ZooKeys 24: 3 1-54 (2009) A peer-rev iewed open-access journal

j 1]
doi: 10.3897/zookeys.24.247 RESEARCH ARTICLE #Zookey S

www.pens oftonline.n et/zoo keys Launched to accelerate biodiversity research

Seven new species of Cephennium Miller &
Kunze (Coleoptera, Staphylinidae, Scydmaeninae,
Cephenniini) from California with a key to native

North American species

Katie J. Hopp*, Michael S. Caterino*

Santa Barbara Museum of Natural History, 2559 Puesta del Sol Rd., Santa Barbara, CA 93105-2998, U.S.A

t urn:lsid:zoobank. org:author:803 BA3DB-84C3-4BF6-96 1 4-64D4E2FB1855
£ urn:lsid:zoobank.org:author:F687B1E2-A07D-4F28-B1F5-4A0DD17B6490

Corresponding author: Katie J. Hopp (katichopp@gmail.com), Michael S. Caterino (mcaterino@sbnature2.org)

Academic editor: Jan Klimaszewski | Received 14 August 2009 | Accepted 17 September 2009 | Published 9 October 2009
urn.lsid:zoobank. org:pub:BB883B 1E-E58F-4074-92EB-7E8 14E78F678

Citation: Hopp KJ, Caterino MS (2009) Seven new species of Cephennium Miller & Kunze (Coleoptera, Staphylini-
dae, Scydmaeninae, Cephenniini) from California with a key to native North American species. ZooKeys 24: 31-54.
doi: 10.3897/zookeys.24.247

Abstract

Seven new species of Cephennium from California are described and illustrated - C. celsifrons, sp. n., C.
mariposae, sp. n., C. grandarboreum, sp. n., C. canestroi, sp. n., C. gilberti, sp. n., C. urbanum, sp. n. and
C. aridum, sp. n. The single known native Nearctic species, C. anophthalmicum Brendel, was known only
from moist coastal forests around the San Francisco Bay area. The new species greatly expand the distribu-
tion of the genus, through central and southern California, occurring in the central Sierra Nevada, south
through the coast ranges and Sierra Nevada to the Santa Monica Mountains and desert foothills of the
San Bernardino and San Jacinto Mountains. A key to all eight species (the entire native Nearctic fauna) of

Cephennium occurring in California is provided.

Keywords

Systematics, Staphylinidae, Scydmaeninae, Cephenniini, Cephennium, California Floristic Province

Introduction

Members of the tribe Cephenniini (Coleoptera: Staphylinidae: Scydmaeninae) are pri-
marily known to occur in the western Palearctic region, and most of the known spe-

Copyright Katie J. Hopp, Michael S. Caterino. This is an open access article distributed under the terms of the Creative Commons Attribution
License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

32 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

cies belong to the genus Cephennium. Newton and Franz (1998) report 124 species
of Cephennium, only one of which is known to occur in the Nearctic (O’Keefe 2001).
This species, Cephennium anophthalmicum, was described by Brendel in 1889 from
Alameda County in coastal, central California, and until this time remains the only
species of this genus described from the Nearctic region (O’Keefe 2001). A second spe-
cies of Cephennium, C. gallicum Ganglbauer, has been reported from maritime north-
eastern North America, but this represents a recent introduction from Europe (Majka
and Klimaszewski 2004).

The California Floristic Province is a biodiversity hotspot, recognized for its unique,
diverse, and threatened biota (Myers et al. 2000). The California Beetle Project, begun
in 2005, was launched to compile a complete inventory of the region’s Coleoptera. ‘This
effort has documented over 8000 species, and uncovered numerous new ones, both un-
detected and undescribed (Caterino 2006; Caterino and Chatzimanolis 2007; Caterino
et al. 2008). Through these efforts the inadequacy of current documentation of the re-
gion’s leaf litter fauna has become ever clearer. Intensive collecting in this microhabitat
by members of the CBP has yielded several new species of Cephennium since 2004,
three of which were discovered in the past two years. Along with the newly collected
material, four species were discovered in the collections of the California State Collec-
tion of Arthropods (CSCA) and the Field Museum of Natural History (FMNH), whose
collectors, Fred Andrews and Art Gilbert (by association, CSCA), and Al Newton and
Margaret Thayer (FMNH) are avid litter sifters. Only one of the species from our newly
collected material and the material we borrowed from other collections overlap, leaving
us with seven new species. These seven new species represent a considerable increase in
the known Nearctic fauna of this tribe and genus, and significantly expand its known

geographic range.

Materials and methods

Specimens examined for this study are deposited in the following institutions and col-
lections (all collection codens follow Evenhuis (2008) and the curators responsible for
borrowed specimens are listed in parentheses):

CASC California Academy of Sciences, San Francisco, CA (David Kavanaugh, Jere
Schweikert).

CSCA California State Collection of Arthropods, Sacramento, CA (Chuck Bellamy).

FMNH Field Museum of Natural History, Chicago, IL (Margaret Thayer, Alfred New-
ton).

LACM Los Angeles County Museum of Natural History, Los Angeles, CA (Brian
Brown).

MCZC Museum of Comparative Zoology, Harvard University, Cambridge, MA
(Philip Perkins).

SBMN Santa Barbara Museum of Natural History, Santa Barbara, CA.

Seven new species of Cephennium Miiller & Kunze 33

Label data from the material examined are verbatim and transcribed following Ivie
(1985): the end of each line on a label is indicated by a “;” (semicolon); the individual
labels are separated by a “/” (backslash).

Specimens were studied with a Leica” MZ9.5 stereomicroscope equipped with a
150w Nikon MKII fiber optic light. SEMs were taken with a Zeiss” EVO 40 XVP
Scanning Electron Microscope. Specimen measurements were taken using an eyepiece
micrometer in a Leica® MZ9.5 stereomicroscope at 3.2x magnification. Length was
measured medially from the base of the pronotum to the apex of the elytra; pronotal
and eytral widths were measured at their widest points.

Male genitalia were extracted by first relaxing the specimen in hot water and then
the entire specimen was placed in a warm solution of 10% KOH. Once the specimen
was cleared, it was removed from the KOH solution and rinsed with distilled water,
and then placed in 100% EtOH for dissection. The specimen was placed on its dorsal
surface and then the abdomen was “pumped” by compressing the abdomen repeatedly
just basad of the medial lobe. This gentle pumping pushed the genitalia out through
the apical abdominal opening. After examination, the genitalia were then placed in a
vial of glycerin that was placed on the pin under the specimen labels. Males can some-
times be recognized when the outline of the genital capsule can be observed through
the translucent cuticle. Otherwise, except in the case of C. celsifrons, where there is a
distinct sexual dimorphism, the sexes are not usually distinguishable.

Key to Nearctic genera of Scydmaeninae

The most recent key to Nearctic genera of Scydmaenidae (=Scydmaeninae, Grebennik-
ov and Newton 2009) (O’ Keefe 2001) must be modified in order to accommodate the
new species of Cephennium described herein. We also revise this to reflect Jatoszynski’s
(2007) recent synonymization of Chelonoidum Strand with Cephennodes Reitter. All
other native Nearctic Cephenniini are now assigned to Cephennodes. Couplet 4(3)
should exclude the presence/absence of eyes because several of our new species possess
reduced eyes. ‘The couplet should read:

4(3) Foveae present in basal pronotal angles (Fig. 27.20); procoxae separated by

prosternal process (Fig. 10.20)... ceseseeeeseeseeeee Cephennodes (Fig. 27.20)
- Foveae absent from basal pronotal angles; procoxae not separated by proster-
HalepeGceses( Fic, “DY pscectvlocedselissesgnesneeoest aes Cephennium (Fig. 28.20)

Morphological characters of significance

There are a few diagnostic characters among the California Cephennium species that
may aid in identifying species collected in the future, and in determining their status

34 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

as described or new. Among the species we examined, characters that were useful in de-
limiting species included the presence of eyes and number of ommatidia, the shape of
the humeral angle of the elytron, the number of scutellar setae, the vestiture and apical
shape of the mesosternal keel, and the male genitalia. Within the aedeagus, structures
that were morphologically useful to delimit species were the shape of the median dorsal
projection and the arrangement of setae arising from the apical collar.

Species descriptions of California Cephennium Miller & Kunze

Cephennium anophthalmicum Brendel, 1889
Bigs LAA, BAL oA, ‘6

Type Material. Not seen. A holotype was not designated for this species. However,
it was described from a single specimen from Alameda County that was sifted from
vegetable debris together with a large number of Pinodytes cryptophagoides (currently
Catopocerus cryptophagoides) by Marie Fuchs (Brendel 1889). We attempted to track
down this specimen but were not able to locate it at either the Academy of Natural Sci-
ences in Philadelphia, the original repository of the Brendel collection, or the Museum
of Comparative Zoology at Harvard University, the current repository for the Brendel
collection. However, we did see a specimen from the MCZC that was determined as C.
anophthalmicum from Alameda County. Because the label data do not exactly match
the information presented by Brendel (1889) we believe this is only a topotype, and
not a primary type. We choose not to designate a neotype here as it is possible that the
original type specimen is still in existence somewhere.

Material Examined. “Alameda; Co. CAL.”/ “Laundry; Farm’/ “H. C. FALL;
COLLECTION”/ “Cephennium; anophthalmicum; Brend.” (1 MCZC); “Mill Valley;
Marin Co. Cal.; 30.V.1952”/ “By sifting; forest duff’/ “H.B. Leech; Collector”/ red
square label/ “Cephennium; sp. 2; Cl. Besuchet; det. V 1961”/ “Collection of the;
CALIFORNIA ACADEMY; of SCIENCES, San; Francisco, Calif.” (1 CASC); “Loma
Mar; SanMateo Co.Calif; [V-29-1970"/ “ex. Redwood; Litter”/ “Collector; T.R.Haig”
(1 FMNH [female, disarticulated]).

Diagnosis. This species can be distinguished from its California congeners
by the character combination of the absence of eyes, humeral angle of elytron
bluntly angulate (Fig. 1A), and the absence of a basal elytral sutural ridge. Ce-
phennium anophthalmicum most closely resembles C. urbanum, but can be sepa-
rated from it by the presence of a basal elytral sutural ridge (Fig. 1G), and the
apex of the mesosternal keel divergent and crescent-shaped in C. urbanum (Fig.
2G). It can be easily distinguished from C. aridum, C. celsifrons, and C. mariposae
by the presence of eyes in these species (Fig. 4A, C-D), and can be separated from
C. grandarboreum, C. canestroi and C. gilberti by the humeral angle of the elytron,
which is raised, dorsally flattened and apically rounded in these three species (Fig.
3D-F).

Seven new species of Cephennium Miiller & Kunze

Figure |. A-H Dorsal habitus SEMs, all to same scale A Cephennium anophthalmicum Brendel B Ce-
phennium celsifrons Hopp & Caterino C Cephennium mariposae Hopp & Caterino D Cephennium grandar-
boreum Hopp & Caterino E Cephennium canestroi Hopp & Caterino F Cephennium gilberti Hopp &
Caterino G Cephennium urbanum Hopp & Caterino H Cephennium aridum Hopp & Caterino.

36 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

Figure 2. A-H Mesosternal keel SEMs A Cephennium anophthalmicum Brendel B Cephennium cel-
sifrons Hopp & Caterino C Cephennium mariposae Hopp & Caterino D Cephennium grandarboreum

Hopp & Caterino E Cephennium canestroi Hopp & Caterino F Cephennium gilberti Hopp & Caterino G
Cephennium urbanum Hopp & Caterino H Cephennium aridum Hopp & Caterino.

Redescription. Male. Length: 0.874 mm; pronotal width: 0.418 mm; elytral
width: 0.475 mm. Body elongate, slender, weakly convex; testaceous; evenly and mod-
erately pubescent; pubescence golden, slender, moderately long, weakly decumbent
(Fig. LA). Dorsal surface of head smooth, weakly pubescent, narrowing anteriorly from
antennal insertions. Eyes absent. Antenna setose, antennomere | and II longer than
broad, antennomeres [II-VI quadrate and smaller than antennomeres II and VII, an-

Seven new species of Cephennium Miiller & Kunze 37

a
DY}
a

Figure 3. A-H Base of left elytron A Cephennium anophthalmicum Brendel B Cephennium celsifrons
Hopp & Caterino C Cephennium mariposae Hopp & Caterino D Cephennium grandarboreum Hopp &
Caterino E Cephennium canestroi Hopp & Caterino F Cephennium gilberti Hopp & Caterino G Cephen-
nium urbanum Hopp & Caterino H Cephennium aridum Hopp & Caterino.

tennomere VIII smaller than antennomeres VII and IX, antennomeres [X-XI gradu-
ally clavate forming a loose club. Pronotum moderately pubescent, broadest between
middle and anterior third, very convex in disc and moderately flattened near each
posterior angle; anterior margin not visible from above; anterior and posterior margin
lacking marginal bead; lateral marginal bead complete, gradually widening towards
base; lateral edge broadly rounded to posterior third, then weakly sinuate to base (Fig.
1A). Hypomeron smooth, sparsely setose towards anterior quarter and along outside
(lateral) edge. Prosternum lacking protuberant nodules anterolaterad procoxal cavities
(Fig. 2A). Elytra smooth, as pubescent as pronotum, covering all abdominal segments;
elytral suture flat; basomedial fovea present on each elytron; fovea moderate in size,
moderately pubescent (Figs. 1A, 3A). Humeral angle of elytron projecting laterally
to blunt point, dorsally raised and flattened (Fig. 3A). Scutellum weakly triangular,
lacking setae (Fig. 3A). Mesosternal keel sparsely setose, lacking scale-like microsculp-

38 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

ture, posterior quarter impunctate, apex weakly bifid (divergent), divergent projec-
tions short, triangulate (Fig. 2A). Metathoracic wings vestigial. Femora strongly clavate
in distal half, tibiae expanded and becoming more densely setose towards distal half.
Five visible abdominal sternites, ventrites V and VI partially fused. Aedeagus strongly
sclerotized, with median lobe basally rounded, pill-shaped; parameres thin, sinuate,
bisetose apically, extending to apex of rather narrow, bluntly triangular median dorsal
process; apical digiform process curving ventrad at apex, extending just beyond apical
collar; membranous apical collar with sclerotized clasper-like processes extending from
apex; membranous lateral flaps present at base of apical collar (Fig. 5A).

Female. Identical to male.

Biology. This species was first described from a single specimen that was sifted
from vegetable debris. An additional specimen was sifted from forest duff. Beyond this,
there is little known about the biology of this species.

Distribution. This species has been collected around the San Francisco Bay Area
in central coastal California (Fig. 6).

Cepehennium celsifrons Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act: FB4E5DDF-D621-4B8E-A233-E2956D8C66EB
Figs 1B, 2B, 3B, 4A-B, 5B, 6

Type Material. Holotype. Male. “CALIF: Calaveras Co.; 3.0 mi NW West Point
[~38.4160°N, 120.5515°W]; 2250 ft., v.20.1976; berl.; litter, mixed hdwd.- Pinus -;
Libocedrus — Abies; for.; A. Newton, M. Thayer”/ “Cephennium sp. 2; A. Newton det.
1978” (FMNH).

Paratypes (8): 1 specimen with same data as holotype (SBMN [gold coated for
SEM]); “CALIF: Calaveras; Co., 3mi NW West; Point 2250’; v.20.1976”/ “A. New-
ton; M. Thayer; collectors”/ “A. Newton; M. Thayer; collectors” (1 MCZC); “CAL.:
Calaveras Co.; 3 mi NE Glencoe 2000’; VI.25.1975 berl. litter,; oak — conifer forest
away; from stream; A. Newton”/ “Cephennium; A. Newton det. 1975” (2 FMNH, fe-
males); “3 miW Michigan Bar; Amador Co., Cal.; I]-3-1971”/ “ex Berlese-; oak duff”/
“Fred G.; Andrews; collector” (1 FMNH, male [disarticulated]); “CALIF: Amador
Co.; Pine Grove; IV-15-1972; Fred G Andrews”/ “Berlesed; Oak; Litter” (1 CSCA));
“CALIF: Amador Co.; 1 mi. W Pine Grove; H-14-1971; R. E Wilkey; Ex. Rotten
wood” (1 CSCA); “Pine Grove; Amador Co., Cal.; [V.15.1972”/ “Berlesed; from; Oak
duff’ (1 CSCA [disarticulated]).

Etymology. This species name is derived from the latin celsus (elevated or lofty)
in combination with frons, in reference to the unique, prominent sexual dimorphism
exhibited by the male.

Diagnosis. This species can be distinguished from all California congeners by the
character combination of the presence of two ommatidia on each side of the head (Fig.
4A), the mesosternal process truncate at the apex (Fig. 2B), and the frons of the male
with a median longitudinal ridge (Fig. 4B). The male frons structure is unique to this

Seven new species of Cephennium Miiller & Kunze ao

Ommatidia

Ommatidia

Figure 4. A-—D Characters on the head, including number of ommatidia and male frons A-B Cephen-
nium celsifrons Hopp & Caterino C Cephennium mariposae Hopp & Caterino D Cephennium aridum
Hopp & Caterino.

species. Cephennium celsifrons and C. mariposae are otherwise very similar but can be
distinguished by the number of ommatidia present (2 vs. 4, respectively, Figs. 4A, C).
Cephennium aridum has a single ommatidium on each side of the head (Fig. 4D), and
no other species exhibit any traces of eyes.

Description. Male. Length: 0.817—0.893 mm; pronotal width: 0.323—0.380 mm;
elytral width: 0.380—0.418 mm. Body broad, ovate, slightly convex, rufo-testaceous
to amber yellow, evenly densely pubescent, pubescence golden, slender, long, moder-
ately decumbent (Fig. 1B). Head small, deflexed, sparsely pubescent, not narrowing
anteriorly from antennal insertions; frons pinched medially forming a medial ridge
(Fig. 4B); two ommatidia present on each side of the head (Fig. 4A). Antenna setose,
antennomere I and II longer than broad, antennomeres HI-VI quadrate and smaller
than antennomeres II and VII, antennomere VIII smaller than antennomeres VII and
IX, antennomeres IX-XI gradually clavate forming loose club. Pronotum densely pu-
bescent, broadest between middle and anterior third, disc very convex medially and

40 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

|
Figure 5.A-F Dorsal and lateral view of the aedeagus A Cephennium anophthalmicum Brendel B Ce-

phennium celsifrons Hopp & Caterino C Cephennium grandarboreum Hopp & Caterino D Cephennium
canestroi Hopp & Caterino E Cephennium urbanum Hopp & Caterino F Cephennium aridum Hopp &

Caterino.

weakly flattened near each posterior angle; anterior margin not visible from above;
anterior and posterior margin lacking marginal bead; marginal bead complete later-
ally, gradually widening towards base; lateral edge broadly rounded to posterior third,

Seven new species of Cephennium Miiller & Kunze 4]

then evenly curved to base (Fig. 1B). Hypomeron smooth, sparsely setose towards up-
per quarter and along outside (lateral) edge, hypomeral bead anterolaterad procoxae
sinuate. Prosternum without nodules anterolaterad procoxal cavities (Fig. 2B). Elytra
impunctate, pubescent as pronotum, covering all abdominal segments, weakly trun-
cate at apex; elytral suture flat; elytral striae absent; basomedial fovea present on each
elytron, fovea small, with dense inwardly directed setae (Figs. 1B, 3B). Humeral angles
of elytron raised, dorsally flattened (plateau-like), laterally rounded; weakly curved
posterad around anterolateral angle (Fig. 3B). Scutellum roundly triangular, without
setae (Fig 3B). Mesosternal keel setose with indentation around each seta; smooth in
the posterior quarter portion, apex straight (Fig. 2B). Metathoracic wings vestigial.
Femora strongly clavate in distal half, tibiae expanded and becoming more densely
setose towards distal half. Six visible abdominal sternites (fusion between ventrites V
and VI incomplete). Aedeagus with median lobe rather lightly sclerotized, bulbous
and deeply emarginate at base; apex with keel-like median dorsal process and dorsally
curved, apically convex digiform process extending over part of apical collar (Fig. 5B).

Female. Identical to male except frons flat.

Biology. This species has been collected from berlesed litters of oak, rotten wood,
mixed oak/conifer, and mixed hardwood. This species apparently favors moist habitats.

Distribution. This species has been collected in the mid-elevations of the central
Sierra Nevada, from localities in Amador and Calaveras Counties (Fig. 6).

Cephennium mariposae Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act:2E36C8F3-E54C-4CD3-AAF4-A5D366014F5E
Figs IO, 2G, 3G, 4G, 6

Type Material. Holotype (sex not determined): “CALIF: Mariposa Co.; Mariposa
[-37.4766°N, 119.9584°W] XI-15-1984; Berlese rotten log; A. J. Gilbert” (CSCA).

Etymology. ‘This species is named for the county and town from which it was col-
lected, Mariposa.

Diagnosis. This species can be distinguished from its California congeners by the
character combination of the presence of four ommatidia on each side of the head
(Fig. 4C), the mesosternal process truncate at the apex (Fig. 2C), and the humeral
angle of the elytron weakly raised and dorsally flattened (Fig. 3C). Cephennium mari-
posae is most similar to C. ce/sifrons but can be distinguished from it by the latter hav-
ing only two ommatidia on each side of the head (Fig. 2A) and a more elongate and
slender body form (Fig. 1B). This species can be easily separated from C. urbanum,
C. anopthalmicum, C. canestroi, C. grandarboreum and C. gilberti by the absence of
ommatidia and the apex of the mesosternal keel being divergent in these species. It
can be further distinguished from C. anophthalmicum, C. urbanum and C. aridum by
the humeral angle of the elytron projecting laterally to a point in these three species

(Fig. 3A, G-H).

42 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

Description. Length: 0.836 mm; pronotal width: 0.380 mm; elytral width:
0.437 mm. Body broad, ovate, slightly convex, testaceous, evenly densely pubescent,
pubescence golden, slender, moderate in length (Fig. 1C). Head small, deflexed,
sparsely pubescent, not narrowing anteriorly from antennal insertions; frons flat;
four ommatidia present on each side of head. Antenna setose, antennomere I and II
longer than broad, antennomeres II-VI quadrate and smaller than antennomeres II
and VII, antennomere VIII smaller than antennomeres VII and IX, antennomeres
IX-XI gradually clavate forming a loose club. Pronotum densely pubescent, broadest
between middle and anterior third, disc convex medially and weakly flattened near
each posterior angle; anterior margin not visible from above; anterior and posterior
margin lacking marginal bead; marginal bead complete laterally, gradually widening
towards base; lateral edge broadly rounded to posterior third, then evenly to base (Fig.
1C). Hypomeron smooth, sparsely setose towards upper quarter and along outside
(lateral) edge; hypomeral bead anterolaterad procoxae sinuate. Prosternum without
nodules anterolaterad procoxal cavities (Fig. 2C). Elytra impunctate, as pubescent as
pronotum, covering all abdominal segments, weakly truncate at apex; elytral suture
flat; elytral striae absent; basomedial fovea present on each elytron, fovea small, with
moderately dense inwardly directed setae (Figs. 1C, 3C). Humeral angles of elytron
raised, dorsally flattened (plateau-like), apically rounded, slender, moderately curv-
ing posterad around anterolateral angles (Fig. 3C). Scutellum roundly triangular,
without setae (Fig. 3C). Mesosternal keel setose with indentation around each seta,
apex not divergent (Fig. 2B). Metathoracic wings vestigial. Femora strongly clavate
in distal half, tibiae expanded and becoming more densely setose towards distal half.
Six visible abdominal sternites (fusion between ventrites V and VI). Aedeagus not
studied.

Female. Identical to male.

Biology. The lone specimen of this species was collected from a rotten log.

Distribution. This species is only known from Mariposa in Mariposa County, CA
(Fig. 6).

Cephennium grandarboreum Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act:8 CDOOFEF-266D-42A 1-8C2F-003 1C5EAAF06
Rigs 2D) SD 25, 6

Type Material. Holotype. Male. “CA: Monterey Co.; 36.2403°N, 121.7781°W;
LPNF: Sycamore Cyn.; II.14.2006; Caterino&¢Chatzimanolis; maple/redwood litter”/
“CA BEETLE PROJ; CBP0041534” (SBMN).

Paratypes (7): 3 specimens with same data as holotype (1 SBMN, CBP0041525
[disarticulated]; 1 FMNH, CBP0041544; 1 CASC, CBP0041516); “CA: Monterey
Co.; 36.0772°N, 121.5923°W; UC Big Creek Reserve; Redwood Camp, iii.28.2004.
M. Caterino, redwood litter” (1 CSCA, CBP0018437; 1 LACM, CBP0018450; 1
SBMN [in freezer for DNA]); “CA: Monterey Co.; 36.0812°N, 121.5974°W; UC Big

Seven new species of Cephennium Miiller & Kunze 43

Creek Reserve; BigCk/BrunetteCk.confl.; ii.7.2003, M. Caterino; redwood litter” (1
SBMN, CBP0006070 [gold coated for SEM)]).

Etymology. This species name is the combination of the Latin words grandis and
arboreus, meaning very large tree, as this species is associated with coast redwoods (Se-
quoia sempervirens).

Diagnosis. ‘This species is most similar to Cephennium gilberti but can be distin-
guished from it by its body size and shape. Cephennium grandarboreum is represented
by the largest specimens (0.988—1.121 mm long) of this genus currently known to
occur in California, and is very robust. C. gilberti is known from the smallest specimen
(0.646 mm long) of this genus known to occur in California and is much more slender
and elongate. These two species can also be separated by the mesosternal keel, which is
strongly divergent in C. grandarboreum, which has the apex ~2.4x as wide as the wid-
est anterior point (Fig. 2D). The mesosternal keel of C. gilberti is weakly divergent and
~2x as wide as the widest anterior point (Fig. 2F). Cephennium grandarboreum can be
separated from all other species by having two scutellar setae (Fig.3D) instead of four
(Fig. 3E) or zero (Figs 3A-C, G-H). Apart from the scutellar setae, C. grandarboreum
and C. canestroi are quite similar, with rounded elytral humeral angles, but the elytral
foveae are large and densely setose in C. grandarboreum (Figs. 1D, 2D), whereas they
are smaller and moderately setose in C. canestroi (Fig. 1E, 2E). The humeral angle of
the elytron is also more slender and posterorly curved in C. grandarboreum (Fig. 3D)
than in C. canestroi (Fig. 3E), and has a more strongly divergent mesosternal keel (Fig.
2D) than C. canestroi (Fig. 2E).

Description. Male. Length: 0.988—1.121 mm; pronotal width: 0.418—0.437 mm;
elytral width: 0.456-0.475 mm. Body broad, ovate, slightly convex, rufo-testaceous to
amber yellow, evenly densely pubescent, pubescence golden, slender, long (Fig. 1D).
Head small, deflexed moderately pubescent, weakly narrowing anteriorly from antennal
insertions; eyes absent. Antenna setose, antennomere I and I longer than broad, anten-
nomeres III-VI quadrate and smaller than antennomeres II and VI, antennomere VIII
smaller than antennomeres VII and IX, antennomeres [X-XI gradually clavate form-
ing a loose club. Pronotum densely pubescent, broadest between middle and anterior
third, disc very convex medially and weakly flattened near each posterior angle; anterior
margin not visible from above; anterior and posterior margin lacking marginal bead;
marginal bead complete laterally, gradually widening towards base; lateral edge broadly
rounded to posterior third, then weakly sinuate at base (Fig. 1D). Hypomeron smooth,
sparsely setose towards upper quarter and along outside (lateral) edge, hypomeral bead
anterolaterad procoxae with small knob at apex. Prosternum with small, weakly pro-
duced egg-shaped nodules anterolaterad procoxal cavities (Fig. 2D). Elytra impunctate,
as pubescent as pronotum, covering all abdominal segments, weakly truncate at apex;
elytral suture flat; elytral striae absent; basomedial fovea present on each elytron, fovea
large, deep, with dense inwardly directed setae (Fig. 1D, 3D). Humeral angle of elytron
raised laterad scutellum to humeral angle, humeral plateau slender, dorsally flattened,
apically rounded; apex reaching posterad past midline of elytral fovea (Fig. 3D). Scutel-
lum roundly triangular, with two setae (Fig. 3D). Mesosternal keel setose, texture ap-

44 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

pearing as fish scales anteriorly, abruptly smooth and impunctate near mesometasternal
suture, apex strongly bifid and divergent, posterior projections long (Fig. 2D). Metatho-
racic wings vestigial. Femora strongly clavate in distal half, tibiae expanded and becom-
ing more densely setose towards distal half. Five visible abdominal sternites (sometimes
appearing as six due to weak fusion between ventrites V and VI). Aedeagus with median
lobe bulbous, heavily sclerotized, median dorsal projection coming to point at apex (tri-
angular), reaching past apical collar; apical collar with dense setae surrounding opening;
dorsal parameres slender, longer than median lobe but not extending past apical collar,
with lateral subapical setae on each side; apical digiform process extending past apical
collar, bent ventrad near apex (Fig. 5C).

Female. Identical to male.

Biology. Specimens were extracted from redwood litter and a combination of ma-
ple and redwood litter with the use of Berlese funnels.

Distribution. ‘This species is known from a few localities near Big Sur, Monterey

County, CA (Fig. 6).

Cephennium canestroi Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act:55F6B353-76A2-4826-8D8E-6FD28BE5671C
Figs VE EOE. 31956

Type Material. Holotype. Male. “CA: San Luis Obispo Co.; 35.5249°N, 121.0719°W;
UC Rancho Marino Res.; II.26.2009, Salix litter; M.S. Caterino”/ “CA BEETLE
PROJ; CBP0087147” (SBMN).

Paratypes (5): “CA: San Luis Obispo Co.; 35.5361°N, 121.0723°W; UC Rancho
Marino Res.; III.10.2009, M.S. Caterino; Heteromeles litter” (2 SBMN, CBP0087853
[gold coated for SEM], CBP0087825 [disarticulated]; 1 CASC, CBP0087841; 1
CSCA, CBP0087830; 1 FMNH, CBP0087815 [male genitalia dissected out and gold
coated for SEM]).

Etymology. We are pleased to name this species for Don Canestro, director of the
Rancho Marino Reserve, in appreciation of his generous assistance with our fieldwork.

Diagnosis. This species can be separated from its California congeners by the char-
acter combination of the absence of eyes, humeral plateau of the elytra raised, dorsally
flattened and apically rounded, and scutellum with four setae (Fig. 3E). Cephennium
canestroi most closely resembles C. grandarboreum and C. gilberti, all having a generally
rounded humeral angle (Figs. 3D-F), but C. canestroi can be distinguished from them
by having four scutellar setae (Fig. 3E) instead of two (Fig. 3D, F). Cephennium canestroi
can be separated from C. anophthalmicum and C. urbanum by the shape of the elytral
humeral angle, which projects laterally to a point in the latter two species (Fig. 3G-H).

Description Male. Length: 0.836—0.922 mm; pronotal width: 0.314-0.323 mm;
elytral width: 0.361—0.380 mm. Body elongate, slender, weakly convex; amber-yellow,
transparent under light, evenly and moderately pubescent, pubescence golden, slender,
moderately long, weakly recumbent (Fig. 1E). Dorsal surface of head impunctate, weakly

Seven new species of Cephennium Miiller & Kunze 45

pubescent, weakly narrowing anteriorly from antennal insertions. Eyes absent. Antenna
setose, antennomeres I and II longer than broad, antennomeres II-VI quadrate and smaller
than antennomeres II and VU, antennomere VIII smaller than antennomeres VII and IX,
antennomeres [X-XI gradually clavate forming a loose club. Pronotum pubescent, broadest
between middle and anterior third, very convex in disc and weakly flattened near each pos-
terior angle; anterior margin not visible from above; anterior and posterior margin lacking
marginal bead; marginal bead complete laterally, gradually widening towards base; lateral
edge broadly rounded to posterior third, then sinuate to base (Fig. 1E). Hypomeron im-
punctate, sparsely setose towards anterior quarter and along outside (lateral) edge (Fig. 2E).
Prosternum with prominent egg-shaped nodules anterolaterad procoxal cavities (Fig. 2E).
Elytra smooth, as pubescent as pronotum, covering all abdominal segments; elytral suture
flat; elytral striae absent, basomedial fovea present on each elytron, fovea small, moder-
ately pubescent (Figs. 1E, 2E). Humeral angle of elytron raised from laterad scutellum
along anterior margin to humeral angle, humeral plateau broad, dorsally flattened, apically
rounded, not reaching past midline of elytral fovea (Fig. 3E). Scutellum nearly semicircular,
with four setae present (two on each side of the midline) (Fig. 3E). Mesosternal keel setose,
texture appearing like fish scales in its entirety until abruptly smooth in the posterior quar-
ter, weakly bifid at mesometasternal suture, divergent projections short (Fig. 2E). Metatho-
racic wings vestigial. Femora strongly clavate in distal half; tibiae expanded and becoming
more densely setose towards distal half. Five visible abdominal sternites, with partial fusion
between the apical two (ventrites V and VI). Aedeagus with median lobe bulbous, strongly
sclerotized, median dorsal projection parallel-sided at base, convergent and bisinuate at
apex, not reaching past apical collar; apical collar with dense setae meeting medially and
forming wick-like setal extension, curving ventrad; dorsal parameres slender, extending be-
yond apex of median lobe but not past apical collar, each with two subapical setae; apical
digiform process present, bent ventrad near apex (Fig. 5D).

Female. Identical to male.

Biology. ‘This species has been collected from Heteromeles (Toyon) and Salix (Wil-
low) litter. The collecting sites occur near a native stand of Monterey Pine (Pinus ra-
diata), and is along the coast near the town of Cambria. Collecting sites were in a rocky
drainage, with sparse shrubby willow cover, facing and only a few hundred meters from
the ocean, and on a shaded northeast facing slope under a very large toyon tree, with a
dense understory of Rubus and Toxicodendron.

Distribution. This species has been only collected from San Luis Obispo County,
CA, in the University of California Rancho Marino Reserve (Fig. 6).

Cephennium gilberti Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act:307 DD4D 1-D511-447D-AFF4-2258EEGDFC10
Bigs PE QELS EG

Type Material. Holotype (unknown sex): “CALIF: Kern Co.; 5 mi. E Glenville [sic]
[- 35.7486°N, 118.5805°W]; Alta Sierra Camp; III-12-1979; A.J. Gilbert”/ “Berlesed;

46 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

Oak; Litter” (CSCA). We did not risk dissecting the type to determine sex due to its
uniqueness and minute size.

Etymology. We name this species in honor of Dr. Art Gilbert, collector of the
unique specimens of two of the species described herein, including this one.

Diagnosis. ‘This species can be distinguished from its California congeners by the char-
acter combination of the eyes absent, humeral angle of the elytron raised, dorsally flattened
and apically rounded (Fig. 3F), scutellum with two setae (Fig. 3F), and the mesosternal
keel weakly divergent at the apex (Fig. 2F). This species most closely resembles C. grandar-
boreum, but can be distinguished from it by its smaller size (0.646 mm long), more elongate
and slender body (Fig. 1F), the posterior angles of the pronotum being sharper in C. gilberti
(Fig. 1F), and the mesosternal keel more weakly divergent at the apex with the apex ap-
proximately only 2x as wide as the widest point anterior to the apex (Fig. 2F), as opposed to
the apex strongly divergent and nearly 2.4x as wide as the widest point anterior to the apex
as in C. grandarboreum (Fig. 2D). C. gilberti can be separated from C. anophthalmicum, C.
urbanum and C. aridum by the humeral angle of the elytron, which projects laterally to a
point in these species (Fig. 3A, G-H). Finally, it can be distinguished from C. celsifrons, C.
mariposae and C. aridum by the presence of ommatidia in these species (Fig. 4A, C-D).

Description (sex unknown). Length: 0.646 mm; pronotal width: 0.228 mm; elytral
width: 0.266 mm. Body elongate, slender, weakly convex, amber yellow, evenly moderately
pubescent, pubescence golden, slender, moderate in length (Fig. 1F). Head small, deflexed,
sparsely pubescent; frons flat; eyes absent. Antenna setose, antennomeres | and II longer than
broad, antennomeres [II-VI quadrate and smaller than antennomeres II and VII, anten-
nomere VIII smaller than antennomeres VII and IX, antennomeres [X-XI gradually clavate
forming a loose club. Pronotum moderately pubescent, broadest between middle and ante-
rior third, disc very convex medially and weakly flattened near each posterior angle; anterior
margin not visible from above; anterior and posterior margin lacking marginal bead; mar-
ginal bead complete laterally, gradually widening towards base; lateral edge broadly rounded
to posterior third, then slightly sinuate to base (Fig. 1F). Hypomeron smooth, sparsely setose
towards upper quarter and along outside (lateral) edge, hypomeral bead anterolaterad pro-
coxae sinuate. Prosternum with prominent, bluntly triangular nodules anterolaterad pro-
coxal cavities (Fig. 2F). Elytra impunctate, as pubescent as pronotum, covering all abdominal
segments, weakly truncate at apex; elytral suture flat; elytral striae absent; basomedial fovea
present on each elytron, fovea large, with moderately dense inwardly directed setae (Figs. 1F
3F). Humeral angles of elytron raised, dorsal plateau apically rounded, slender, barely reach-
ing beyond anterior portion of the elytral fovae (Fig. 3F). Scutellum roundly triangular, with
two short setae, one on each side of the midline (Fig. 3F). Mesosternal keel setose with scale-
like microsculpture over most of length, abruptly smooth near meso-metasternal junction,
apex weakly divergent, angulate (Fig. 2F). Metathoracic wings vestigial. Femora strongly
clavate in distal half, tibiae expanded and becoming more densely setose towards distal half.
Six visible abdominal sternites (fusion between ventrites V and VI not evident).

Biology. ‘This species has been collected only from berlesed oak litter.

Distribution. ‘This species is known from a single locality five miles east of Glen-
nville, at about 1800m in the southern Sierra Nevada of Kern County, CA (Fig. 6).

Seven new species of Cephennium Miiller & Kunze 47

Cephennium urbanum Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act:5640D 12 1-C792-4274-B02B-01EA234F177E
Figs 1G, 2G, 3G, 5E, 6

Type Material. Holotype. Male. “CA: Los Angeles Co.; 34.0824°N, 118.5660°W;
Topanga SP, Santa Ynez; Cyn, III.30.2009; K.J. Hopp; Quercus! Platanus litter”/ “CA
BEETLE PROJ; CBP0088983” (SBMN).

Paratypes (20): 9 specimens with same data as holotype (8 SBEMN, CBP0088984
[male genitalia dissected], CBP0088982, CBP0088981, CBP0088989, CBP0088986,
CBP0088990, CBP0088988 [male genitalia gold coated for SEM], CBP0088987 [dis-
articulated]; 1 LACM, CBP0088985); “CA: Los Angeles Co.; 34.0768°N, 118.8163°W;
Santa Monica Mts:; Zuma Cyn, IV.14.2009; M.S. Caterino & K.J. Hopp; Heterome-
les litter” (2 SBMN, CBP0089582, CBP0089583 [gold coated for SEM]); “CA: Los
Angeles Co.; 34.0760°N, 118.8170°W; Santa Monica Mts:; Zuma Cyn, IV.14.2009;
M.S. Caterino & K.J. Hopp; Heteromeles! Ceanothus litter” (1 SBMN, CBP0089561);
“CA: Los Angeles Co.; 34.1258°N, 118.6567°W; SAMO, Calabasas Cold; Creek
Trail, [V.22.2009; K.J. Hopp & M. Polihronakis; Quercus! Neotoma litter” (2 SBMN,
CBP0090114, CBP0090115); “CA: Los Angeles Co.; 34.0561°N, 118.8800°W;
Charmlee Park, IV.14.2009; M.S. Caterino & K.J. Hopp; Quercus litter” (1 SBMN,
CBP0089831); “CA: Los Angeles Co.; 34.0723°N, 118.6846°W; Santa Monica Mts
NRA; Piuma Rd, HI.30.2009; K.J. Hopp, Quercus! Ceanothus/ Heteromeles litter” (1
SBMN, CBP0089190; 1 CSCA, CBP0089191); “CA: Los Angeles Co.; 34.0948°N,
118.7242°W; Santa Monica Mts:; Malibu Ck SP; Quercus/ Rhus litter, I.8.2009; M.S.
Caterino & K.J. Hopp” (1 FMNH, CBP0086147); “CA: Los Angeles Co.; 34.0969°N,
118.7276°W; Santa Monica Mts:; Malibu Ck SP; Quercus litter, I1.8.2009; M.S. Ca-
terino & K.J. Hopp” (1 CASC, CBP0086092); “Pasadena; Cal.”/ “H. C. FALL; COL-
LECTION” (1 MCZ©).

Etymology. The species name urbanum signifies the proximity of the Santa Monica
Mountains to the urban development of greater Los Angeles. These mountains repre-
sent an extremely important island of native biodiversity surrounded by development.

Diagnosis. ‘This species can be distinguished from its California congeners by the
character combination of the absence of eyes, humeral angles of elytral project laterally
to a sharp point (Fig. 3G), presence of a basal elytral sutural ridge (Fig. 1G), and the
apex of the mesosternal keel crescent-shaped (Fig. 2G). This species most closely re-
sembles C. anophthalmicum but they can be separated by the basal sutural ridge, absent
from C. anopthalmicum, and the apex of the mesosternal keel, which is divergent and
crescent-shaped in C. urbanum (Fig. 2G) and very weakly divergent and angulate in
C. anophthalmicum (Fig. 2A). It can be easily distinguished from C. celsifrons, C. mari-
posae and C. aridum by the presence of eyes in these species (Fig. 4A, C-D). Finally, it
can be distinguished from C. grandarboreum, C. canestroi and C. gilberti by the humer-
al angle of the elytron, which is raised and rounded in these three species (Fig. 3D-F).

Description. Male. Length: 0.760—0.874 mm; pronotal width: 0.304—0.342 mm;
elytral width: 0.352-0.399 mm. Body elongate, slender, weakly convex; amber-yellow,

48 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

translucent; evenly and moderately pubescent; pubescence golden, slender, moderately
long, weakly decumbent (Fig. 1G). Dorsal surface of head smooth, weakly pubescent, nar-
rowing anteriorly from antennal insertions. Eyes absent. Antennae setose, antennomeres |
and II longer than broad, antennomeres IIJ-VI quadrate and smaller than antennomeres
II and VII, antennomere VIII smaller than antennomeres VII and IX, antennomeres IX-
XI gradually clavate forming a loose club. Pronotum moderately pubescent, broadest be-
tween middle and anterior third, very convex in disc and moderately flattened near each
posterior angle; anterior margin not visible from above; anterior and posterior margin
lacking marginal bead; lateral marginal bead complete, gradually widening towards base;
lateral edge broadly rounded to posterior third, then weakly sinuate to base (Fig. 1G). Hy-
pomeron smooth, sparsely setose towards anterior quarter and along outside (lateral) edge.
Prosternum with large, protruding, obliquely oriented, ovate nodules anterolaterad pro-
coxal cavities (Fig. 2G). Elytra smooth, as pubescent as pronotum, covering all abdominal
segments; elytral sutural margin raised from base of scutellum to < 1/3 the length of elytra;
basomedial fovea present on each elytron; fovea small, moderately pubescent (Figs. 1G,
3G). Humeral angle of elytron projecting laterally to a sharp point (Fig. 3G). Scutellum
weakly triangular, lacking setae (Fig. 3G). Mesosternal keel setose, anterior quarter with
scale-like microsculpture, between coxae with large, round setose punctures, lacking scale-
like microsculpture, and with posterior quarter impunctate, apex weakly bifid (divergent),
divergent projections short, posterior margin crescent-shaped (Fig. 2G). Metathoracic
wings vestigial. Femora strongly clavate in distal half, tibiae expanded and becoming more
densely setose towards distal half. Five visible abdominal sternites, ventrites VI and VII
partially fused. Aedeagus with median lobe bulbous, heavily sclerotized, median dorsal
process triangular lacking parallel-sided base, not reaching past apical collar; apical collar
membranous, long, with thin dorsally and ventrally articulated projections; dorsal para-
meres slender, extending just beyond end of median lobe but not past apical collar, each
with two subapical seta; apical digiform process bent weakly ventrad near apex (Fig. 5E).

Female. Identical to male.

Biology. ‘This species has been sifted and extracted by Berlese funnels from Quercus
(oak) and Heteromeles (Toyon) litter as well as mixed litters of Quercus and Rhus (sumac),
Quercus and Salix (willow), Quercus and Platanus (sycamore), Quercus, Ceanothus (Cali-
fornia lilac), and Heteromeles, and from a Neotoma (woodrat) nest pile under Quercus.

Distribution. This species has mainly been collected in the Santa Monica Moun-
tains in Los Angeles County, CA, with a single record from “Pasadena in the western

foothills of the San Gabriel Mountains (Fig. 6).

Cephennium aridum Hopp & Caterino, sp. n.
urn:lsid:zoobank.org:act:859E5725-2579-406B-8E7A-7AB07AE6739B
Figs 1H, 2H, 3H, 4D, 5E 6

Type Material. Holotype. Male. “CA: San Bernardino Co.; 34.1405°N, 116.4541°W;
UC Burns Reserve; Railroad Cyn, IV.1.2008; sifted from grass & flood debris; M. Ca-

Seven new species of Cephennium Miiller & Kunze 49

debris; M. Caterino & R. Leschen”/ “CA BEETLE PROJ; CBP0072926” (SBMN
[gold coated for SEM]).

Paratypes (9): “CALIF: Riverside Co; 4mi SE Valle Vista; H-1978 K.Cooper; Oak
litter 78-57”/ “CEPHENNIUM; spp.; S. O7KEEFE DET “90” (1 SBMN); “CALIF:
Riverside Co; Palm Canyon Aqua [sic]; Caliente Reservation; IV-27-1978 78-67;
K.W.Cooper’/ “berlese litter; at base of Washingtonia filifera” (1 CSCA); “CALIF:
Riverside Co.; Whitewater cyn. Palm Oasis; V-4-1987 to XI-27-1988; Rolf L. Aalbu
coll.”/ “Ethylene-glycol pitfall; trap at base of Washingtonia filifera” (2 CSCA); “CAL-
IF: Riverside Co.; 2 mi. E Gilman Hot; Springs, Lamb Canyon; V-18-1978/XI-13-
1979; Fred G. Andrews; Antifreeze pit trap” (5 CSCA).

Etymology. ‘This species name refers to its occurrence in the arid regions of the
western Mojave/Colorado desert transition zone.

Diagnosis. This species can be immediately distinguished from all of its California
congeners by the presence of a single ommatidium on each side of the head (Fig. 4D).
Besides this somewhat cryptic character, C. aridum shares a sharply angulate humeral
process only with C. urbanum. The mesosternal keel of C. aridum, however, has scale-
like microsculpture between the mesocoxae (Fig. 2H), whereas that of C. urbanum has
only setose punctures between the coxae, its scale-like microsculpture being restricted
to the extreme anterior end (Fig. 2G). The mesosternal process of C. urbanum is also
distinctly crescent-shaped (Fig. 2G), where that of C. aridum is angulate (Fig. 2H).

Description. Male. Length: 0.817—0.912 mm; pronotal width: 0.304—0.342
mm; elytral width: 0.352-0.418 mm. Body elongate, slender, weakly convex;
amber-yellow, translucent; evenly and moderately pubescent; pubescence golden,
slender, moderately long, weakly decumbent (Fig. 1H). Dorsal surface of head
smooth, sparsely pubescent, narrowing anteriorly from antennal insertions. Sin-
gle ommatidium present on each side of head (Fig. 4D). Antenna setose, anten-
nomeres I and II longer than broad, antennomeres II-VI quadrate and smaller
than antennomeres II and VII, antennomere VIII smaller than antennomeres VII
and IX, antennomeres IX-XI gradually clavate forming a loose club. Pronotum
moderately pubescent, broadest between middle and anterior third, very convex
in disc part and moderately flattened near each posterior angle; anterior margin
not visible from above; anterior and posterior margin lacking marginal bead; mar-
ginal bead complete laterally, gradually widening towards base; lateral edge broadly
rounded to posterior third, then weakly sinuate to base (Fig. 1H). Hypomeron
smooth, sparsely setose towards anterior quarter and along outside (lateral) edge.
Prosternum with small, weakly protruding, oblique narrowly ovate nodules ante-
rolaterad procoxal cavities (Fig. 2H). Elytra smooth, lacking striae, as pubescent
as pronotum, covering all abdominal segments; elytral suture absent; basomedial
fovea present on each elytron; fovea small, moderately pubescent (Figs. 1H, 3H).
Humeral angle of elytron projecting laterally to sharp point (Fig. 3H). Scutel-
lum weakly triangular, lacking setae (Fig. 3H). Mesosternal keel sparsely setose,
with scale-like microsculpture over most of its length, abruptly smooth in posterior
extreme, apex moderately bifid, divergent projections short, diverging at ~90° an-

50 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

OC. anophthalmicum
O C. celsifrons

+ C. mariposae

¥& C. grandarboreum

AC. canestroi
@ C. gilberti
@ C. urbanum
@ C. aridum

Figure 6. Distribution map for all California species of Cephennium.

gle (Fig. 2H). Metathoracic wings vestigial. Femora strongly clavate in distal half;
tibiae expanded and becoming more densely setose towards distal half. Five visible
abdominal sternites, ventrites V and VI largely fused. Aedeagus with median lobe
bulbous, heavily sclerotized, truncate at apex, with very short median dorsal proc-
ess, weakly emarginate for reception of apex of apical digiform process; apical collar
not apparent; dorsal parameres slender, extending well beyond end of median lobe,
each with two subapical setae (Fig. 1H).

Female. Unknown.

Biology. The type of this species was sifted from grass and flood debris in a small,
spring-fed, but frequently dry drainage. Other specimens were sifted from litter of the
native California Fan Palm (Washingtonia filifera), which also occur in scattered spring-
fed drainages in the Colorado (and broader Sonoran) desert.

Distribution. This species is known from several localities in the far northeastern

Colorado desert, in San Bernardino and Riverside Counties, CA (Fig. 6).

Seven new species of Cephennium Miiller & Kunze 51

Key to the species of California Cephennium Miller and Kunze

Eyesapresent, reduced (Pig, AAs CD ))ae wc ivecaues cntestiveuer bineneeerenensaunnvnceniens Z
FV CRTAD SEIN cards phase Sey haalae oneese cideedstnaneremnterus pauet rete phichanr eds ietoe epic tne ciate &
Mesosternal process straight, not divergent at apex (Fig. 2B); frons of males
with medial ridge (Fig. 4B) ............ C. celsifrons Hopp & Caterino, sp. n.
Mesosternal process divergent at apex (Fig. 2A, D-H); frons of males flat,
WAtHOUL MECHA Toe Msc au micled Avecpnelee Masur con atieee an edapel vnsunt chon edteeh read 3
Single ommatidium present on each side of head (Fig. 4D); humeral angle of
elytron apically pointed (Fig. 3H); body elongate, slender... eee

OE Fes ad Ro oa AME Rta) TE ae 8 C. aridum Hopp & Caterino, sp. n.
Four ommatidia present on each side of head (Fig. 4C); humeral angle of
elytron apically rounded (Fig. 3C); body robust........ceceseseeecseeeeeseeeseeseees

dec Rach ated lta nh theD el Pae nas 2b C. mariposae Hopp & Caterino, sp. n.
Humeral angle of elytron apically pointed (Fig. 3A, G-H) oo. eeeeeeee 5
Humeral angle of elytron apically rounded (Fig. 3B-F) wees ee eeeereeees 6
Basal elytral sutural ridge present (Fig. 1G); apex of mesosternal keel cres-
ecnt-shaped. (MicP 2G) s cstt vena acssusire C. urbanum Hopp & Caterino, sp. n.
Basal elytral sutural ridge absent; apex of mesosternal keel triangulate (Fig.
21k Mee EPI ea Mh: Pek Aa AR PO C. anophthalmicum Brendel
Seutellimwithetworsetac dees SN: BY flees wanala inns Ruadbaitenss'suate's bapaldti be chensieseead 7
Scutellum with four setae (Fig. 3E) ...... C. canestroi Hopp & Caterino, sp. n.
Body robust; moderately large, 0.988—1.121 mm in length; mesosternal keel
strongly divergent at apex, apex 2.4x as wide as widest point anteriorly (Fig.
2D); scutellar setae long, extending past posterior margin of scutellum (Fig.
DUD ces sessaies eset ouadeastvsn A ieazsshes C. grandarboreum Hopp & Caterino, n.sp.
Body elongate, slender; very small, 0.646 mm in length; mesosternal keel
weakly divergent at apex, apex 2x as wide as widest point anteriorly (Fig. 2F);
scutellar setae short, not extending past posterior margin of scutellum (Fig.

BE \ivitedaaet udhenns Mpowat doovets Baxetecboncpanpsenp ies C. gilberti Hopp & Caterino, n.sp.

Checklist of the California species of Cephennium Miller and Kunze

Cephennium anophthalmicum Brendel, 1889
Alameda Co., Marin Co. and San Mateo Co.

Cephennium celsifrons Hopp & Caterino, sp. n.
Amador Co. and Calaveras Co.

Cephennium mariposae Hopp & Caterino, sp. n.
Mariposa Co.

Cephennium grandarboreum Hopp & Caterino, sp. n.
Monterey Co.

Cephennium canestroi Hopp & Caterino, sp. n.

52 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

San Luis Obispo Co.

Cephennium gilberti Hopp & Caterino, sp. n.
Kern Co.

Cephennium urbanum Hopp & Caterino, sp. n.
Los Angeles Co.

Cephennium aridum Hopp & Caterino, sp. n.
Riverside Co. and San Bernardino Co.

Discussion

The discovery of these new species constitutes a significant expansion of the known
range of Cephennium, and Cephennini in general, in the New World. Previously
represented by a single species, confined to moist coastal forests of central Califor-
nia, Cephennium appeared to have a minimal, possibly relictual, presence in North
America. The discovery of several additional species, across widely separated and
ecologically varied habitats instead indicates that the group has undergone consid-
erable diversification in the region. ‘The type localities of C. grandarboreum and C.
canestroi, in Monterey and San Luis Obispo Counties, alone would represent only
a slight extension of range down the coast, into an area otherwise known to host
several southernmost records for various beetles (Caterino, unpublished data). The
cool and foggy coastal environments conceivably provide similar microhabitats to
those inhabited by the genus further northward. The discovery of Cephennium in
the Santa Monica Mountains, however, represents a major disjunction, not only
in distance, but into much drier environments, mostly dominated by chaparral
and relatively sparse oak woodland. The presence of C. aridum on the fringes of
the Colorado desert takes this to an extreme, occurring in areas receiving less than
10 inches of rain/year. Finding additional, localized species in the Sierra Nevada
also extends the range of the genus into true montane habitats. Together these new
localities indicate surprising adaptability in ecological requirements, and suggest
that many other populations of these beetles will be discovered through diligent
searching.

Aside from the above habitat-related observations, we know very little about the
natural history of these beetles, as all were collected through Berlese extraction of sifted
leaf litter. The litters represent a variety of plants and plant communities, from fan
palm oases to chaparral, redwood forest, and oak woodland. In these habitats, species
of Cephennium are expected to be armored mite predators, with suction disks on their
labium, and rasp-like mandibles to slowly grind a hole in the thick dorsal surface of
the mite (Schmid 1988; Newton and Franz 1998; O’Keefe 2001; Jatoszyriski 2009).
We have observed similar labial suction disks to those described for European species.
Thus, it is probably a safe speculation that the new species are also specialist oribatid
mite predators.

Seven new species of Cephennium Miiller & Kunze 53

The biogeographic history of the California Floristic Province is complex and
poorly understood. While broad scale distributional patterns have long been evi-
dent, understanding the finer scale movements of lineages through time has suffered
from very fragmentary data on distribution patterns for those smaller, more seden-
tary taxa that would be most expected to reflect deep history. The cryptic fauna of
leaf litter stands to greatly increase not only our knowledge of gross biodiversity, but
also the evolutionary processes responsible for generating and maintaining it. The
insect fauna of this region in general remains woefully underdocumented. Increased
survey effort focusing on the more cryptic elements of the regional fauna would be
repaid many times over in biogeographic understanding and in management ap-
plicability.

Acknowledgements

We are grateful to the managers of the University of California Big Creek Reserve
(John Smiley, Kurt Merg, Feynner Arias), the UC Burns Pifion Ridge Reserve (Wil-
liam Bretz) and the UC Kenneth S. Norris Rancho Marino Reserve (Don Can-
estro), the Santa Monica Mountains National Recreation Area (Lena Lee), and the
California Department of Fish & Game for field permissions. We are also grateful
to Stylianos Chatzimanolis, Richard Leschen, and Maxi Polihronakis for field as-
sistance. This study could not have been completed without the response to loan
requests from all of the curators and collection managers who facilitated specimen
loans (listed above under Materials and methods). Finally, we are appreciative of
discussion of male genitalia morphological terminology with Al Newton. This work
was supported in part by National Science Foundation grants DEB0447694 and
MRI0420726 to M. Caterino, and the Santa Monica Mountains Fund, a non-profit
organization working in partnership with the National Park Service, grant to M.

Caterino and K. Hopp.

References

Brendel E (1889) Descriptions of new Scydmaenidae and Pselaphidae. Entomologica Ameri-
cana 5: 193-197. http://www. biodiversitylibrary.org/item/42392

Caterino M (2006) California beetle faunistics: 100 years after Fall. Coleopterists Bulletin
60(2): 177-191.

Caterino M, Chatzimanolis S (2007) Newly recorded and noteworthy California Staphylini-
dae. Coleopterists Bulletin 61:398—407.

Caterino M, Leschen RAB, Johnson C (2008) A new genus of Caenoscelini (Cryptophagi-
dae: Cryptophaginae) from California, with two new species. Coleopterists Bulletin 62(4):
509-523.

54 Katie J. Hopp & Michael S. Caterino / ZooKeys 24: 31-54 (2009)

Evenhuis N (2008) The insect and spider collections of the world website. Available at: http://
hbs.bishopmuseum.org/codens/ [Last accessed: August 11, 2009].

Grebennikov VV, Newton AF (2009) Good-bye Scydmaenidae, or why the ant-like stone beetles
should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal
of Entomology 106: 275-301. http://www.eje.cz/pdfarticles/ 145 1/eje_106_2_275_Gre-
bennikov.pdf

Ivie MA (1985) Nomenclatoral notes on West Indian Elaphidiini (Coleoptera: Cerambycidae).
Pan-Pacific Entomologist 61(4): 303-314.

Jatoszyniski P (2007) The Cephenniini of China. II. Cephennodes Reitter of southern provinces,
with taxonomic notes on the Cephennodes-Chelonoidum complex (Coleoptera: Scydmaeni-
dae). Genus 18(1): 7-101.

Jatoszynski P (2009) Trurlia, a new Oriental genus of the tribe Cephenniini (Coleoptera: Scy-
dmaenidae). European Journal of Entomology 106: 261-274.

Majka C, Klimaszewski J (2004) Phloeocharis subtilissima Mannerheim (Staphylinidae: Phloeo-
charinae) and Cephennium gallicum Ganglbauer (Scydmaenidae) new to North America: a
case study in the introduction of exotic Coleoptera to the port of Halifax, with new records
of other species. Zootaxa 781: 1-15. http://www.mapress.com/zootaxa/2004f/zt00781.pdf

Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity
hotspots for conservation priorities. Nature 403: 853-858.

Newton AK, Franz H (1998) World catalog of the genera of Scydmaenidae. Koleopterologische
Rundschau 68: 137-165.

O’Keefe ST (2001) Scydmaenidae Leach, 1815. In: Arnett, R. H., M. C. Thomas (Eds), Vol.
1 American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia.
CRC Press, Boca Raton, Florida, 259-266.

Schmid R (1988) Morphologische Anpassungen in einem Rauber-Beute-System: Ameisenkafer
(Scydmaenidae, Staphylinoidea) und gepanzerte Milben (Acari). Zoologische Jahrbiicher,
Abteilung fiir Systematik, Okologie und Geographie der Tiere 115: 207—228.