the
elytral suture straight in most of the genera
of the Osoriinae, Piestinae, and Omaliinae.
Variation from these conditions in the Oxy-
telinae are derived (table 1).

No other genera of the Oxytelinae have the
expanded protibiae. The configuration of the
confluence of the gular sutures as found in
Bledius and Eppelsheimius (fig. 30) is also
found in the genera of the Carpelimus and
Thinobius lineages (see Herman, 1970, p.
354). Dehiscent elytra are found in Thinobius
and in some species of Platystethus. Species
of Coprophilus and one of Manda lack a neck.
Geniculate antennae are found only in Ble-
dius and Eppelsheimius. The dorsal epipha-
ryngeal seta is found in one species of Manda.

The hypothesis that Eppelsheimius is the
sister taxon of Bledius rather than of Manda
and Planeustomus is supported by the fact

that it shares six apomorphic features with
Bledius but only two with the two other gen-
era. Except as noted in the preceding para-
graph, Eppelsheimius shares no other appar-
ently derived features with other genera that
are not shared by most genera of the subfam-
ily. This hypothesis along with the relation-
ship of Eppelsheimius and Bledius to other
oxyteline genera is discussed further in a pa-
per dealing with the phylogeny of Bledius
(Herman, in prep.). The present paper is a
first step toward understanding the relation-
ships of the species of Bledius and of the ge-
nus to other genera of the subfamily. Many
questions remain. What is the sister group to
Bledius and Eppelsheimius? How are other
genera, such as Aploderus, Pareiobledius,
Blediotrogus, and Xerophygus, that share
some derived characters with Bledius and
Eppelsheimius related to them? What are the
relationships of the genera of the Carpelimus
and Thinobius lineages to Bledius and Ep-
pelsheimius and to other genera of the
subfamily? These and similar problems are
beyond the scope of the present paper.

Based on similarities to Bledius, that is the
expanded protibiae, the absence of a neck,
and the subcylindrical body, I expect that
species of Eppelsheimius make burrows in
the soil. Eppelsheimius, as does Bledius, may
use their elongate mandibles to excavate bur-
rows. The absence ofa neck permits the great-
er range of movement needed to pick up and
deposit sand. The presence of a neck restricts
the degree of rotation of the head. The ex-
panded protibiae are not used by Bledius to
construct a burrow, and it is unlikely that
Eppelsheimius uses them in that way. Species
of both genera may use them to facilitate agile
movement in the burrow.

If Eppelsheimius and Bledius are sister taxa,
several interesting comparisons can be made.
Bledius has 420 species found in nearly all
parts of the world. They burrow into the soil
adjacent to fresh and saline water of rivers,
lakes, ponds, and the ocean; some live in tem-
porarily wet soil that is separated from stand-
ing water. They feed on algae. Bledius is high-
ly variable anatomically; the degree of
variation permits recognition of many species
groups based on numerous characters. These
groups often have more characters separating

AMERICAN MUSEUM NOVITATES

NO. 2764

them than do some genera of the Oxytelinae.
The species within each group may be ex-
tremely difficult to distinguish (Herman,
1972, 1976, in press, and in prep.).

By contrast, although Eppelsheimius might

-share the saline habitat with Bledius, it has

a more restricted geographical and ecological
distribution and is a relatively minuscule ge-
nus. However, the two species of Eppel-
sheimius are separated by as many characters
as are some species groups of Bledius. This
divergence in Eppelsheimius coupled with the
intraspecific variation leads me to wonder
whether the two species actually represent
complexes of subtly distinguishable species.
I was unable to detect discrete variation with-
in the two recognized species with samples
available to me.

Information on the habitat of species of
Eppelsheimius has not been published or in-
cluded on the specimen labels. Since all the
localities from which the species have been
taken are near salt water—either near the sea
or near inland salt lakes, flats, and marshes
it is probable that they live in saline habitats.
Although the species have been collected only
near salt marshes of northern Africa, along
the coasts of the Mediterranean and Red seas,
and at a few probably saline habitats in Saudi
Arabia, Iraq, Iran, and the USSR (fig. 2), they
are likely to occur near the salt lakes and salt
marshes that are found across arid parts of
Asia to China. The species are attracted to
light.

In addition to querying the extent of the
geographical range, further collection can de-
termine where the species live, whether they
make burrows, and if so how, what they eat,
where the larvae live, and what their anatom-
ical features are. The collection sites of Ep-
pelsheimius are widely scattered (fig. 2) but
near the Chott Djerid, a large salt marsh in
Tunisia, 141 of the 217 specimens studied
were collected. Chott Djerid would be an ap-
propriate place to try to answer the preceding
questions and perhaps others.

ACKNOWLEDGMENTS AND
ABBREVIATIONS

Specimens used in this study were bor-
rowed from the individuals and institutions

HERMAN: EPPELSHEIMIUS

1983

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6 AMERICAN MUSEUM NOVITATES

listed below. Abbreviations preceding the
name of each institution are used in the text
to indicate the location of specimens. The
name of the person who lent the material
follows the name of the institution. I grate-
fully thank them for their assistance. I es-
pecially thank Dr. Heinrich Sch6nmann, who
lent types, Miss Beatrice Brewster, who trans-
lated articles from French and German, and
Miss Joan Whelan, who took the scanning
electron micrographs.

AMNH, American Museum of Natural History.

BMNH, British Museum (Natural History), Lon-
don; Mr. Peter Hammond.

FMNH, Field Museum of Natural History, Chi-
cago; Dr. Larry Watrous.

HCC, Dr. H. Coiffait, Toulouse.

IRSN, Institut Royal des Sciences Naturelles,
Brussels; Dr. Léon Baert.

MGF, Museum G. Frey, via. Dr. Gerhard Scherer.

MHMV, Naturhistorisches Museum Wien, Vi-
enna; Dr. Heinrich Schénmann.

MNHN, Muséum National d’Histoire Naturelle,
Paris; Miss Nicole Berti.

MSNM, Museo Civico di Storia Naturale, Milano;
Dr. Carlo Leonardi.

MTC, Marc Tronquet, Paris.

NHMEB, Naturhistorisches Museum Basel, Basel;
Dr. M. Brancucci.

EPPELSHEIMIUS BERNHAUER
Figures 1-49

Eppelsheimius Bernhauer, 1915, p. 270. Scheer-
peltz, 1958, pp. 14-17. Herman, 1970, p. 369.
Type Species: Eppelsheimius pirazzolii (Ep-
pelsheim).

Oncophorus Eppelsheim, 1885, p. 46; preoccu-
pied. Type Species: Oncophorus pirazzolii Ep-
pelsheim.

Oncogenys Champion, 1919, p. 154. TYPE SPECIEs:
Oncogenys pirazzolii (Eppelsheim).

DIAGNOSIS: Eppelsheimius is separated
from other genera of the Oxytelinae by the
presence of an epistomal suture (fig. 14), ge-
niculate antennae (fig. 26), open procoxal fis-
sure (figs. 12, 46), spiniform submental pro-
cesses (figs. 13, 16, 42, 45), confluent gular
sutures (fig. 30), dehiscent elytra suture (fig.
1), absence of elytral epipleural ridges, pres-
ence of five tarsomeres, and expanded, spi-
nous protibiae (fig. 6).

The genus is similar to Bledius in form and
shares with it many characters. Except for the

NO. 2764

presence of five tarsomeres and the submen-
tal processes the diagnostic features cited in
the preceding paragraph are found in all or
some of the species of Bledius. The species
of Eppelsheimius are more “loosely con-
structed”’ than are those of Bledius. The me-
sotibia of Bledius has two rows of spinelike
setae; that of Eppelsheimius has one row of
elongate setae among the scattered ones. The
postprocoxal lobe is larger in Bledius than in
Eppelsheimius. The genital appendages of the
females of Bledius consist of a pair of elon-
gate, flattened sclerites; those of Eppelsheim-
ius are paired, elongate, transversely divided
(into a coxite and a valvifer) and compressed
(fig. 33).

DESCRIPTION: Length 3.1 to 6.8 mm.

Color pale to dark reddish brown with yel-
lowish brown elytra. Form slender and sub-
cylindrical. Body sparsely pubescent.

Head (figs. 14, 41) sparsely pubescent and
punctate; lateral margins gradually conver-
gent from eyes toward base; neck absent,
postocular transverse groove absent. Clypeal
length variable. Epistomal suture straight; su-
ture approximately even with anterior mar-
gin of supra-antennal ridge. Eyes (figs. 14, 41)
slightly to moderately protruding from sides
of head, not extending onto ventral surface.
Supra-antennal ridge small. Anterior and
dorsal tentorial arms present. Antenna (fig.
26) with first segment elongate and genicu-
late; segments 9, 10, and | 1 expanded to form
loose, well-defined club. Gular sutures (fig.
30) confluent for most of length, sutures
sharply divergent at base. Submentum (figs.
13, 16, 42, 45) with short to long, slender to
stout spiniform process on anterior lateral
edge. Labium (fig. 21) with trapezoidal men-
tum; palpal segments subequal. Hypophar-
ynx as in figures 3 and 4. Labrum (figs. 23,
36) fused, without midlongitudinal fissure;
surface polished and with long, stout setae
near anterior margin; posterior margin with
long, posteriorly directed, internal strut on
lateral edge; ventroposterior median surface
with triangular internal strut. Epipharynx
(figs. 23, 36) with short to long membranous,
anteriorly directed lobe; lobe with long, slen-
der, apically bifurcate process on mesial mar-
gin; epipharyngeal lobe with long, stout setae
near base of dorsal surface. Maxilla as in fig-

1983 HERMAN: EPPELSHEIMIUS 7

me”

* > , a " mye ’
ox SOA * mn 7 Ptr
ws “4 > ® Af - =
H fe Fe asi ee Gi
(Bet te. ok WGA &

Fics. 3-7. Eppelsheimius pirazzolii. 3. Hypopharynx, 270X. 4. Hypopharynx, 674. 5. Scutellum,
220X. 6. Protibia and tarsus, 105X. 7. Metatibia and tarsus, 121.

ure 8; fourth segment of maxillary palpus stout 22, 38) with short to long spine on anterior
(fig. 9); galea (figs. 27, 35) with row of stout margin of dorsal surface. Mandibles (figs. 25,
spinelike setae on dorsal surface; stipes (figs. 47) edentate, long, curved mesially, and

8 AMERICAN MUSEUM NOVITATES

NO. 2764

Fics. 8-11.
collapsed, 479X. 10. Epipharynx, ventral surface, 156X. 11. Epipharynx, ventral surface, 491.

touching at apices when closed, but not cross-
ing.

Pronotum (figs. 17, 44) longer than wide;
lateral margins gradually curved to nearly
straight or strongly sinuous with bulge near
basal third; surface sparsely pubescent and
punctate with shallow midlongitudinal
groove. Pronotal lateral marginal bead pres-
ent (fig. 12) or absent (fig. 46). Prohypomeron
impressed anterior to coxae; postprocoxal
portion present as small (fig. 46) to large (fig.
12) lobe. Protergosternal (figs. 12, 46) suture
evident as weak ridge. Procoxal fissure pres-
ent and widely open (figs. 12, 46). Protro-
chantin exposed. Prosternum (fig. 18) with
long, stout setae anterior to procoxae; setig-
erous pit absent. Prosternal intercoxal pro-

Eppelsheimius pirazzolii. 8. Maxilla, 206.9. Maxillary palpus, fourth segment, partially

cess (figs. 12, 46) elongate, carinate and ven-
troposteriorly directed; ventral edge straight.
Scutellum with apex slightly exposed to con-
cealed under pronotum; surface with elongate
oval impression (fig. 5). Elytra elongate; su-
ture dehiscent (fig. 1); posterior margin with-
out membranous lobe; epipleural ridge ab-
sent; surface sparsely punctate and pubescent
and without longitudinal striae. Mesosternal
(fig. 15) process short to moderately long and
extending between coxae; process broad and
tapered to acute apex. Mesocoxae (fig. 15)
separated by mesosternal process and broad,
rounded metasternal ridge. Mesendosternite
(fig. 28) with expanded blunt apex; latero-
posterior arm absent. Metendosternite (fig.
29) with long, narrow stalk and furcal arms;

1983

HERMAN: EPPELSHEIMIUS 9

Fics. 12-20. Eppelsheimius pirazzolii. 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 anterolaterally di-
rected; anterior tendons close to one another
and arising from elongate median stalk.
Procoxae broad and strongly expanded.
Protibiae (fig. 6) strongly expanded; posterior
surface with numerous spinelike setae and
other stout setae; apex with pair of thick,

spinelike setae (figs. 20, 37). Mesotibiae with
one row of, and many scattered, elongate se-
tae. Tarsal formula 5-5-5.

Abdomen with pair of elongate narrow
sclerites (sternite I?) anterior to sternite II.
Sternite II well developed; midlongitudinal
carina absent. Terga without basolateral

10 AMERICAN MUSEUM NOVITATES NO. 2764

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Fics. 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. Mesendosternite. 29. Metendosternite. 30. Head, ventral view.

ridges. Tergum VII with fringe on posterior
margin. Tergum VIII with posterior margin
broadly and feebly emarginate. Segments. II
to VII each with one or two pairs of latero-
sternites. Tergum X (figs. 31, 34) nearly di-

vided ‘dorsally by tergum X, each half con-

nected by narrow, sclerotized strap anterior
to tergum X; strap fused to middle of anterior
margin of tergum X; opening for abdominal
glands with membranous dorsal surface. Ter-

1983 HERMAN: EPPELSHEIMIUS 11

Fics. 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.

Fics. 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
anterior margin of ventromedial edges. Ster-
num IX of male flattened and of variable
shape (figs. 19, 43). Tergum IX (fig. 31) of

female with struts on anterior margin of ven-
tromedial edge. Coxites ovoid and flattened
(fig. 33). Valvifers flattened and of irregular
shape (fig. 33). Stylus absent (fig. 33).

12 AMERICAN MUSEUM NOVITATES

TABLE 2
Measurements (in Millimeters) for Eppelsheimius pirazzolii

Width Interocular
of head width
pirazzolii
(All specimens, excluding 0.767 0.04% 0.56 0.03

type of persicus) (N = 40)

(Type) 0.81 0.61

(Iranian specimen) 0.78 0.57
persicus

(Type) 0.72 0.50

«Mean; ” standard deviation; © range.

Aedeagus (fig. 32) trilobed. Parameres stout,
elongate and unmodified. Median lobe broad
basally and with acute apex.

Spermatheca (fig. 40) membranous; sper-
mathecal gland present.

DISTRIBUTION: Eppelsheimius is known
from Morocco eastward across northern Af-
rica to Buchara (fig. 2).

Eppelsheimius pirazzolii (Eppelsheim)
Figures 1-34; Table 2

Oncophorus pirazzolii Eppelsheim, 1885, p. 47.
Fauvel, 1902, p. 73. (Type locality: Tunis, Tu-
nisia.)

Eppelsheimius pirazzolii (Eppelsheim): Bernhauer,
1915, p. 270. Koch, 1934, p. 35; 1936, pp. 119,
134; 1937, p. 229. Scheerpeltz, 1958, pp. 14—-
16.

Eppelshéimius persicus Scheerpeltz, 1958, p. 14.
(Type locality: Iran, Sultanabad. Type in Natur-
historisches Museum, Wien.) NEW SYNONYM.

DiAGnosis: Eppelsheimius pirazzolii can
be separated from miricollis by the pointed
mandibular apex (figs. 24, 25), long clypeus
(fig. 14), presence of a pronotal lateral mar-
ginal bead (fig. 12), long postprocoxal lobe
(fig. 12), and presence of two pairs of para-
tergites per abdominal segment. Numerous
other characters that differentiate the two
species are given in the description.

DESCRIPTION: Length 3.6 to 5.1 mm.

Color pale to dark reddish brown with yel-
lowish brown elytra. Head black to pale red-
dish brown. Prothorax dark reddish brown
to pale reddish brown. Elytra yellowish brown
with pale reddish brown infusions along su-

(0.65-0.84)* (0.48-0.63) (0.55-0.76)

NO. 2764
Pronotal
width at Head width Head width
anterior Pronotal Interocular Pronotal
margin length width width
0.56 0.04 0.68 0.04 1.36 0.03 1.16 0.02
(0.57-0.77) (1.30-1.41) (1.10-1.21)
0.69 0.74 1.33 1.17
0.65 0.68 1.37 1.20
0.58 0.60 1.44 1.24

ture. Abdomen dark reddish brown to pale
reddish brown; segments VII to X usually
darker than anterior segments. Legs and an-
tennae reddish brown.

Dorsum of head with moderately strong to
weak ground sculpturing and surface shining
dully to polished; pubescence (fig. 14) mod-
erately dense. Head (fig. 14) moderately
strongly convergent toward base. Clypeus less
than 1.5 times wider than long (fig. 14); sur-
face sparsely pubescent. Eyes protruding
slightly from head (fig. 14). Submentum (figs.
13, 16) with anteriorly directed, straight pro-
cess on each side; males (fig. 13) with long
processes, female (fig. 16) with short pro-
cesses. Maxilla with short anteriorly directed
process on dorsal surface of stipes (fig. 22);
galea with short row of five spiniform setae
(fig. 27). Mandible (figs. 24, 25) tapered to
apical point. First segment of antenna mod-
erately sinuate in lateral view; segments 9,
10, 11 nearly cylindrical in cross section.

Pronotum with feeble ground sculpturing
and moderately shining or without ground
sculpturing and polished; surface (fig. 17) with
moderately dense pubescence. Pronotum (fig.
17) with lateral margins gradually convergent
from anterior angles to basal angles; lateral
margin nearly straight to subsinuate; prono-
tum widest near anterior margin. Pronotal
lateral marginal bead present (fig. 12); post-
procoxal lobe present and moderately long
(fig. 12). Protergosternal suture evident as
feeble ridge. Prosternal intercoxal process
short and extending between coxae (fig. 12).

1983

Protibia (fig. 20) with short apical process
anterior to thick curved apical spinelike seta.
Mesosternal intercoxal process (fig. 15) mod-
erately long and tapered apically; apex acute.
Elytra with moderately dense short pubes-
cence; surface without long setae.

Abdomen moderately densely pubescent.
Abdominal segments II to VII each with two
pairs of paratergites. "

Male. Tergum IX (fig. 34) with slender an-
teriorly directed struts on dorsomedian edge;
dorsal surface with pair of moderately long
and slender, anteriorly directed struts on an-
terior margin. Sternum IX (fig. 19) broad,
with pair of anteriorly directed lobes on an-
terior margin.

Female. Tergum IX without struts (fig. 31)
on dorsomedian edge. Tergum X with mod-
erately long and slender, anteriorly directed
strut on anterior margin.

SEXUAL DIMORPHISM: The males have
longer submental processes (fig. 13) than the
females (fig. 16).

VARIATION: In spite of the description of
an Iranian specimen of pirazzolii as a new
species (Scheerpeltz, 1958), there is relatively
little notable variation, certainly less than for
miricollis. The characters used by Scheerpeltz
are found widely in the species and in differ-
ent combinations. Some of the variation of
size 1s shown in table 2.

SyNonyMyY: Eppelsheimius persicus, de-
scribed by Scheerpeltz (1958) from one spec-
imen, was the second report of the genus from
the region of the southern border of Iran and
Iraq (Koch, 1937). Until Koch’s citation the
genus was known from Algeria, Tunisia,
Egypt, and French Somaliland (specimens
from the last locality are E. miricollis).
Scheerpeltz, possibly impressed by the mag-
nitude of the geographical disjunction, sought
characters that would distinguish the Iranian
specimen from other species of the genus.
With only one Iranian specimen and perhaps
few of pirazzolii (only six were seen from the
Scheerpeltz collection), it was inevitable that
little consideration would be given to varia-
tion. All the characters that purport to dis-
tinguish persicus and pirazzolii can be bridged
by specimens of pirazzolii. Although the types
of persicus and pirazzolii can be distin-
guished by characters cited by Scheerpeltz
(relative size of the eyes, antennal segments

HERMAN: EPPELSHEIMIUS 13

2 and 3, pronotal, cephalic and elytral width,
pronotal length and cephalic punctation and
sculpturing), other specimens have these
characters in a variety of combinations.
Scheerpeltz indicates that the eyes of persicus
are larger and bulge (laterally) more than the
eyes of pirazzolii. To evaluate this I measured
the width of the head across the eyes and the
interocular width and made aratio of the two,
dividing the head width by the interocular
width (table 2). The higher the resulting num-
ber, the more bulging the eyes. The head
width/interocular width of persicus is 1.44. I
measured 40 specimens of pirazzolii and in-
cluded specimens from all localities. The
range for head width is 0.65 to 0.84 mm., for
interocular width 0.48 to 0.63 mm., and the
ratio is 1.30—1.41. The head-interocular ratio
is 1.40 to 1.41 for five specimens from five
localities. The only other specimen known
from Iran has a ratio of 1.37; the specimen
from Iraq has a ratio of 1.38. The type of
persicus does in fact have more bulging eyes
but only barely.

Another character that is easily analyzed
is the width of the head compared with the
width of the anterior pronotal margin. Di-
viding the width of the head by the width of
the pronotum gives a figure of 1.24 for per-
sicus and a range of 1.10 to 1.21 for 40 spec-
imens of pirazzolii. Four specimens from four
localities G@ncluding the second Iranian lo-
cality) have a ratio larger than 1.20; many
others are near 1.20. The head of persicus is
relatively wider and the pronotum relatively
narrower than specimens of pirazzolii, but
again only slightly. In absolute measure-
ments, persicus falls near the upper part of
the range of variation seen for pirazzolii.

The characters used by Scheerpeltz are
variable in pirazzolii and not greatly different
in persicus. Further collecting, particularly
from Iran and Iraq, probably will reveal suf-
ficient variation to bridge the gap more de-
finitively. I therefore regard persicus and pi-
razzolii to be conspecific.

Parenthetically, the holotype of persicus is
a female, not a male as indicated by Scheer-
peltz. Although I did not dissect the holotype,
the coxites of the genital segment can be seen
without dissection. The submental processes
are short, as they are in other females of pi-
razzolii.

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 1.177 0.04 0.87 0.04 0.95 0.04 1.02 0.04 1.35 0.02
(N = 10; Tunisia) (1.11-1.23)° (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)

* Mean; ® standard deviation; ‘ range. The numbers are in the same sequence throughout the table.

HABITAT AND DISTRIBUTION: The species
is known from Morocco across north Africa
to Egypt then again in southeastern Iraq and
southwestern and south-central Iran (fig. 2,
see Material Examined).

None of the specimens have habitat data
associated with them; a few specimens were
collected at light. All the localities from which
the species was collected were at or near sa-
line habitats.

MATERIAL EXAMINED: 170 specimens.

Africa: (Type of pirazzolii; 1 female
MHMV.) Algeria: Touggourt (1 MNHN),
May 1898, L. Vareilles (1 BMNH), 7/2/29,
A. Schatzmayr (1 BMNH; 2 MSNM), Le-
vasseur (1 BMNH); Tidikelt, In Salah, central
Sahara, April 24-30, 1912, Hartert and Hilg
(1 BMNH); Massif du Hoggar, Peyerimhoff
(4 MHMV); Oasis Beni-Abbes, sur Orane-
sado, J. Mateu (3 HCC); Beni Abbes, 4/5/65,
R. Gauthier, at light (1 MNHN), April 13,
1963, J. Mateu, at light (1 MNHN), July 23,
1969 (1 MNHN); Zerhamra, northwest Sa-
hara, 60 Km de [word illegible] Atlas, May
1961, F. Pierre (2 MNHN); M’Guebra, April
1887 (1 MNHN),; El Golea (2 MNHN). Egypt:
Mariut [Maryut], April 26, 1932, Priesner (1
MSNM; specimen missing head and pro-
thorax). Iran: Sultanabad [=Soltanabad],
Bodemeyer (Type of persicus; MHMV); Bal-
out Chab lut [=Baluch Ab, about 100 km.
NE Bam],? March 9, 1969, F. Pierre (1

2 The location of Baluch Ab was provided by Ms. N.
Berti who contacted F. Pierre, the collector.

MNHN). Iraq: Basra, April 1, 1936, Frey (1
MGF). Morocco: Tenouchan, November 3,
1971, H. Coiffait (10 HCC). Saudi Arabia:
Dammam [Ad Dammam], May 18, 1976, W.
Buettiker (1 NMMB). Tunisia: Bel Assel: (1
MNHN); Chott de Tozeur (1 MNHN); To-
zeur, G. C. Champion (1 FMNH; 1 MHMV;
73 BMNH), May 1954, R. Demoflys (5 MTC;
5 AMNH); Kebili (3 IRSN), April 1887 (1
MNHN), R. Demoflys (10 MTC; 5 AMNH);
Gabes (1 IRSN); Matmata, May 1939, R.
Demoflys (1 HCC); Tunis (5 BMNH); El
Hamma, May 1939, R. Demoflys (8 MTC;
2 AMNH); Kriz, April 1949, R. Demoflys,
(1 MTC); DJ [Djebel]; Tebaga, May 1939, R.
Demoflys (6 MTC); Zarzis, March 1950, R.
Demoflys (2 MTC).

Eppelsheimius miricollis (Fauvel)
Figures 2, 35-49; Table 3

Oncophorus miricollis Fauvel, 1898, p. 96. Fair-
maire, 1892, p. 79 (cited as pirazzolii). Fauvel,
1902, p. 73. (Type locality: I have not examined
the type. In the original description two locali-
ties, ‘“Province de Constantine” and “‘Tunisie,”’
are cited.)

Eppelsheimius miricollis (Fauvel): Koch, 1934, p.
35; 1936, pp. 119, 134. Scheerpeltz, 1958, pp.
15-16.

DiaGcnosis: Eppelsheimius miricollis can
be separated from pirazzolii by the truncate,
flattened mandibular apex (figs. 47, 48), short
clypeus (fig. 41), absence of a pronotal lateral
marginal bead (fig. 46), short postprocoxal
lobe (fig. 46), and presence of one pair of

1983 HERMAN: EPPELSHEIMIUS 15

Fics. 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

brown infusions along suture. Abdomen red-
dish yellow to reddish brown to dark reddish
brown. Legs and antennae pale reddish brown.

Dorsum of head with feeble ground sculp-
turing and surface strongly shining to pol-
ished on central portion; lateral region ad-
jacent to eye with strong ground sculpturing
and dully shining; pubescence sparse (fig. 41).
Head strongly convergent toward base (fig.
41). Clypeus more than 2 times wider than
long (fig. 41); surface sparsely pubescent. Eyes
(fig. 41) protruding moderately strongly from
head. Submentum (figs. 42, 45) with ante-
riorly directed, sinuate (fig. 42) to straight (fig.
45) process on each side. Maxilla with long,
anteriorly directed process on dorsal surface
of stipes (fig. 38); galea with long row of 10
spiniform setae (fig. 35). Mandible (figs. 47,
48) tapered to blunt, emarginate apex. First
segment of antenna strongly sinuate in lateral
view; segments 9, 10, 11 compressed in cross
section.

Pronotum with feeble ground sculpturing,
surface polished; surface sparsely pubescent
(fig. 44). Pronotum with lateral margins sin-
uous and widest just behind middle (fig. 44).
Pronotal lateral marginal bead absent (fig. 46);
postprocoxal lobe present and small (fig. 46).
Protergosternal suture evident as feeble ridge.
Prosternal intercoxal process long and ex-
tending between coxae (fig. 46). Protibia with
long apical process (fig. 37) anterior to thick
curved apical spinelike seta (fig. 39). Mesos-
ternal intercoxal process short and tapered
apically; apex acute to rounded. Elytra with
sparse short pubescence; surface with a few,
scattered long setae.

Abdomen sparsely pubescent. Abdominal
segments II to VII each with one pair of para-
tergites.

Male. Tergum IX (fig. 49) with broad an-
teriorly directed struts on dorsomedian edge;
dorsal surface with pair of short, broad, an-
teriorly directed struts on anterior margin;
sternum IX (fig. 43) broad posteriorly, ta-
pered anteriorly, then expanded near anterior
margin; anterior margin with pair of antero-
laterally directed lobes.

Female. Tergum IX without strut on dor-
somedian edge. Tergum X with short, broad,
anteriorly directed strut on anterior margin.

SEXUAL DIMORPHISM: None.

NO. 2764

VARIATION: The size and several structures
are notably variable. Table 3 describes some
of the variation of size.

The species varies from 3.1 to 6.8 mm.
long. The head width is 0.71 to 1.23 mm.,
and the pronotal length 0.69 to 1.07 mm. The
largest individuals (5.5 to 6.8 mm. long) are
from Zarzis and Kebili (Tunisia), interme-
diate ones are from Iran. The smallest indi-
viduals (3.1 to 4.9 mm. long) are found at
many localities including Zarzis, Tunisia. At
Zarzis four specimens, two of the large form
and two of the small, were collected in Oc-
tober 1946; attached to them is the same
handwritten label: ‘““B-3380.” All these spec-
imens were presumably collected together.
Other than size and the form of the submen-
tal processes, nothing separates them.

The form of the submental process varies
from strongly sinuous (fig. 42) to straight (fig.
45). The large form has the sinuous processes,
the small ones the straight ones. However,
some of the small and intermediate speci-
mens have weakly sinuous processes.

The mesosternal intercoxal process has an
acute apex in most specimens of the large
form, one has a blunt apex. The intercoxal
process of the small form has a blunt apex,
a few have an acute apex.

Specimens from most localities have a dark
reddish brown abdomen; specimens from
Obock, French Somaliland and Al Qunfud-
hah, Saudi Arabia have a reddish yellow ab-
domen.

HABITAT AND DISTRIBUTION: Eppelsheim-
ius miricollis is known from 10 localities scat-
tered from Algeria to Iran, the Soviet Union,
and French Somaliland (fig. 2; see Material
Examined). A few specimens were collected
at light. Eppelsheimius miricollis and E. pi-
razzolii probably live in similar saline habi-
tats; both species were collected at three lo-
calities in Tunisia and one in Iran.

DISCUSSION: The variation is sufficiently
great as to suggest the possibility that E. mi-
ricollis is two species. At Zarzis and Kebili,
Tunisia, seven specimens ofa large form with
sinuous submental processes were collected
(see Variation). Two other specimens, much
smaller and with straight submental process-
es were collected from Zarzis with two of the
larger form. At all of the other localities the

1983

specimens are small or intermediate between
the two forms. The occurrence of a few spec-
imens that seem to bridge the gaps between
the large and small forms supports the hy-
pothesis that FE. miricollis is one variable
species, not two. Further collecting with an
emphasis on long series will permit testing of
this idea. As of this writing only 45 specimens
have been examined.

A specimen from Repetek, Buchara, al-
though lacking a head and pronotum, can be
identified by the presence of only one pair of
paratergites per abdominal segment which is
characteristic of E. miricollis.

MATERIAL EXAMINED: 45 specimens.

Algeria: Chott Melrhir, May 1891 (6
MHMV; 1 FMNH; small form). French So-
maliland: Obock, G. Hardy (6 MNHN; small
form). Iran: Balout Chab lut [=Baluch Ab,
about 100 km. NE Bam],” March 9, 1969, F.
Pierre (5 MNHN;; intermediate form). Saudi
Arabia: Hofuf, April 19, 1977, W. Buettiker
(1 BMNH; 3 NHMB; small form); near El
Gumfuda [=A] Qunfudhah], March 1936, R.
C. M. Darling, at light (1 BMNH; small form).
Sudan: Mersa Halaib, January 1, 1933, Pries-
ner (1 MSNM). Tunisia: Kebili, May 1952,
R. Demoflys (4 MTC; 3 AMNH; large form);
Tozeur, May 1954, R. Demoflys (1 MTC;
small form); Zarzis, October 1946 (4 MTC;
large and small forms), December 1950 (1
MTC; large form), R. Demoflys. U.S.S.R.:
Buchara: Repetek, April 1900, F. Hauser (1
FMNH; small form; head and prothorax
missing).

In addition to the preceding specimens, I
have studied one with the following data: Gr.
Balachan, Dschebell, 1898, F. Hauser
(FMNH). Dschebell is most probably Djebel
for Mountain, the remainder is obscure. Two
other specimens are labeled: Dschebell
(MHMV).

LITERATURE CITED

Bernhauer, Max
1915. Beitrag zur Kenntnis der Palaarktischen
Staphyliniden-Fauna IV. Miinchener
Kol. Zeitschr., vol. 4, pp. 262-270.
Champion, George C.
1919. On some new species of the staphylinid
genus Planeustomus Duv. from India,

HERMAN: EPPELSHEIMIUS 17

with notes on certain allied forms. Ent.
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Eppelsheim, Eduard
1885. Eine neue Oxytelinen-Gattung der
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1892. Coléoptéres d’Obock. Troisiéme partie.
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Fauvel, Albert

1898. Catalogue des Staphylinides de Barba-
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vages et Canaries. Rev. d’Ent., vol. 17,

pp. 93-113.
1902. Catalogue des staphylinides de la Bar-

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Hammond, Peter M.

1975. Reports of the Lund University Expe-
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A review of the genus Anotylus C. G.

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Bull. Brit. Mus. (Nat. Hist.). Ent. vol.

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Herman, Lee H., Jr.

1970. Phylogeny and reclassification of the
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1-73, table 1.

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genera (Coleoptera, Staphylinidae, Ox-
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vol. 149, art. 2, pp. 111-254, figs. 1-

451, maps 1-17, tables 1-5.

Revision of B/edius and related genera.

Part II. The armatus, basalis, and me-

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Staphylinidae, Oxytelinae). Bull. Amer.

Mus. Nat. Hist. vol. 157, art. 2, pp. 71-

172, figs. 1-391, tables 1-2.

[In press] Revision of Bledius, Part III]. The an-
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groups. Bull. Amer. Mus. Nat. Hist.

Koch, Carl

1934. Wissenschaftliche Ergebnisse der En-

tomologischen Expeditionen seiner
Durchlaucht des Fiirsten Alessandro C.
della Torre e Tasso nach Aegypten und

1976.

1972.

1976.

18

1936.

1937.

AMERICAN MUSEUM NOVITATES

auf die Halbinsel Sinae. IV. Staphyli-
nidae: Coleoptera. Bull. Soc. Roy. Ent.
Egypte. vol. 18, pp. 33-91.

Wissenschaftliche Ergebnisse der En-
tomologischen Expeditionen seiner
Durchlaucht des Fiirsten Alessandro C.
della Torre e Tasso nach Aegypten und
auf die Halbinsel Sinae. XIII. Staphy-
linidae. Pubbl. Mus. Ent. Pietro Rossi,
vol. 1, pp. 115-232.

Uber einige Staphylinidae aus dem Ost-
lichen Mediterrangebiet. Pubbl. Mus.
Ent. Pietro Rossi, vol. 2, pp. 229-264.

NO. 2764

Newton, Alfred

1982.

A new genus and species of Oxytelinae
from Australia, with a description of its
larva, systematic position, and phylo-
genetic relationships (Coleoptera,
Staphylinidae). Amer. Mus. Novitates,
no. 2744, pp. 1-24, figs. 1-41, table 1.

Scheerpeltz, Otto

1958.

Eine neue Art der Gattung Eppelsheim-
ius Bernh. (Col., Staphylinidae) nebst
einer Bestimmungstabelle der bis heute
bekannt gawordenen Arten dieser Gat-
tung. Kol. Rundsch., vol. 35 (1957), pp.
14-17.