ZooKeys 524: 103—I 36 (20 | 5) A peer-reviewed open-access journal
doi: 10.3897/zookeys.524.6105 RESEARCH ARTICLE #ZooKey S

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A review of Canadian and Alaskan species of the
genera Clusiota Casey and Atheta Thomson,
subgenus Microdota Mulsant & Rey
(Coleoptera, Staphylinidae, Aleocharinae)

Jan Klimaszewski', Reginald P. Webster’, Derek Sikes,
Caroline Bourdon', Myriam Labrecque'

| Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, 1055 du PE.PS., RO. Box
10380, Stn. Sainte-Foy, Québec, Quebec, Canada GIV 4C7 2 24 Mill Stream Drive, Charters Settlement,
New Brunswick, Canada E3C 1X1 3 University of Alaska Museum, 907 Yukon Dr., Fairbanks, Alaska, USA,
99775-6960

Corresponding author: Jan Klimaszewski (jan. klimaszewski@canada.ca)

Academic editor: Volker Assing | Received 2 July 2015 | Accepted 2 September 2015 | Published 30 September 2015
http://zoobank. ore/FO7CD 1B4-D0B0-4048-837E-1B731168C5EC

Citation: Klimaszewski J, Webster RP, Sikes D, Bourdon C, Labrecque M (2015) A review of Canadian and Alaskan
species of the genera Clusiota Casey and Atheta Thomson, subgenus Microdota Mulsant & Rey (Coleoptera, Staphylinidae,
Aleocharinae). ZooKeys 524: 103-136. doi: 10.3897/zookeys.524.6105

Abstract

This paper treats 13 species of the subgenus Microdota Mulsant & Rey of Atheta Thomson and 3 species
of the genus Clusiota Casey in Canada and Alaska. We report here 4 species new to science, and 3 new
provincial records. The following species are new to science: A. (M/.) curtipenis Klimaszewski & Webster,
sp. n., A. (M.) formicaensis Klimaszewski & Webster, sp. n., A. (Z.) macesi Klimaszewski & Webster,
sp. n., and Clusiota grandipenis Klimaszewski & Webster, sp. n. The new provincial records are: A. (/.)
pseudosubtilis Klimaszewski & Langor, new to AB, and A. (.) subtilis (Scriba), an adventive Palaearctic
species new to North America, first reported in LB and NB. The two Clusiota Casey species are reviewed,
and their distribution is revised. A female C. impressicollis was discovered in Ontario and is illustrated here
for the first time. A key to all Canadian species of the subgenus Microdota and genus Clusiota are provided.
Atheta (Microdota) holmbergi Bernhauer and A. (M.) alesi Klimaszewski & Brunke are transferred here to
the subgenus Dimetrota Mulsant & Rey.

Keywords

Alaska, Canada, Coleoptera, Staphylinidae, Aleocharinae, Clusiota, Microdota, new records, adventive species

Copyright Her Majesty the Queen in Right of Canada. This is an open access article distributed under the terms of the Creative Commons Attribution
License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

104 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Introduction

Aleocharines are species rich in the boreal forest of Canada but knowledge of them,
despite recent progress (Klimaszewski et al. 2015), is still fragmentary and there are
many species likely to be discovered as new to science or as new records of adventive or
formerly known species from the USA (Klimaszewski et al. 2015).

This paper deals with Canadian species of the subgenus Microdota Mulsant and
Rey of the genus Atheta Thomson and Rey, and the genus C/usiota Casey occurring in
Canada and Alaska. The subgenus Microdota contains about 215 species in the Palae-
arctic region (Lee and Ahn 2015). In the Nearctic region the true number of species
is unknown but Ashe (2000) reported 27 species. Microdota species may be confused
with those of Clusiota due to their small size, and other superficial similarities. That is
why both groups are treated here. Microdota species may also be confused with mem-
bers of the subgenus Datomicra Mulsant and Rey from which they may be separated
by having a fully exposed pronotal hypomeron in lateral view, whereas in Datomicra
it is only partially exposed (Seevers 1978, Ashe 2000). In Canada we recognize 13
Microdota species including 3 species described here as new to science, and 3 species
of Clusiota, including one species new to science. These C/usiota species constitute all
known Nearctic species of the genus. We provide diagnoses of new or newly recorded
species, illustrations of habitus and genital structures of all Microdota species, and keys
to their identification. We hope that this publication will lead to the proper identi-
fication of species in this difficult group and will make them available for ecological,
environmental, and other studies.

Materials and methods

About 100 adults of the genera Microdota and Clusiota from Canada and Alaska were
studied, and most specimens were dissected to examine the genitalic structures that
were dehydrated in absolute alcohol, mounted in Canada balsam on celluloid mi-
croslides, and pinned with the specimens from which they originated. Images of the
entire body and the genital structures were taken using an image processing system
(Nikon SMZ 1500 stereoscopic microscope; Nikon Digit-like Camera DXM 1200F
and Adobe Photoshop software).

Morphological terms mainly follow those used by Seevers (1978), Ashe (2000),
and Klimaszewski et al. (2011). The ventral side of the median lobe of the aedeagus is
considered to be the side of the bulbus containing the foramen mediale, the entrance
of the ductus ejaculatorius, and the adjacent ventral side of the tubus of the median
lobe with internal sac and its structures (this part is referred to as the parameral side
in some recent publications); the opposite side is referred to as the dorsal part. In the
species descriptions, microsculpture refers to the surface of the upper forebody (head,
pronotum and elytra).

A review of Canadian and Alaskan species of the genera Clusiota Casey... 105

Distribution. Each species is cited with its currently known distribution in Cana-
da and Alaska. The following abbreviations are used in the text for Canadian provinces
and territories:

AB — Alberta, BC — British Columbia, LB — Labrador, MB — Manitoba, NB — New
Brunswick, NF — Newfoundland (island), NS — Nova Scotia, NT — Northwest Ter-
ritories, NU — Nunavut, ON — Ontario, PE — Prince Edward Island, QC — Quebec,
SK — Saskatchewan, YT — Yukon Territory.

USA state abbreviations follow those of the US Postal Service.

Two labels were used on some specimens (RWC), one that included the locality,
collection date, and collector, and one with macro and micro habitat data and collec-
tion method. Information from the two labels is separated by a // in the data presented
for these specimens.

Depository/institutional abbreviations

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture
and Agri-Food Canada, Ottawa, Ontario, Canada.

LFC Natural Resources Canada, Canadian Forest Service, Laurentian Forestry
Centre, R. Martineau Insectarium, Quebec City, Quebec, Canada.

RWC _ Reginald Webster Collection, Charters Settlement, New Brunswick, Canada.

UAM_ University of Alaska Museum, University of Alaska, Fairbanks, Alaska, U.S.A.
http://dx.doi.org/doi:10.7299/X75D8S0H

ZMB_ Zoological Museum of Humboldt University, Berlin, Germany.

ZML Museum of Zoology, Lund University, Lund, Sweden.

Checklist of Canadian Microdota and Clusiota species

New jurisdictional records are indicated in bold type.

Genus Atheta Thomson, 1858

Subgenus Microdota Mulsant & Rey, 1873

1) Atheta (Microdota) amicula (Stephens, 1832). Palaearctic; adventive in Canada:
NF, NS. USA: WA.

2) Atheta (Microdota) curtipenis Klimaszewski & Webster, sp. n. Canada: NB.

3) Atheta (Microdota) festinans (Erichson, 1839). Canada: ON. USA: IN, ME, MI, PA.

4) Atheta (Microdota) formicaensis Klimaszewski & Webster, sp. n. Canada: NB.

5) Atheta (Microdota) macesi Klimaszewski & Webster, sp. n. Canada: NB.

6) Atheta (Microdota) microelytrata Klimaszewski & Godin, 2012. Canada: YT.

7) Atheta (Microdota) pennsylvanica Bernhauer, 1907. Canada: LB, NB, NF, NS,
ON, QC. USA: IN, PA, RI, VA.

106 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

8) Atheta (Microdota) platonoffi Brundin, 1948. Holarctic; Canada: AB, BC, LB, NB,
NF, NS, ON, YT. USA: AK.

9) Atheta (Microdota) pratensis (Maklin, 1852). USA: AK, WA.

10) Atheta (Microdota) pseudosubtilis Klimaszewski & Langor, 2011. Canada: AB, LB,
NB, NF, QC, YT.

11) Atheta (Microdota) riparia Klimaszewski & Godin, 2012. Canada: YT.

12) Atheta (Microdota) sculptisoma Klimaszewski & Langor, 2011. Canada: NF, QC.

13) Atheta (Microdota) subtilis (Scriba, 1866). Palaearctic, adventive in Canada: LB, NB.

Species transferred to the subgenus Dimetrota Mulsant & Rey
14) Atheta (Microdota) alesi (Klimaszewski & Brunke, 2012). Canada: ON.
15) Atheta (Microdota) holmbergi Bernhauer, 1907. Canada: BC. USA: AK.

Species removed from NF and LB species list (misidentification for A. subtilis Mulsant
and Rey)
16) Atheta (Microdota) pratensis (Maklin, 1852). Canada: YT. USA: AK.

Subgenus Clusiota Casey, 1910

17) Clusiota antennalis Klimaszewski and Godin, 2008. Canada: BC. USA: AK.
18) Clusiota impressicollis (Bernhauer, 1907). Canada: BC, ON, NB, NF.

19) Clusiota grandipenis Klimaszewski and Webster, sp. n. Canada: NB.

Taxonomic review

Genus Atheta Thomson

Subgenus Microdota Mulsant & Rey
(Mouthparts illustrated by Lee and Ahn 2015)

Diagnosis. The following combination of characters is distinctive for Microdota: small
and subparallel body (Figs 1, 9, 17, 25, 33, 41, 49, 56, 61, 69, 80, 84), length 1.5-2.7
mm, antennomere | enlarged but not extremely swollen, longer than II, II longer than
III, V-X slightly to strongly transverse; median region of prementum very narrow
and without pseudopores; labial palpus with 3 articles; glossa split apically forming
Y-shaped structure; maxillary palpus with 3 articles, last one narrowly elongate; pro-
notum transverse, more than 1.2 times as wide as long, midline pubescence directed
in most specimens anteriorly and laterad elsewhere (Figs 1, 9, 17, 25, 33, 41, 49, 56,
61, 69, 80, 84); pronotal hypomeron fully visible laterally; elytra in some species with
wavy pattern of pubescence in postero-sutural section of disc (Figs 33, 49); median
lobe of aedeagus with large bulbus and triangularly shaped apex of tubus in dorsal
view, internal sac of median lobe of aedeagus with well-developed complex structures

A review of Canadian and Alaskan species of the genera Clusiota Casey... 107

(Figs 4, 5, 12, 13, 20, 21, 28, 29, 36, 37, 44, 45, 51, 52, 64, 65, 72-75, 87); sper-
matheca of variable shape, L- or S-shaped, capsule usually in a form of a narrow sac
or club-shaped, and stem often sinuate (Figs 8, 16, 24, 32, 40, 48, 55, 59, 60, 68, 78,
79, 83); male tergite VIII in most species simple, truncate apically and without teeth,
sometimes with minute crenulation and small pairs of teeth (Figs 2, 10, 18, 26, 34,
42,50, 62, 70, 85).

Some species of Microdota, due to small body size and some superficial external
similarity, may be confused in collections with members of the subgenus Datomicra
Mulsant and Rey, from which they may be distinguished by having a fully exposed
pronotal hypomeron in lateral view; the pronotal hypomeron is only partially visible in
Datomicra. Many species of Datomicra also have a more densely and coarsely punctate
forebody than that of Microdota.

Microdota may be distinguished from Dimetrota by the following combination of
characters: body usually parallel-sided, small, on average 2 mm long (Microdota — 1.5—
2.8 mm; Dimetrota — 1.8-3.8 mm, with elytra usually distinctly broader than prono-
tum); glossae Y-shaped (deeply split in Dimetrota); pronotum with sparse to moderately
dense and slightly asperate punctation (dense and strongly asperate in Dimetrota); lat-
eral margins of pronotum and elytra, and middle and hind tibiae with moderately pro-
nounced macrosetae (strong bristles in Dimetrota); hypomera fully visible in lateral view
(partially to less often fully visible in Dimetrota); and male tergite VII truncate apically,
rarely crenulated, and usually without large lateral teeth (with two large lateral teeth
and often distinctive form of margin between them or with pattern of smaller teeth in
Dimetrota). Details on diagnostics of Microdota are provided by Brundin (1948), and
Lee and Ahn (2015). Species of Nearctic Dimetrota badly need revision.

Clusiota may be distinguished from Microdota by the following combination of
characters: basal antennal article swollen (some species); antennal articles V-X strongly
transverse; glossae deeply split medially; pronotum narrower than elytra; elytra flat-
tened, truncate posteriorly and without distinct lateral emargination; abdomen often
swollen; spermatheca more or less sinuate with narrowly pear-shaped capsule and small
and short apical invagination; and by the median lobe of the aedeagus with large bul-
bus, strongly ventrally produced tubus bearing elongate subapical part, and with crista
apicalis located on elevated part of bulbus.

Key to Canadian and Alaskan species of Microdota

1 Elytra at suture about as long as or shorter than pronotum (Figs 9, 17, 25, 56,
GARD AT Nias. eee IO ache BRD aaah at Chae OM ULE sachs RAO SLAM 2
= Elytra at suture longer than pronotum (Figs 1, 33, 41, 49, 80, 84)... 7

Body bicoloured, head and abdomen dark brown, and pronotum and elytra
or elytra only yellowish brown or orange brown (Figs 61, 69); genitalia as il-
hustraredi(higs G4, GO Ss 72279 Oi Uy sssraates su slesonvencs oxseccsthevecos veces uses 3
- Body approximately uniformly brown to black; genitalia differently shaped.. 4

108

Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Pronotum in most specimens orange and elytra yellowish brown (Fig. 69);
median lobe of aedeagus and spermatheca as illustrated (Figs 72-75, 78, 79)...
Riana Ao Re eh aa set ene ecient oes Atheta (Microdota) pennsylvanica Bernhauer
Pronotum brown to light brown and entire elytra or only central part of disc
yellowish (Fig. 61); median lobe of aedeagus and spermatheca as illustrated
(Pigsi643- 65268) eri iwnthct ona Sate Ate taent h ecaee ce pee dete cea Ee
each Atheta (Microdota) formicaensis Klimaszewski & Webster, sp. n.
Elytra distinctly broader than maximum width of pronotum (Figs 17, 56);

genitalia asalltistrated: (Ries 20,21, 24) 595-00) vieiccss ecenveaeecosteveneetoncostmentes 5
Elytra about as wide as maximum width of pronotum (Figs 9, 25); genitalia
ditterentivishaped! (Fissal) <aS AG; 28,0955 2) 4.1). cen tess th oc iis: AOR 6

Pubescence on forebody sparse, on elytra directed straight posteriad (Fig. 56);
spermatheca as illustrated (Figs 59, 60); male unknown... eee eeeeeeeeeeereeeeee
Aen Rr RAR es Dts Bed Oboe Weces a Atheta (Microdota) festinans (Erichson)
Pubescence on forebody dense, on elytra directed obliquely posteriad (Fig.
7) eenttaliacas illustrated (Figs D021 D4) i oh wets ne cect etary
ERPS sss Atheta (Microdota) curtipenis Klimaszewski & Webster, sp. n.
Pronotum as broad as head (Fig. 25); abdomen subparallel (Fig. 25); anten-
nal articles VI-X moderately transverse (Fig. 25); genitalia as illustrated (Figs
2B 9232) hel cee Atheta (Microdota) microelytrata Klimaszewski & Godin
Pronotum broader than head (Fig. 9); abdomen swollen apically (Fig. 9);
antennal articles VI-X strongly transverse (Fig. 9); genitalia as illustrated
LST Feas5) BZ DOL Tar 6) i Decne eT ne eRe OR eva CuURU ec PPRT td a, Onc Rey Mie CURT ORT RATS
heated be sk Atheta (Microdota) pseudosubtilis Klimaszewski & Langor
Elytra almost twice as long as pronotum; median lobe of aedeagus with en-
larged oval bulbus and narrow tubus that is ventrally produced at apex; sper-

PALME CARS ACEH ob ee Near eadee ae? Atheta (Microdota) pratensis (Maklin)
Elytra 1.2-1.3 times longer than pronotum (Figs 1, 33, 41, 49, 80, 84); geni-
talia differenthy-shapediis.ccsezse:aonetacceneasenaictt evs scanner Manes celacuscerdanetmenehhecstesoate 8

Pronotum dark brown to black and elytra light brown to yellowish-brown
(Fig. 49); antennae with articles VII-X strongly transverse, at least twice as
wide as long (Fig. 49); pubescence of forebody not soft in appearance; geni-
talia as illustrated (Figs 51, 52, 55) [males absent in North America]............
pater TREN Lp Maer ta cel Lae bench Atheta (Microdota) amicula (Stephens)
Pronotum and elytra about the same colour, dark brown with elytra in some
specimens only slightly paler (Figs 1, 33, 41, 80, 84) [group of species dif-
heultte cis tirvouishrextenial lytic. tes rete enueertasned Ae NAS le Sidhe ey 9
Elytra 1.2 times broader than pronotum, shoulders moderately angular (Fig.
80); spermatheca with broad, sac-shaped capsule without apparent invagina-
tion, stem straight, narrow, half looped posteriorly and slightly twisted at
apex (Fig §83)5 male wi wWar sh ac ceaenrcctnchce cpus Meets Meant ny tencshaeueneapencenteconee son’
w Seasbanericatsneseeates Atheta (Microdota) sculptisoma Klimaszewski & Langor

10

11]

12

A review of Canadian and Alaskan species of the genera Clusiota Casey... 109

Elytra at least 1.3 times broader than pronotum, shoulders strongly angular
(Pigs 33741584); spermathecaofasditierent shape. sactevescedasbaccidunerast 10
Pronotal punctation coarse and sparse (Fig. 41); spermatheca with club-
shaped capsule bearing deep apical invagination, stem sinuate and twisted
apically (Fig. 48); median lobe of aedeagus in lateral view with approximately
oval bulbus dorsally (Fig. 45), and sinuate and narrowly elongate tubus
laterally (Fig. 44) ....... Atheta (Microdota) riparia Klimaszewski & Godin
Pronotal punctation fine and dense (Figs 1, 33, 84); genitalia of a different
SETS e tase, Manet rc pe esse acct catac eetae teste Betas ae sete peel ne ee ac pecan tena 11
Body dark brown, almost black, strongly glossy, with dense, meshed and
strongly pronounced microsculpture, punctation and pubescence sparse (Fig.
84); male tergite VIII truncate apically, with two small lateral teeth and min-
ute crenulation on apical margin of disc (Fig. 85); median lobe of aedeagus as
ilbuserated (Pie /8:7)5 female Ua HOW i aniston ae oSbonlatsdclsatann ated Seiisanignatet
Dra ded an rod Bak Atheta (Microdota) macesi Klimaszewski & Webster, sp. n.
Body dark brown, moderately glossy, meshed microsculpture present but not
strongly pronounced, punctation and pubescence dense (Figs 1, 33): male
reteite-V Il andicenialis, ditterently shapeds tas ceri ote Masccvstseeocys pecvvmsccks 12
Pubescence on elytra forming wavy pattern (Fig. 33); median lobe of aedea-
gus with sinuate venter of tubus in lateral view (Fig. 36), and internal sac with
two apical lobes in dorsal view (Fig. 37); spermatheca with tubular capsule,
deep and narrow invagination and S-shaped stem (Fig. 40)... eee eens
Sindh Se ween en swe LON imate Wn Atheta (Microdota) platonoffi Brundin
Pubescence on elytra directed obliquely posteriad from midline of disc (Fig.
1); median lobe of aedeagus with large bulbus and moderately long tubus, its
ventral margin arcuate and narrowly elongate apically (Fig. 4); spermatheca
with long, sac-shaped, tubular capsule bearing shallow but broad apical in-
vagination, stem broad and club-shaped (Fig. 8) [adventive in Canada]........
este a tts Hate cate Btn ora ears Az 8 Ra AEE Atheta (Microdota) subtilis (Scriba)

Subtilis species group (new)

Species of this group are characterized by: elytra at suture at least as long as prono-
tum (Figs 1, 9), male tergite VIII truncate and sometimes slightly emarginated me-
dially (Figs 2, 10), median lobe of aedeagus with broadly oval bulbus streamlined
with broadly triangular tubus in dorsal view (Figs 5, 13), in lateral view tubus straight
medio-basally and narrowly elongate and strongly produced ventrally at apex (Figs 4,

12), internal sac structures complex with two prominent elongate structures in bulbus

(Figs 4, 5, 12, 13), spermatheca L-shaped with long, broad sac-shaped capsule bearing

wide and shallow apical invagination and with short and swollen apically stem (Figs 8,

16). Two species are known from eastern Canada.

eli) Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Atheta (Microdota) subtilis (Scriba)
Figs 1-8

Homalota subtilis Scriba 1866: 128. As Atheta (Microdota): Brundin 1948, Palm 1970,
Benick and Lohse 1974, Smetana 2004 (review of literature and description).

Material examined. Canada, Labrador, Goose Bay, Rts. 500 and 520 jct., 53°16.9
N, 60°24.6 W, 13-26.VHI.2001, S. and J. Peck, Flight Intercept Trap, elevation 10 m,
spruce-poplar forest (LFC) 3 females, 1 male; Goose Bay, Goose River Bridge, 53°22.2
N, 60°26.2 W, 15-20.VII.2001, S. and J. Peck, elevation 10 m, spruce-birch forest
(LFC) 1 male. New Brunswick, Albert Co., Caledonia Gorge P.N.A., 45.7941°N,
64.7736°W, 13.[X.2011, R.P. Webster // near Crooked Creek, mixed forest (red spruce
and yellow birch) in decaying gilled mushrooms (RWC) 1 male; Carleton Co., Wake-
field, Meduxnekeag Valley Nature Preserve, 46.1940°N, 67.6800°W, 3.VII.2006,
R.P. Webster coll. // mixed forest on Pleurotus sp. on dead standing Populus tremu-
loides (RWC) 1 male; York Co., New Maryland, Charters Settlement, 45.8331°N,
66.7410°W, 27.VII.2005, R.P Webster coll. // mixed forest on flowers of Spiraea
alba (LFC) 1 male; Restigouche Co., off Bellone Road, 47.7755°N, 68.2501°W,
24.VIII.2011, R. Webster and M. Turgeon // Old spruce and fir forest, mossy forest
floor, in gilled mushrooms in various stages of decay (RWC) 1 female.

Diagnosis. Body length 1.5-2.0 mm, subparallel, flattened, reddish brown to
dark brown, head and abdomen darker than pronotum and elytra in some specimens,
legs yellowish brown (Fig. 1); integument moderately glossy, densely punctate and
densely pubescent on forebody and less so on abdomen, microsculpture fine; head
slightly narrower than pronotum, strongly narrowed posteriad, eyes large and about
as long as postocular area dorsally; pronotum transverse, narrower than elytra; elytra
wider and longer than pronotum; abdomen subparallel. MALE. Tergite VHI truncate
apically (Fig. 2); sternite VII broadly rounded apically (Fig. 3); median lobe of aedea-
gus narrow, and strongly ventrally produced apically in lateral view (Fig. 4); internal
sac structures complex (Figs 4, 5). FEMALE. Tergite VIII broadly emarginated api-
cally (Fig. 6); sternite VIII slightly emarginated apically (Fig. 7); spermatheca L-shaped
with long, broad sac-shaped capsule bearing wide and shallow apical invagination and
club-shaped short and swollen apically stem (Fig. 8).

Natural history. The LB specimens were collected in flight intercept traps set in
spruce-poplar forest. The NB specimens were found in gilled mushrooms at various
stages of decay in spruce/fir forest, in Pleurotus sp. on dead standing P. tremuloides, in
mixed forest on flowers of S. alba, and in a mixed forest with red spruce and yellow
birch. Adults were captured from July to September.

Distribution. Atheta (M.) subtilis is a Palaearctic species (for details, see Brundin
1948, Palm 1970, Benick and Lohse 1974, Smetana 2004), and it is reported here as
adventive for the first time from Canada (LB, NB) and North America.

Comments. Adults of A. subtilis from LB were captured in association with A.
pseudosubtilis Klimaszewski and Langor. Some females of the former species, because

A review of Canadian and Alaskan species of the genera Clusiota Casey... 111

Atheta subtilis

Figures |-8. Azheta (Microdota) subtilis (Scriba): | habitus in dorsal view 2 male tergite VIUI3 male sternite
VII 4 median lobe of aedeagus in lateral view 5 median lobe of aedeagus in dorsal view 6 female tergite VIII

7 female sternite VII 8 spermatheca. Scale bar for habitus = 1 mm; remaining scale bars = 0.2 mm.

of similarly shaped spermatheca and poorly preserved body, were misidentified as the
latter species. We have compared European specimens of A. subtilis with those from
Canada (LB, NB) and found no significant differences in external morphology and
shape and structures of genitalia.

112 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Atheta (Microdota) pseudosubtilis Klimaszewski & Langor
Figs 9-16

(For diagnosis, see Klimaszewski et al. 2011)

Distribution. Recorded from NF and LB, NB (Klimaszewski et al. 2011, Webster et
al. 2012).

Comments. The taxonomic position of A. pseudosubtilis is somewhat unclear. The
shape of the spermatheca is very similar to those of A. subtilis and C. antennalis. Ex-
ternally it is similar to C. antennalis but does not have a swollen basal antennal article.
The median lobe of the aedeagus has internal sac structures very similar to those of
A, subtilis. Externally, A. pseudosubtilis is readily distinguished from A. subtilis by the
much shorter elytra (Figs 1, 9). DNA studies of all these species would be very useful
in revealing their true relationships.

Platonoffi species group (new)

Species of this group are characterized by elytra at suture ranging from shorter to
longer than pronotum (Figs 17, 25, 33, 41), male tergite VII truncate and sometimes
slightly emarginated medially or slightly crenulate apically (Figs 18, 26, 34, 42), me-
dian lobe of aedeagus with broadly oval bulbus clearly demarcated from triangular
tubus in dorsal view (Figs 20, 29, 37, 45), and in lateral view, tubus straight medio-
basally, arcuate or sinuate and moderately to strongly produced ventrally at apex (Figs
21, 28, 36, 44), internal sac structures complex (Figs 20, 21, 28, 29, 36, 37, 44, 45);
spermatheca S-shaped with long, elongate club-shaped capsule bearing narrow and
deep apical invagination and long sinuate stem (Figs 24, 32, 40, 48). Five species are
known from Canada and Alaska.

Atheta (Microdota) curtipenis Klimaszewski & Webster, sp. n.
http://zoobank.org/08F3959E-4933-47 1 F-B4FB-7D29A49665F5
Figs 17-24

Holotype (male). Canada, New Brunswick, Saint John Co., ca 2 km NE of Maces
Bay, 45.1168°N, 66.4552°W, 8.V.2006, R.P. Webster, coll. // eastern white cedar
swamp, under moose dung (LFC). Paratypes: labelled as the holotype (RWC) 1 male,
1 female.

Etymology. The specific name curtipenis refers to a short median lobe of aedeagus
of this species.

Diagnosis. Body length 2.0—2.3 mm, subparallel, moderately convex, dark brown,
abdomen slightly darker than remainder of the body, legs yellowish brown (Fig. 17);
integument glossy, densely punctate and densely pubescent on forebody and less so

A review of Canadian and Alaskan species of the genera Clusiota Casey... 113

Atheta pseudosubtilis

Figures 9-16. Atheta (Microdota) pseudosubtilis Klimaszewski & Langor: 9 habitus in dorsal view
10 male tergite VIII 11 male sternite VIII 12 median lobe of aedeagus in lateral view 13 median lobe
of aedeagus in dorsal view 14 female tergite VIII 15 female sternite VIII 16 spermatheca. Scale bar for

habitus = 1 mm; remaining scale bars = 0.2 mm.

on head and abdomen, microsculpture of forebody fine; head slightly narrower than
pronotum, strongly narrowed posteriad and slightly angular posteriorly, eyes large
and slightly shorter than postocular area dorsally; pronotum transverse, narrower than

114 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Atheta curtipenis

Figures | 7-24. Azheta (Microdota) curtipenis Klimaszewski & Webster, sp. n.: 17 habitus in dorsal view
18 male tergite VIII 19 male sternite VI 20 median lobe of aedeagus in dorsal view 21 median lobe
of aedeagus in lateral view 22 female tergite VUI 23 female sternite VIII 24 spermatheca. Scale bar for

habitus = 1 mm; remaining scale bars = 0.2 mm.

elytra; elytra wider and longer than pronotum; abdomen subparallel. MALE. Tergite
VII truncate apically (Fig. 18); sternite VIII broadly rounded apically (Fig. 19); me-
dian lobe of aedeagus narrowly oval in dorsal view (Fig. 20), tubus sinuate basally and

A review of Canadian and Alaskan species of the genera Clusiota Casey... Ts

then straight and rounded apically in lateral view (Fig. 21); internal sac structures as
illustrated (Figs 20, 21). FEMALE. Tergite VII truncate apically (Fig. 22); sternite
VIII slightly emarginated apically (Fig. 23); spermatheca compressed S-shaped, cap-
sule spherical with short and narrow apical invagination, stem narrow and U-formed
posteriorly (Fig. 24).

Natural history. Adults were found in an eastern white cedar swamp under moose
dung in May.

Distribution. Known only from NB, Canada.

Comments. This species is distinguished by the moderately transverse pronotum,
and the shape of the median lobe of the aedeagus and spermatheca.

Atheta (Microdota) microelytrata Klimaszewski & Godin
Figs 25-32

(For diagnosis, see Klimaszewski et al. 2012)

Distribution. Recorded only from YT (Klimaszewski et al. 2012).

Atheta (Microdota) platonoffi Brundin
Figs 33-40

(For diagnosis, see Klimaszewski et al. 2011)

Distribution. In Canada, recorded from AB, BC, LB, NB, NE NS, ON, SK, YT, and
in USA from AK (Klimaszewski et al. 2011, 2015).

Atheta (Microdota) pratensis (Maklin, 1852)

Homalota pratensis Maklin, 1852: 308. As Atheta (Microdota): Moore and Legner
1975, Klimaszewski et al. 2011; Bousquet et al. 2013.

Syntypes. USA, Alaska: Kenai; Holmberg; pratensis Maklin; Mus. Zool. Helsinki, No.
14517 (ZMH) 1 male; same labels except No. 17518 (ZMH) 1 female.

Diagnosis. This species may be readily separated from other Nearctic congen-
ers by the following combination of characters: pronotum rounded and margined, as
wide as head and at least 1.5 times narrower than elytra; elytra elongate about twice
as long as pronotum with wavy pattern of pubescence posteriorly; male tergite VI
truncate apically; sternite VII rounded apically; median lobe of aedeagus with large
oval bulbus and small triangular tubus in dorsal view, apical part of tubus narrow and

116 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

32b 32a
Atheta microelytrata

Figures 25-32. Atheta (Microdota) microelytrata Klimaszewski & Godin: 25 habitus in dorsal view
26 male tergite VIII 27 male sternite VIII 28 median lobe of aedeagus in lateral view 29 median lobe of
aedeagus in dorsal view 30 female tergite VII 31 female sternite VIH 32a, b spermatheca. Scale bar for

habitus = 1 mm; remaining scale bars = 0.2 mm.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 117

Atheta platonoffi

Figures 33-40. Atheta (Microdota) platonoffi Bernhauer: 33 habitus in dorsal view 34 male tergite VIII
35 male sternite VII] 36 median lobe of aedeagus in lateral view 37 median lobe of aedeagus in dorsal
view 38 female tergite VIII 39 female sternite VIII 40 spermatheca. Scale bar for habitus = 1 mm; remain-

ing scale bars = 0.2 mm.

118 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

produced ventrally in lateral view; female tergite VIII truncate apically, and sternite
VIII rounded apically and with antecostal suture sinuate and pointed medially; sperm-
atheca S-shaped with club-shaped capsule bearing deep invagination and sinuate stem
looped posteriorly, similar to that of A. platonoffi.

Distribution. AK, WA (Maklin 1852, Moore and Legner 1975).

Comments. This species is somewhat similar to A. subtilis but may readily be dis-
tinguished externally by having elytra about 1.5 times wider and almost twice longer
than pronotum. We examined the type series but the specimens were in poor shape
and therefore were not illustrated.

Atheta (Microdota) riparia Klimaszewski & Godin
Figs 41-48

(For diagnosis, see Klimaszewski et al. 2012)
Atheta (Microdota) riparia Klimaszewski & Godin, 2012: 225.

Distribution. Recorded only from YT in Canada (Klimaszewski et al. 2012).

Pennsylvanica species group (new)

Species of this group are characterized by elytra at suture ranging from as long as or
longer than pronotum (Figs 49, 56, 61, 69), male tergite VUI truncate or slightly
emarginated medially and slightly crenulate apically (Figs 50, 70), median lobe of
aedeagus with broadly oval bulbus streamlined with broad basally triangular tubus in
dorsal view (Figs 52, 65, 74, 75), in lateral view tubus straight and slightly narrowly
triangularly produced ventrally (Figs 51, 64, 72, 73), internal sac structures complex
(Figs 51, 52, 64, 65, 72-75); spermatheca S-shaped with club-shaped capsule bearing
narrow and shallow apical invagination and long, posteriorly looped stem (Figs 55, 59,

60, 68, 78, 79). Four species are known from Canada.

Atheta (Microdota) amicula (Stephens)
Figs 49-55

(For diagnosis, see Klimaszewski et al. 2011, Lee and Ahn 2015)
Aleochara amicula Stephens, 1832: 132. As Atheta (Microdota): Brundin 1948, Palm
1970, Benick and Lohse 1974, Smetana 2004, Lee and Ahn 2015.

Distribution. Atheta (M.) amicula is a Palaearctic species adventive in North America.
It was reported in Canada based only on female specimens from NF and NS (Majka

A review of Canadian and Alaskan species of the genera Clusiota Casey... 119

Atheta riparia

Figures 41-48. Atheta (Microdota) riparia Klimaszewski & Godin: 41 habitus in dorsal view 42 male
tergite VIII 43 male sternite VIII 44 median lobe of aedeagus in lateral view 45 median lobe of aedeagus
in dorsal view 46 female tergite VIII 47 female sternite VIII 48 spermatheca. Scale bar for habitus = 1

mm; remaining scale bars = 0.2 mm.

and Klimaszewski 2008, Klimaszewski et al. 2011). In USA, it was recorded from WA
(Moore and Legner 1975). For Palaearctic distribution and synonymy of this species, see
Lee and Ahn (2015).

120 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Atheta amicula

Figures 49-55. Azheta (Microdota) amicula (Stephens): 49 habitus in dorsal view 50 male tergite VIII
(based on European specimen) 51 median lobe of aedeagus in lateral view (after Brundin 1948) 52 me-
dian lobe of aedeagus in dorsal view (after Brundin 1948) 53 female tergite VIII 54 female sternite VII
55 spermatheca. Scale bar for habitus = 1 mm; remaining scale bars = 0.2 mm.

Atheta (Microdota) festinans (Erichson)
Figs 56-60

(For diagnosis and synonymy, see Gusarov 2003b)
Homalota festinans Erichson, 1839: 112. As Atheta (Microdota): Gusarov 2003a, b,
Brunke et al. 2012.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 121

56

Atheta festinans

Figures 56-60. Atheta (Microdota) festinans (Erichson): 56 habitus in dorsal view 57 female tergite VIII

58 female sternite VIII 59, 60 spermatheca. Scale bar for habitus = 1 mm; remaining scale bars = 0.2 mm.

Material examined. Canada, Quebec, Berthier Co., Berthierville, 20.X1.2004, Michel
Racine coll., sous débris de bois, dans sabliére, avec Carabe Dyschiriodes sp. (LFC) 1 female.
Natural history. ‘This is the first record with habitat data for this species. The QC

specimen was captured from woody debris in a sandy pit in association with Dyschirius

sp. (Carabidae).

122 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Distribution. Erichson (1839) described this species from PA in USA. Gusarov
(2003b) recorded it from AZ, CT, IA, IN, KY, NY, PA, and RI. Bernhauer (1907)
reported A. festinans from ON, and Brunke et al. (2012) confirmed occurrence of
this species in Waterloo Reg., ON. Here, we provide the first record of this species
from QC.

Comments. All known Canadian specimens of Atheta festinans are females. Gusa-
rov (2003b) remarked that all specimens seen of this species were females and sug-
gested that this species may be parthenogenetic.

Atheta (Microdota) formicaensis Klimaszewski & Webster, sp. n.
http://zoobank.org/A4C2D2A2-735F-4D0B-94B0-D46C3D2BDB8C
Figs 61-68

Holotype (male). Canada, New Brunswick, York Co., New Maryland, Charters Set-
tlement, 45.8395°N, 66.7391°W, 19.V.2006, R.P Webster coll. // mixed forest, on
surface of nest of black Formica sp. (LFC) 1 male. Paratypes: labelled as holotype
(RWC) 1 male, 1 female; Charters Settlement, 45.8395°N, 66.7391°W, 29.IV 2004,
R.P. Webster coll. // mixed forest, on surface of nest of black Formica sp. (RWC) 3 sex
undetermined; same data except: 30.IV.2005 // mixed forest in nest of black Formica
sp., sifting nest material (RWC) 2 sex undetermined; Queens Co., Cranberry Lake
PN.A., 46.1125°N, 65.6075°W, 13.V.2011, R.P Webster coll. // old red oak forest, in
nest of black mound-building Formica species, near surface of mound (LFC, RWC) 1
female, 2 sex undetermined.

Etymology. The specific name formicaensis is a feminine adjective derived from
the generic name Formica, an ant genus found in association with the type series.

Diagnosis. Body length 2.6—2.8 mm, subparallel, moderately convex, head and
posterior part of abdomen dark brown, pronotum medium to dark brown, elytra with
centre of disc yellowish brown and darker edges, base of abdomen light brown, legs yel-
lowish brown (Fig. 61); integument glossy, sparsely punctate and sparsely pubescent,
microsculpture distinct and stronger on pronotum and elytra; head slightly narrower
than pronotum, rounded and slightly angular posteriorly, eyes small and shorter than
postocular area dorsally; antennal articles V—X from subquadrate to slightly transverse;
pronotum transverse, slightly narrower than elytra; elytra wider and as long as prono-
tum; abdomen subparallel. MALE. Tergite VIII truncate apically (Fig. 62); sternite
VIII broadly rounded apically and slightly pointed medially (Fig. 63); median lobe of
aedeagus narrowly oval in dorsal view with short and triangular tubus (Fig. 65), in lat-
eral view tubus sinuate basally and then straight and rounded apically (Fig. 64); internal
sac structures as illustrated (Figs 64, 65). FEMALE. Tergite VIII truncate apically (Fig.
66); sternite VII broadly rounded apically (Fig. 67); spermatheca small, S-shaped, cap-
sule spherical without apparent apical invagination, stem narrow and sinuate (Fig. 68).

Natural history. Adults were found in association with nests of black ants in the
genus Formica in April and May.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 123

67

Atheta formicaensis

Figures 61-68. Azheta (Microdota) formicaensis Klimaszewski & Webster, sp. n.: 61 habitus in dorsal
view 62 male tergite VIII 63 male sternite VIII 64 median lobe of aedeagus in lateral view 65 median
lobe of aedeagus in dorsal view 66 female tergite VIII 67 female sternite VIII 68 spermatheca. Scale bar

for habitus = 1 mm; remaining scale bars = 0.2 mm.

124 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Distribution. Known only from NB, Canada.

Comments. This species is probably closely associated with nests of the ant genus
Formica. It is distinguished from all other Nearctic species of Microdota by the shape
of the median lobe of the aedeagus and spermatheca. ‘The shape of the spermatheca is
similar to that of Palaearctic A. (.) glabricula Thomson (Palm 1970).

Atheta (Microdota) pennsylvanica Bernhauer
Figs 69-79

(For diagnosis, see Klimaszewski et al. 2011, and for synonymy, Gusarov 2003b)
Bernhauer 1907: 388. As Atheta (Microdota): Gusarov 2003b.

Distribution. This species was recorded in Canada from NB, NE, NS, ON, QC, and
in the USA from MN, NY, PA, VT (Gusarov 2003b, Klimaszewski et al. 2011).

Sculptisoma species group (new)

Species of this group are characterized by elytra at suture at least as long as pronotum
(Fig. 80), male unknown; spermatheca pipe-shaped with hemispherical capsule nar-
rowed basally and without apparent apical invagination, and with long stem that is
looped posteriorly and twisted apically (Fig. 83). One species belongs to this group.

Atheta (Microdota) sculptisoma Klimaszewski & Langor
Figs 80-83

(For diagnosis, see Klimaszewski et al. 2011)
Klimaszewski et al. 2011: 148.

Distribution. This native Nearctic species was recorded only from the type locality in
southeastern NF in Canada (Klimaszewski et al. 2011).

Macesi species group (new)

Species of this group are characterized by the strongly glossy body, elytra at suture
slightly longer than pronotum (Fig. 84), male tergite VIII truncate apically and with
two small lateral teeth (Fig. 85), median lobe of aedeagus with small bulbus and elon-
gate tubus, in lateral view tubus straight, apex narrowly triangular and slightly pointed
(Fig. 87), internal sac structures pronounced (Fig. 87); female unknown. One species
belongs to this group.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 125

Atheta pennsylvanica

Figures 69-79. Atheta (Microdota) pennsylvanica Bernhauer: 69 habitus in dorsal view 70 male tergite
VII (after Gusarov 2003b) 71 male sternite VIII (after Gusarov 2003b) 72 median lobe of aedeagus in
lateral view (after Gusarov 2003b), and 73 based on Canadian specimen 74 median lobe of aedeagus
in ventral view (after Gusarov 2003b), and 75 in dorsal view (based on Canadian specimen) 76 female
tergite VIII (after Gusarov 2003b) 77 female sternite VIII (after Gusarov 2003b) 78 spermatheca (after
Gusarov 2003b), and 79 based on Canadian specimen. Scale bar for habitus = 1 mm; remaining scale

bars = 0.2 mm.

126 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Atheta sculptisoma

Figures 80-83. Atheta (Microdota) sculptisoma Klimaszewski & Langor: 80 habitus in dorsal view 81 fe-
male tergite VII 82 female sternite VIII 83 spermatheca. Scale bar for habitus = 1 mm; remaining scale

bars = 0.2 mm.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 127

Atheta (Microdota) macesi Klimaszewski & Webster, sp. n.
http://zoobank.org/A4599D2A-246D-4AD 1-A2F4-214C4585DED9
Figs 84-87

Holotype (male). Canada, New Brunswick, Saint John Co., ca 2 km NE of Maces
Bay, 45.1161 N, 66.4560 W, 8.V.2006, R.P. Webster, coll. // Eastern white cedar
swamp, in sphagnum and litter near brook (LFC).

Etymology. The specific name macesi is an adjective derived from Maces Bay in
NB, where the holotype specimen was found.

Diagnosis. Body length 2.7 mm, subparallel, flattened, brownish-black, tibiae and
tarsi brown (Fig. 84); integument glossy and more so on abdomen, sparsely punctate
and pubescent, except for pronotum and elytra; microsculpture of forebody dense and
strong, meshed with hexagonal sculpticells; head about as wide as pronotum, slightly
angular posteriorly, eyes large and as long as postocular area dorsally; antennae with
articles V-X moderately to strongly transverse; pronotum broadest in apical third and
narrowest at base, rounded laterally and basally, transverse, narrower than elytra; elytra
wider and slightly longer than pronotum; abdomen subparallel. MALE. Tergite VII
truncate apically and with two large lateral teeth (Fig. 85); sternite VII rounded api-
cally (Fig. 86); median lobe of aedeagus with small bulbus and long tubus, in lateral
view tubus straight and apex slightly produced ventrally, apex narrowly triangular and
slightly pointed (Fig. 87), internal sac structures well defined (Fig. 87). FEMALE.
Unknown.

Natural history. A single male was found in eastern white cedar in sphagnum and
litter near a brook, in May.

Distribution. Known only from NB, Canada.

Comments. ‘This species is known only from a single male collected in sphagnum
and litter.

Genus Clusiota Casey

Clusiota Casey, 1910: 119; Moore and Legner 1975: 347.

Diagnosis. The following combination of characters is distinctive for Clusiota: small
and subparallel body (Figs 96, 88, 104), length 1.5-2.5 mm, antennomere I swollen
(Figs 96, 88) except for C. grandipenis (Fig. 104), and longer than H, V-X strongly
transverse (Figs 96, 88, 104); labial palps with 3 articles; glossae narrow, deeply split
forming V-shaped structure; maxillary palpus with 3 articles, last one narrowly elon-
gate; pronotum transverse, about 1.2 times as wide as long, pubescence at midline
directed apically in most specimens anteriorly and laterad elsewhere (Figs 96, 88, 104);
pronotal hypomeron fully visible medially in lateral view; elytra with pubescence di-
rected obliquely postero-laterad from midline of disc (Figs 96, 88, 104); abdomen
slightly swollen posteriorly; male tergite VIII emarginate medially (Figs 89, 97, 105);

128 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

86

Atheta macesi

Figures 84-87. Atheta (Microdota) macesi Klimaszewski & Webster, sp. n.: 84 habitus in dorsal view
85 male tergite VII 86 male sternite VIII 87 median lobe of aedeagus in lateral view. Scale bar for habitus

= 1 mm; remaining scale bars = 0.2 mm.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 129)

median lobe of aedeagus with large bulbus and moderately narrow, and triangularly
shaped apically tubus in dorsal view (Figs 91, 99, 107), crista apicalis of bulbus large
(Figs 92, 100, 108); spermatheca L-shaped or S-shaped with club-shaped tubular cap-
sule, and short sinuate stem (Figs 95, 103, 111).

Comments. Species of this genus may be confused with Microdota species, from
which they may be readily distinguished by the swollen first basal antennal articles
(except for C. grandipenis), and shape of genitalia, with median lobe bearing large
crista apicalis of bulbus. Casey (1910) believed this genus was related to the subgenus
Datomicra of Atheta.

Key to Nearctic species of Clusiota

ib Elytra longer than pronotum and at least 1.3 times broader than pronotum
ig SeS LOA YK cM dk. ce cten teat calcite Muh ita shel pts ath, Pus etoatnis nha ea esd, odd SoS Melt 2
- Elytra at most as long as pronotum and about 1.1 times broader than prono-
tum (Fig. 96); median lobe of aedeagus with tubus straight medially and api-
cal part broad in lateral view (Fig. 100), bulbus without distinct large sclerites
in dorsal view (Fig. 99); male tergite VHI with shallowly emarginated apical
margin (Fig. 97); spermatheca consisting of club-shaped and elongate capsule
and sinuate, short ‘and broad:stenn (Pig 4V03 i. <x.ncoteet ature aueboesneat beceeh cate
ier, Mee oanitiasiete to gon tecesneet ieee Clusiota antennalis Klimaszewski & Godin
De Body reddish-brown (Fig. 88); median lobe of aedeagus with tubus sinuate
medially and apical part narrow in lateral view (Fig. 92), bulbus with two
strong elongate sclerites in dorsal view (Fig. 91); male tergite VIII with deeply
emarginated apical margin (Fig. 89); spermatheca sinuate (Fig. 95)..........0..
sariga var eaknnacatie fod eaten Men tat eek cae ance oaees Clusiota impressicollis (Bernhauer)
= Body dark brown (Fig. 104), median lobe of aedeagus with tubus scarcely
sinuate medially and apical part broad in lateral view (Fig. 108), bulbus with
differently shaped structures (Fig. 107); spermatheca S-shaped (Fig. 111) ....
Er es Resnet diese Anau Clusiota grandipenis Klimaszewski & Webster, sp. n.

Clusiota impressicollis (Bernhauer)

Figs 88-95

(For diagnosis, see Casey 1910, Klimaszewski et al. 2011)

Atheta impressicollis Bernhauer 1907: 389. As Clusiota: Klimaszewski et al. 2011, Brunke
et al. 2012, Bousquet et al. 2013.

Clusiota claviventris Casey 1910: 119. Synonymized by Gusarov 2003a.

Material examined (additional locality data). Canada, Ontario, Sudbury Co.,
Mattagami, 25.VHI.1980, R. Baranowski (ZML) 2 males, 1 female; Nipissing Co.,
Algonquin Provincial Park, near Brent, 21.VIII.1980, R. Baranowski (ZML) 1 male.

130 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

Clusiota impressicollis

Figures 88-95. Clusiota impressicollis (Bernhauer): 88 habitus in dorsal view 89 male tergite VIII
90 male sternite VIII 91 median lobe of aedeagus in dorsal view 92 median lobe of aedeagus in lateral
view 93 female tergite VIII 94 female sternite VIII 95 spermatheca. Scale bar for habitus = 1 mm; remain-
ing scale bars = 0.2 mm.

Distribution. BC, NB, NF, ON (Casey 1910, Klimaszewski et al. 2011, Bousquet
et al. 2013).

Natural history. The specimens from ON were collected in August. The NF speci-
mens were captured in a light flight intercept trap in fir-deciduous forest in July/August
(Klimaszewski et al. 2011). These records indicate that adults occur late in the season.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 131

Comments. This species was originally reported by Bernhauer (1907) from Bar-
ing, WA and Pasadena, CA. ‘The previously unknown female is illustrated here for the
first time from a specimen from Mattagami, ON.

Clusiota antennalis Klimaszewski & Godin
Figs 96-103

(For diagnosis, see Klimaszewski et al. 2008)
Clusiota antennalis K\limaszewski & Godin in Klimaszewski et al. 2008.

Material examined. Canada, British Columbia, Cooper River Valley, A31698/F4-1-
1, 4.VU-7.VII.1996, +/- 20 m. pitfall trap, J. Lemieux (LFC) 1 female, 1 sex? same
data except: A37541/P2-1-5, 6.VI-4.VII (LFC) 2 females; A36435/04-1-1, 7.VI-6.
VII.1996 (LFC) 2 males; Vancouver Island, Mt. Cain: 50.14°N, 126.21°W, 5.VI-27.
VI1.1996, 16.6 PIT2, N. Winchester (ZML) 2 females; 50.14°N, 126.21°W, 27, VI-13.
VII.1966, 16.6.PIT 3, N. Winchester (ZML) 3 females; 50.13°N, 126.21°W, 23.VI-
7.VII.1997, 16.6 PIT2, N. Winchester (ZML) 18 females; 50.13°N, 126.21°W,
7.VII—20.VII.1997, 16.6 PIT8, N. Winchester (ZML) 7 females; 50.13°N, 126.21°W,
20.VII—5.VIII.1997, 16.6 PIT 6, N. Winchester (ZML) 1 female; 50.13°N, 126.21°W,
+/- 20 m, 1.IX—19.IX.1997, 16.6 PIT8, N. Winchester (ZML) 5 males; same data ex-
cept: 15.[X—28.IX.1996, 17.4 PAN 3 (ZML) 1 female; 28.[X-12.X.1996, 16.6 PAN
1 (ZML) 3 females; 12.X—1.X1.1996, 16.6 PAN3 (ZML) 1 male; 50.15°N, 126.25°W,
19.IX—4.X.1997, 17.4 PIT 7 (ZML) 1 male, 1 female. U.S.A., Alaska, Prince of Wales
Is.: Staney Ck., 41-45 m el., 55.79901°N, 133.11782°W, old growth, pitfall 2, 25.VI—
9. VII.2012, J. Stockbridge et al. UAM100340147 (UAM) 1 female; same data except:
11—25.V1.2012, VAM100338700 (UAM) 1 female; 14—28.V.2012 UAM100338413,
UAM 100338412, UAM100338414 (UAM) 3 females; Luck Lk. 2 Rd., old growth,
105 m el., 55.96855°N, 132.79615°W, +/- 10 m pitfall 3, 29.VI-8.VII.2010, J.
Stockbridge UAM100278064 (UAM) 1 female; Luck Lk. 1 Rd., old growth, 101
m el., 55.97805°N, 132.75456°W, +/- 10 m. pitfall, 8-30.VII.2010, J. Stockbridge,
C. Bickford UAM100279634, UAM100279635 (UAM) 2 females; same data ex-
cept: 27.VI-11.VII.2012 UAM100343219, UAM100343218, UAM100343246
(UAM) 3 females; pitfall 1, 13-27.VI.2012, J. Stockbridge et al. VAM100339932
(UAM) 1 female; Hatchery Ck. 1, 2"! growth, 49 m el., 55.92654°N, 132.95645°N,
+/- 10 m pitfall 3, 18.V—4.VI.2010, J. Stockbridge, C. Bickford UAM100262426
(UAM) 1 female; Dall Is., p. 12 el. 688 m., 54.99342°N, 133.01688°N, +/-4 m krum-
mholtz, 7suga mertensiana, Vaccinium ovalifolium, pitfall, 15—-16.VH.2011, D.S. Sikes
UAM 100329960, UAM100329969 (UAM) 2 females. UAM data can be downloaded
at http://arctos.database.museum/saved/Clusiota_antennalis.

Distribution. Originally described from Dyea, AK (Klimaszewski et al. 2008). It
is here newly reported in Canada from BC, and from new localities in AK. Clusiota
antennalis is a western Nearctic species known only from western BC and AK.

132 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

i nN
MS "
\

Clusiota antennalis

96

Figures 96-103. Clusiota antennalis (Klimaszewski & Langor): 96 habitus in dorsal view 97 male tergite
VIII 98 male sternite VIII 99 median lobe of aedeagus in dorsal view 100 median lobe of aedeagus in lat-
eral view 101 female tergite VII 102 female sternite VIII 103 spermatheca. Scale bar for habitus = 1 mm;
remaining scale bars = 0.2 mm.

Natural history. In AK, the holotype was captured in alder litter, and the recent
AK specimens were collected primarily in pitfall traps from old-growth Pacific rain
forests at low elevations although two specimens were collected in an alpine habitat

A review of Canadian and Alaskan species of the genera Clusiota Casey... 133

above 650 m elevation. In BC, specimens from Vancouver Island were captured con-
tinuously from June to November in pitfall and pan traps.

Comments. Females of this species have a spermatheca extremely similar to those
of Atheta (M.) subtilis and A. (M.) pseudosubtilis. Clusiota antennalis is easily separated
from the two species by the swollen basal antennal article (Fig. 96), and from A. subtilis
by much shorter elytra (Fig. 96). It differs from all Nearctic Clusiota and Microdota
species by the shape of the genitalia.

Clusiota grandipenis Klimaszewski & Webster, sp. n.
http://zoobank.org/0424783F-C865-4166-A976-1 D60D3AA10AF
Figs 104-111

Holotype (male). Canada, New Brunswick, Westmorland Co., Sackville near
Ogden Mill, 45.92155°N, 64.38925°W, 12.V.2006, Scott Makepeace coll. // Black
spruce forest, in nest contents of Great Horned Owl — Bubo virginensis (LFC). Para-
type: Canada, New Brunswick, Northumberland Co., ca. 2.5 km W of Sevogle,
47.0876N, 65.8613W, 27.VIII.2013, old jack pine forest, in decaying gilled mush-
room, R.P. Webster (RWC) 1 female.

Etymology. ‘The specific name grandipenis, meaning large penis, refers to the large
tubus of the median lobe of the aedeagus of this species.

Diagnosis. Body length 2.2 mm, subparallel, flattened, dark brown, abdo-
men slightly darker than remainder of the body, legs yellowish brown (Fig. 104);
integument glossy, densely punctate and densely pubescent on forebody and less
so on head and particularly on abdomen, microsculpture of forebody fine, meshed
with hexagonal sculpticells; head about as wide as pronotum, slightly angular pos-
teriorly, eyes large and as long as postocular area dorsally; pronotum rounded
laterally and basally, transverse, narrower than elytra; elytra wider and longer than
pronotum; abdomen subparallel. MALE. Tergite VIII slightly emarginate apical-
ly (Fig. 105); sternite VHI broadly rounded apically (Fig. 106); median lobe of
aedeagus with broadly oval bulbus streamlined with apically narrowly triangular
tubus in dorsal view (Fig. 107), in lateral view tubus strongly produced ventrally,
apex narrowly triangular and slightly pointed (Fig. 108), internal sac structures
pronounced (Figs 107, 108). FEMALE. Tergite VIII with shallow apical median
emargination (Fig. 109); sternite VIII rounded apically (Fig. 110); spermatheca S-
shaped, capsule broadly club-shaped with deep median invagination, stem sinuate
with posterior loop (Fig. 111).

Natural history. One adult was found in the nest contents of a Great Horned
Owl, — Bubo virginensis in a black spruce forest in May and another from a decaying
gilled mushroom in a jack pine forest during August.

Distribution. Known only from NB, Canada.

134 Jan Klimaszewski et al. / ZooKeys 524: 103-136 (2015)

104
Clusiota grandipenis

Figures 104-I 11. Clusiota grandipenis Klimaszewski & Webster, sp. n.: 104 habitus in dorsal view
105 male tergite VII 106 male sternite VHI 107 median lobe of aedeagus in dorsal view 108 median
lobe of aedeagus in lateral view 109 female tergite VIII 110 female sternite VIII 111 spermatheca. Scale

bar for habitus = 1 mm; remaining scale bars = 0.2 mm.

A review of Canadian and Alaskan species of the genera Clusiota Casey... 135

Acknowledgments

We thank J. Frisch (ZMB), C. Fagerstr6m (ZML) for providing some specimens for
this study, and P. Fortin for providing eight images (Figs 61-68). We appreciate the
editorial assistance of P. Cheers (LFC) and the text formatting by D. Paquet (LFC).
We thank Scott Makepeace for collecting the nest contents from owl nests in New
Brunswick, and field technicians S. Meierotto and I. MacDougall. We are obliged to R.
Pace (Verona, Italy) for some taxonomic advice. This research was funded by Natural
Resources Canada and the Alaska Department of Fish and Game.

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