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M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger
(arthropodes excl. insectes).

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REVUE SUISSE DE ZOOLOGIE

TOME 118— FASCICULE 3

Publication subventionnée par:
ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT)
VILLE DE GENÈVE
SOCIÉTÉ SUISSE DE ZOOLOGIE

Comité de rédaction

DANIELLE DECROUEZ
Directrice du Muséum d'histoire naturelle de Genève

ALICE CIBOIS, PETER SCHUCHERT
Chargés de recherche au Muséum d'histoire naturelle de Genève

Comité de lecture

A. Cibois (oiseaux), G. Cuccodoro (coléoptères), S. Fisch-Muller (poissons),
B. Merz (insectes, excl. coléoptères), J. Mariaux (invertébrés excl. arthropodes),
M. Ruedi (mammifères), A. Schmitz (amphibiens, reptiles), P. Schwendinger
(arthropodes excl. insectes).

Le comité soumet chaque manuscrit pour évaluation à des experts d’institutions
suisses ou étrangères selon le sujet étudié.

La préférence sera donnée aux travaux concernant les domaines suivants: taxonomie,
systématique, faunistique, phylogénie, évolution, morphologie et anatomie comparée.

Administration

MUSÉUM D'HISTOIRE NATURELLE
1211 GENÈVE 6

Internet: http:/www.ville-ge.ch/musinfo/mhng/page/rsz.htm

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REVUE SUISSE DE ZOOLOGIE 118 (3): 401-412; septembre 2011

Microphorella cassari sp. n., a new species of Microphorella Becker
(Diptera: Dolichopodidae) from Tunisia

Paul GATT

Research Associate, Institute of Earth Systems, Chemistry Building 314 Floor,
University of Malta, Msida MSD 2080, Malta.

E-mail: paulgatt@onvol.net

Microphorella cassari Sp. nov., a new species of Microphorella Becker
(Diptera: Dolichopodidae) from Tunisia. - A new species of Micro -
phorella Becker, Microphorella cassari Sp. n., is described from Tunisia.
The strikingly long, strap-like, lanceolate antennal stylus and the long,
spine-like setae on the mid-portion of the costal vein of the male are unique
amongst previously described species of Microphorella. Likewise, the
completely divided sternite 10 of the female has not hitherto been recorded
in other species of this genus.

Keywords: Diptera - Dolichopodidae - Parathalassiinae - new species -
Mediterranean.

INTRODUCTION

Microphorella Becker is one of a small group of genera, both fossil and extant,
which are currently placed in the subfamily Parathalassiinae of the Dolichopodidae
s. lato (Ulrich, 2003; Sinclair & Cumming, 2006). This almost cosmopolitan genus
presently comprises 16 species of which 5 are Nearctic (Melander, 1928), 5 Palaearctic
(Chväla, 1988; Gatt, 2003; Shamshev, 2004), 5 Oriental (Shamshev & Grootaert, 2004)
and 1 Australian (Colless, 1963). Three species - M. curtipes (Becker, 1910) (Corsica
& Sardinia), M. ulrichi Gatt, 2003 (Tunisia & Morocco) and M. merzi Gatt, 2003
(Malta, Cyprus & Turkey) - are currently known from the Mediterranean, the latter two
exclusively so. Undescribed species are known from the Mediterranean (Gatt, in prep.)
and it appears that this subregion is more diverse in species than previously thought.

Species of Microphorella are minute, cryptic flies with inconspicuous habits
and which blend perfectly well with their surroundings (sandy beaches, gravel and
sand in river beds, and moist rocks in streams). They are therefore not often collected,
and very rare in collections.

MATERIAL AND METHODS

AII specimens were collected by the author from coastal sand dunes or estuaries
in two different localities in the Tabarka embayment (northwest coast of Tunisia) in the
spring of 2005 and 2007. Most specimens were collected by lying prone on the sand
and carefully aspirating the minute flies as they appeared. Some were collected by
sweeping very close to the sand using slow, deliberate movements.

Manuscript accepted 29.11.2010

402 P. GATT

Dissected specimens on which the drawings are based are stored in glycerine
microvials mounted on pins in the author’s collection. Temporary slide mounts of in-
sect parts were prepared in Berlese fluid, as described by Disney (1983). Drawings
were made with the aid of a x250 stereomicroscope and drawing tube. For some figures
(Figs 6-8, 13) a compound microscope with built in ocular grid was employed.
Drawings of complex structures made from slide mounts studied in transmitted light
(eg. Fig. 5) fail to make a clear distinction between internal and external structures, and
cannot show how the various parts are interconnected.

In descriptions of the abdomen and hypopygium, the terms “dorsal” and
“ventral” refer to the morphological position after genital rotation and flexion, 1.e. as
they appear in the intact specimen.

The following abbreviations are used in the text figures 1-13:

as accessory sclerites of genital fork (sternite 9)
& female cercus

Ce male right cercus

ds dorsal sclerite of genital fork

EA ejaculatory apodeme

EPA epandrium

fo foramen from segment 8

HA hypandrium

hs10 female abdominal hemisternite 10
hti0 female abdominal hemitergite 10
PH phallus

S spermatheca

si-s8 abdominal sternites 1-8

ti-t8 abdominal tergites 1-8

vs ventral sclerite of genital fork (sternite 9)

The holotype labels are cited verbatim. The text of each separate label is
enclosed in quotation marks, whilst individual lines on each label are separated by
slash lines.

Specimens are deposited as indicated by the following acronyms in brackets
under ‘material examined”:

IRSNB Institut Royal des Sciences Naturelles, Bruxelles, Belgium

MHNG Museum d'histoire naturelle, Genève

NMWC National Museum Wales, Cardiff

PGS Private collection of P. Gatt, Sliema, Malta

ZIN Zoological Institute of the Russian Academy of Sciences, St. Petersburg

ZMHB Museum für Naturkunde, Berlin (Zoologisches Museum der Humboldt
Universität Berlin)

The terminology used in this account follows Merz & Haenni (2000).
Homologies for the male terminalia follows Sinclair (2000).

A NEW MICROPHORELLA FROM TUNISIA 403

TAXONOMIC TREATMENT

Microphorella cassari sp. n. Figs 1-13

MATERIAL EXAMINED (13 GG 14 99: Holotype, male: “TUNISIA: Tabarka, / Oued
Berkoukech, / dunes, 31.11.2007 / P. Gatt”, “HOLOTYPUS / Microphorella / cassari sp. n. /
Gatt.” (IRSNB). The holotype is preserved in alcohol and is in perfect condition. — Paratypes, 9
G G and 12 © ® same data as holotype: 1 © Tunisia, Tabarka: Oued Berkoukech, beach, 13.iv.
2005, P. Gatt; 2 G à , Tunisia, Tabarka, Oued Bouterfess, beach, estuary, 14.iv. 2005, P. Gatt; 1
& and 1 © , Tunisia, Tabarka, Oued Bouterfess, dunes, 31.11.2007, P. Gatt (IRSNB, MHNG.,
NMWC, PGS, ZIN, ZMHB). Paratypes are preserved in alcohol or dry (double mounted on pins)
and are in good to excellent condition. — Nontype material: 2 & 4 and 1 9 , same data as holo-
type; 1 ©, Tunisia, Tabarka, Oued Berkoukech, estuary, 13.iv.2005, P. Gatt (PGS).

ETYMOLOGY: This species 1s dedicated to my dear friend and colleague Dr.
Louis F. Cassar who introduced me to the type locality, and in the company of whom

I have spent many happy hours of fieldwork in North Africa.

DIAGNOSIS: À small (1.3 mm), brownish-grey microtrichose, sexually dimorphic
species with white, lanceolate macrosetae and infuscated wings. Male: antennal style
long, strap-like, acuminate; fore and mid tibia, posteriorly, with long setae; mid femur
ventrally with a dense tuft of very short setulae and some long setae; mid basitarsus
ventrally with a long row of hook-like spinules; anterior costal margin with spine-like
setae. Female: tergite 10 and sternite 10 completely divided; tergite 10 bearing setae.

DESCRIPTION

Male

Length. Body 1.3 mm, wing 1.2 mm (specimen in alcohol).

Head (Fig. 1). Black in ground colour, brownish-grey microtrichose, higher and
wider than deep. Eyes widely separated on frons, ommatidia equal in size, sparsely
covered with minute ommatrichia. Neck inserted high on head. Occiput greyish-brown
microtrichose with some metallic reflections, scarcely projecting beyond posterior
margin of eye, concave above neck, convex below. Gena very narrow. Face silvery
grey microtrichose, long and narrow, narrower than frons above antennae, wWidening
below. Clypeus grey microtrichose, long and convex. Antenna (Fig. 2) black, brownish
microtrichose and clothed with short, pale setulae; placed at middle of head in profile,
and as long as head is deep; scape cup-shaped; pedicel globular, with a circlet of sub-
apical setulae; postpedicel long, bulbous in distal half and bearing a sensory pit at both
ends; stylus uniarticulate, long and strap-like, subequal in length to pedicel, widening
apically and terminating acutely; postpedicel and stylus covered with adpressed, spine-
like microtrichia. Proboscis short, lustrous brown, projecting downwards. Palpus
small, greyish microtrichose, with several long apical setulae and one longer seta on
lateral surface; sensory pit present. Chaetotaxy: cephalic setae white, thick and
flattened, somewhat lanceolate; one pair of strong, lateroclinate anterior ocellars; one
pair of weak, proclinate posterior ocellars; one pair of inclinate anterior fronto-orbitals;
one pair of lateroclinate posterior fronto-orbitals; one pair of long, inclinate medial
verticals; one pair of shorter, lateroclinate lateral verticals; postocular occipital setae
uniserial above neck, becoming longer and irregularly multiserial below; several
longer setae present just posterior to mouth opening, including postgena.

Thorax. Black in ground colour, generally grey microtrichose; scutum slightly
brown microtrichose anteriorly, With some faint metallic reflections; dorsal surface of

404 P. GATT

FIGs 1-3

Microphorella cassari sp. n.(1) 4 head, viewed from above and in front. (2) & antenna, lateral
view. (3) 4 , wing. Scale bars, Figs 1 and 3: 0.5 mm, Fig. 2: 0.2 mm.

mesoscutum moderately arched; prescutellar depression distinct; complete prothoracic
precoxal bridge present. Mesopleuron bare, greyish brown microtrichose. Chaetotaxy:
thoracic setae white, thick and flattened. Antepronotum with 4 setulae. Propleuron with

À NEW MICROPHORELLA FROM TUNISIA 405

ME au PE 180;

FIG. 4

Microphorella cassari sp. n., &, abdomen, ventral view. Scale bar 0.5 mm. Abbreviations
explained in the text.

one pair of setulae. Postpronotum with 1 setula. A single pair of long, acrostichal setae
on anterior slope of mesoscutum. Five (2+3) pairs of subequal dorsocentral setae,
1 prescutellar pair longer, wider apart, and as strong as 1 pair of long, inclinate scu -
tellar setae; dorsocentral setae sometimes preceded by 1-2 setulae; additional setulae
(2-3) sometimes present outside dorsocentral row on intra-alar line; 1 strong supra-alar
seta, preceded by a row of 2-3 setulae; 1 postalar seta; 2 notopleural setae.

Legs. Long and slender, black in ground colour, greyish microtrichose and
clothed with very short, white setulae; trochanters, apices of femora and all tibiae and
tarsi paler. Femora equally thick, not much stronger than tibiae. Basitarsi as long as
following tarsal segments combined. Coxae (especially fore pair) with some longer
setae. Hind trochanter with 1 long, ventral seta. AIT tarsomeres with short, spine-like

406 P. GATT

FIG. 5

Microphorella cassari sp. n., 8 abdomen, dorsal view. Scale bar 0.5 mm. Abbreviations
explained in the text.

subapical setae, stronger and in groups of 4 on middle segments. Tarsal claws, pulvilli
and empodium developed on all legs; pulvilli and empodium haired. Fore tibia with a
posterior row of 8-10 long posteriorly curved setae, becoming shorter apically; bearing
anterior apical comb; pigmented spinulated tubercles absent. Mid femur with a dense
tuft of very short setulae along middle third of ventral surface, and with a fringe of
some 8 long, pale, ventral setae, as long as or longer than femur is deep. Mid tibia with
a posterior row of 4 long setae, and one spine-like subapical seta. Mid basitarsus with
2 spine-like setae ventrally at base; curved, convex laterally, and with a long row of
some 23 short ventral spinules having curved, hook-like apices. Hind leg simple. Hind
femur with a dorsal row of longer setae. Hind tibia with an apical posterior comb of
closely set spinules. Hind basitarsus with an apical posterior comb of short setae.

A NEW MICROPHORELLA FROM TUNISIA 407

FIG. 6

Microphorella cassari sp. n., & , hypopygium, viewed ventrally and from the right. Scale bar
0.2 mm. Abbreviations explained in the text.

Wing (Fig. 3). Two and a half times longer than broad, axillary lobe hardly
developed; wing membrane brown, darker on anterior half of wing, covered with
microtrichia (including veins); macrosetae white; hind marginal fringe longest at base
of wing; wing veins brown, stigma absent; C circumambient; C with 2 strong basal
setae, anterior costal margin with a row of spine-like setae, longer and stronger at
middle of wing; Sc parallel to R,, upturning to € before merging imperceptibly into
membrane very close to R,; humeral crossvein indistinct; R, originating opposite

408 P. GATT

FIG. 7

Microphorella cassari sp. n., 4 hypopygium, viewed dorsally and from the left. Scale bar
0.3 mm. Abbreviations explained in the text.

humeral crossvein; R, meeting C opposite tip of discal cell; R;,,; sinuous, upcurved at
its junction with C; R4.s sinuous, ending in € at tip of wing; crossvein R-M present,
sometimes indistinct; discal cell incompletely separated from second basal cell by
incomplete crossvein BM-Cu, closed distally by crossvein DM-Cu and emitting 3
veins to Wing margin; base of M, complete; crossvein DM-Cu complete; CuA, curved,
Al+CuA, absent. A, present. Alula absent. Squama brown, short, with a fringe of long,
pale hairs. Haltere brown, stem darker than knob, the latter large and quadrate.
Abdomen (Figs 4-5): Brownish black in ground colour, less intensely grey
microtrichose than thorax, rather subshining; sclerites with sparse, white setulae on
posterior margins and on disc. Abdominal muscle plaques distinct. Tergites 1-4 and
sternites 1-3 simple, unmodified. Postabdomen beginning with sternite 5, rotated and
lateroflexed to the right. Sternite 1 very short, narrowly sclerotised only on posterior
and lateral margins, bare except for a covering of microtrichia. Sternites 2 and 3
setulose on disc especially along posterior margins; sternite 4 with a posteromedian
membranous area, fringed on either side with a number of long setae. Sternite 5 short,
bare, with a robust, well sclerotised posteromedian projection. Sternites 6 and 7 bare,
simple. Sternite 8 large, subrectangular; tergite 8 atrophied. Terminalia (Figs 6-7)

A NEW MICROPHORELLA FROM TUNISIA 409

FiGs 8-10

Microphorella cassari sp. n.(8) & phallus and associated structures, lateral view. (9) © antenna,
lateral view. (10) ® abdomen, lateral view (membranes omitted). Scale bars, Figs 8 and 9:
0.2 mm, Fig. 10: 0.5 mm. Abbreviations explained in the text.

lateroflexed to the right, inverted and with caudal pole directed forward, asymmetrical;
hypandrium large, separated from epandrium, produced at apex and with several
accessory processes, microtrichose. Cerci large, the right cercus larger, both deeply

410 P. GATT

incised apically and clothed with setulae, longer setose anteriorly; each with 3 short,
inclinate spine-like setae medially and 2 long, hair-like setae apically.

Phallus and associated structures as in Fig 8; phallus directed forwards, blunt
tipped.

Female

Length. Body 1.4 mm, wing 1.3 mm (specimen in alcohol).

Resembling male, including dichoptic condition of eyes, flattened cephalic and
thoracic setae, apical combs on fore and hind tibia and hind basitarsus, but differing in
the following:

Body and wings somewhat larger. Colour, including legs, darker. Antenna
(Fig. 9) darker, pedicel as long as scape, postpedicel not strap-like, uniformly tapering.
Cephalic and thoracic macrosetae longer, stronger. Mesoscutum with a less micro-
trichose stripe between acrostichal and dorsocentral lines on each side, appearing as a
pair of parallel, longitudinal vittae extending from anterior edge of mesoscutum to
prescutellar depression. AÏl legs with short, undifferentiated setulae. No differentiated
spine-like setae on anterior margin of costa near middle of wing. Abdomen (Fig. 10):
gradually tapering, segments 1-6 forming preabdomen into which posterior segments
are retracted; terminalia not acanthophorous. Tergite 1 short, tergites 2-5 normal.
Tergite 6 with a fringe of long setae on posterior margin. Tergite 7 very narrowly scle-
rotised dorsally, broadening and extending laterally. Tergite 8 (Figs 10-11) long, emar-
ginate anteriorly, not divided medially, depigmented posteriorly. Tergite 10 very short
and narrowly pigmented posteriorly, fused to cerci, complete but depigmented
medially or divided into 2 hemitergites each bearing 3 long setae not forming spines
(Figs 10-11). Sternite 1 short, sclerotised only posteriorly and laterally. Sternites 2-5
normal. Sternite 6 with a fringe of longer setae on posterior margin. Sternite 7 short,
membranous or only very narrowly sclerotised. Sternite 8 (Figs 10, 12) long, produced
posteriorly, depigmented posteromedially; genital fork (= sternite 9) represented (in
part) by 2 small quadrate to rectangular accessory sclerites (Figs 10, 12), separated or
narrowly connected medially, posterior to spermatecha. Sternite 10 divided into 2
sinuous, strap-like hemisternites (Figs 10-11). Cercus (Figs 10-11) broad, bearing setae
of varying lengths on dorsal and lateral surfaces, longest at tip. Spermatheca (Figs 10,
12-13) tubular, receptacle pigmented; middle part of spermathecal duct cylindrical,
pigmented and with tracheated surface; minute dorsal and ventral sclerites (part of
genital fork) posterior to receptacle.

BIOLOGY: AIl specimens were collected from sand dunes or estuaries in spring,
suggesting that this species inhabits sandy, coastal biotopes.

DISTRIBUTION: Hitherto known only from two localities —- Oued Berkoukech and
Oued Bouterfess — on the Tabarka embayment, northwest coast of Tunisia.

REMARKS: The new species described in this paper shows a set of remarkable
morphological features. The strikingly long, strap-like, lanceolate antennal stylus of
the male is unique amongst previously described species of Microphorella, both fossil
and extant. The costal vein of the male bears several long, spine-like setae on the
middle portion that also have not been recorded in other species of this genus. Among

À NEW MICROPHORELLA FROM TUNISIA 411

FIGs 11-13

Microphorella cassari sp. n. (11) ® tip of abdomen, dorsal view. (12) © sternite 8 with
(internally) parts of genital fork and spermatheca, dorsal view. (13) © spermatheca and parts of
genital fork, lateral view. Scale bars, Fig. 11: 0.3mm, Figs 12-13: O.2mm. Abbreviations
explained in the text.

“Microphorinae + Parathalassiinae” modified costal setation is known in the mono-
typic genus Thalassophorus Saigusa, 1986. Although divided female tergites 8 and 10
have been described, a completely divided female sternite 10 has not hitherto been
reported to occur in Microphorella. Similarly, exclusively setose tergites 10 have been
previously reported in only three species of Microphorella, recently described from
Southeast Asia and New Guinea (Shamshev & Grootaert, 2004).

Similar widened and flat cephalic bristles have been described and figured for
Plesiothalassius capensis (Smith) by Ulrich (1991) but have not been known to occur
so far in Microphorella.

412 P. GATT

ACKNOWLEDGMENTS

I am grateful to the MEDCORE (Mediterranean Coastal River Ecosystems)
Project co-ordinators, in particular Prof. Felicita Scapini (Dipartimento di Biologia
Animale e Genetica ‘Leo Pardi’, University of Florence, Italy) for facilitating my
participation in the project. Dr. Louis F. Cassar (Institute of Earth Systems, University
of Malta) is thanked for introducing me to the type locality of this species. I am also
indebted to Dr. Igov Shamshev (AlÏl-Russian Institute of Plant Protection, St.
Petersburg, Russia) for kindly reading a draft version of the manuscript and suggesting
improvements. My trip to Tunisia in 2005 was funded by the European Commission
INCO-DC Programme (MEDCORE Project ICA-3-CT 2002-10003).

REFERENCES

CHVALA, M. 1988. Revision of Palaearctic Microphoridae (Diptera) 3. Parathalassiinae
(Parathalassius Mik and Microphorella Becker). Acta Entomologica Bohemoslovaca
85: 352-372.

COLLESS, D. H. 1963. An Australian species of Microphorella (Diptera: Empididae), with notes
on the phylogenetic significance of the genus. Proceedings of the Linnean Society of
New South Wales 88: 320-323.

DISNEY, R. H. L. 1983. Scuttle Flies. Diptera, Phoridae (except Megaselia). Handbooks for the
Identification of British Insects 10 (6): 1-81; London.

GATT, P. 2003. New species and records of Microphorella Becker (Diptera: Empidoidea,
Dolichopodidae) from the Mediterranean region. Revue suisse de Zoologie 110 (4):
669-684.

MELANDER, À. L. 1928 (1927). Diptera, Fam. Empididae. Zn: WYTSMAN, P. (ed.). Genera insec-
torum 185: 1-434 & 8 plates. Bruxelles.

MERZ, B. & HAENNI, J. P. 2000. Morphology and terminology of adult Diptera (other than
terminalia) (pp. 21-51). Zn: PAPP, L. & DARVAS, B. (eds). Contributions to a Manual of
Palaearctic Diptera 1: 1-978. Science Herald, Budapest.

SHAMSHEV, I. V. 2004. A new species of the genus Microphorella Becker from the Far East of
Russia, with notes on some morphological features in the Microphorinae (Diptera:
Empidoidea). Studia Dipterologica 10(2) (2003): 527-535.

SHAMSHEV, I. V. & GROOTAERT, P. 2004. Descriptions of four new species of the genus
Microphorella Becker (Diptera: Empidoidea, Microphoridae, Parathalassiini) from
southeast Asia and New Guinea, with notes on the relationships within the genus. The
Raffles Bulletin of Zoology 52(1): 45-58.

SINCLAIR, B. J. 2000. Morphology and terminology of Diptera male terminalia (pp. 53-74). /n:
PaApPp, L. & DARVAS, B. (eds). Contributions to a Manual of Palaearctic Diptera 1:
1-978. Science Herald, Budapest.

SINCLAIR, B. J. & CUMMING, J. M. 2006. The morphology, higher-level phylogeny and classifi-
cation of the Empidoidea (Diptera). Zootaxa 1180: 1-172.

ULRICH, H, 1991. Two new genera of parathalassiine-like flies from South Africa (Diptera,
Empodoidea). Bonner zoologische Beiträge 42(2): 187-216.

ULRICH, H. 2003. How recent are the Empidoïidea of Baltic amber? Srudia Dipterologica 10:
321-327.

REVUE SUISSE DE ZOOLOGIE 118 (3): 413-421; septembre 2011

Redescription of Rhacophorus chuyangsinensis Orlov, Nguyen &
Ho, 2008 (Anura: Rhacophoridae) based on new collections from
new south Vietnamese provincial records: Lam Dong and Khanh
Hoa

Dao Thi Anh TRAN!#, Tao Thien NGUYEN?, Trung My PHUNG#, Tri LY!,

Wolfgang BÔHME#, and Thomas ZIEGLER

l Vietnam National University, Ho Chi Minh City, University of Science, Faculty of
Biology, Department of Ecology & Evolutionary Biology, 227 Nguyen Van Cu,
District 5, Ho Chi Minh City, Vietnam. E-mail: ttadao.hemuns@ gmail.com

2 Vietnam National Museum of Nature, 18 Hoang Quoc Viet, Hanoï, Vietnam.

39A Dong Khoi, Tam Hiep, Bien Hoa, Dong Nai Province, Vietnam.

4 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160,
D-53113 Bonn, Germany. E-mail: w.boehme.zfmk@uni-bonn.de

5 AG Zoologischer Garten Kôln, Riehler Strasse 173, D-50735 Cologne, Germany.
E-mail: ziegler@koelnerzoo.de

Redescription of Rhacophorus chuyangsinensis Orlov, Nguyen & Ho,
2008 (Anura: Rhacophoridae) based on new collections from new south
Vietnamese provincial records: Lam Dong and Khanh Hoa. -
Rhacophorus chuyangsinensis Orlov, Nguyen, & Ho, 2008, which was de-
scribed based on a type series consisting of three adult males only, is re-
described based on extensive new collections from southern Vietnam. Our
new records consist of 17 individuals, among them the first two females to
become known. In our extended description we deal for the first time with
adult female morphology and with so far unknown colour pattern in life.
Our new records of R. chuyangsinensis for Lam Dong and Khanh Hoa
provinces expand the originally known distribution of this species about 81
km to the Southeast of 1ts type locality (Chu Yang Sin National Park, Dak
Lak Province, southern Vietnam, 1,600 m a.s.1.). We further add additional
information on the natural history of À. chuyangsinensis, which inhabits
rocky forest streams at altitudes between 1,320-1 ,600 m a.s.l.

Keywords: Anura: Rhacophoridae: Rhacophorus chuyangsinensis - mor-
phology, taxonomy, new distribution data, natural history - Vietnam: Lang
Bian Plateau.

INTRODUCTION

Rhacophorus chuyangsinensis Was recently described by Orlov ef al. (2008)
based on a type series consisting of three adult males from Kon Tum Plateau, Vietnam.
This species was so far known only from the type locality in Chu Yang Sin National
Park, Dak Lak Province, Central Vietnam (Nguyen ef al. 2009). During recent

Manuscript accepted 20.06.2011

414 D. T. A. TRAN ET AL

herpetological surveys in the Southeast of the type locality, viz. in Bi Doup-Nui Ba
National Park (Lam Dong Province) and Hon Ba Nature Reserve (Khanh Hoa
Province), southern Vietnam (Fig. 1), extensive collections of this species took place,
among them the first females known to science and a previously unknown colour
pattern. Therefore, we herein provide an extended description of R. chuyangsinensis
based on our new records from southern Vietnam.

MATERIAL AND METHODS

Specimens were collected in the evergreen forests of Bi Doup-Nui Ba National
Park, (Lac Duong District, Lam Dong Province: 12°00°-12°52°N, 108°17-108°42’E;
600-2000 m a.s.1.) between March and June 2010 (periods of 18-25 March, 26-30
April, 20-24 May, and 07-19 June) and of Hon Ba Nature Reserve (Dien Khanh
District, Khanh Hoa Province: 12°02°-12°15°N, 108°57-109°05’E; approximate
1600 m a.s.1) in September 2010 (period of 10-15 September).

Frogs were collected by hand from 19:00-23:00. After taking photographs,
specimens were anaëesthetized, fixed in 80% ethanol for few hours, and subsequently
preserved in 70% ethanol. Specimens finally were deposited in the collections of the
University of Science, Ho Chi Minh City, Vietnam (US); the Vietnam National
Museum of Nature (VNMN), Ha Noi, Vietnam and the Zoologisches Forschungs-
museum Alexander Koenig (ZFMK), Bonn, Germany. Adults (4 4): UNS 00500-
00508, VNMN 965, ZFMK91517-91522; Adults (9 © ): UNS 00509, VNMN 960.

Measurements were taken with a digital caliper to the nearest 0.1 mm: SVL
(snout-vent length): distance between tip of snout and vent; HW (head width): distance
between angles of jaws; HL (head length): distance between angle of jaws and snout
tip; SNL (snout length): distance between anterior corner of eye where the upper and
lower lids meet together and the tip of snout; NS (distance between nostril and snout
tip): distance between middle of nostril and tip of snout; IN (internarial distance): dis-
tance between nostrils; IO (interorbital distance): least distance between upper eyelids:
UEW (upper eyelid width): greatest width of upper eyelid; ED (eye diameter): hori-
zontal width of eye at its widest point; DFE (distance between front of eyes): distance
between anterior points of eyes; DBE (distance between back of eyes): distance
between posterior points of eyes; TD (tympanum diameter): horizontal width of tym-
panum at its widest point; E-T (distance between eye and tympanum): distance
between posteriormost point of eye and anteriormost edge of tympanum; distance
between axilla and groin (A-G): distance between posterior edge of forelimb at its
insertion to body and anterior edge of hind limb at its insertion to body; length of upper
arm (UAL): distance between axilla and elbow; length of lower arm (LAL): distance
between elbow and posteriormost margin of inner palmar tubercle; length of hand
(HAL): distance between proximal edge of palmar tubercle and tip of the third finger;
length of thigh (THL): distance between center of knee and center of hindlimb
insertion; length of tibia (TBL): distance between center of knee and center of heel;
foot length (FOL): distance between base of inner metatarsal tubercle and tip of the
fourth toe; tarsus-foot length (TFOL): distance between base of tarsus and tip of the
fourth toe; length of finger or toe: distance between posterior margin of most proximal
subarticular tubercle or crease of articulation and tip of finger or toe; first finger length

REDESCRIPTION OF RHACOPHORUS CHUYANGSINENSIS 415

104° 108° 112°

104° 108° 112°

FIG. 1

Map showing the distribution of Rhacophorus chuyangsisensis in Vietnam (blue: type locality:
red: new records from Lam Dong and Khanh Hoa provinces, Vietnam).

(FEL); first toe length (FTL); width of disc on finger or toe: greatest width of terminal
disc on finger or toe; third finger’s disc width (TFPW); length of inner metatarsal
tubercle (IMTL); length of nuptial pad (NPL); formula of webbing followed Glaw and
Vences (2007). The sex of the specimens was superficially determined based on the

416 D. T. A. TRAN ET AL

absence or presence of male nuptial pads; female sex was proven by the examination
of the reproductive organs after dissection. Morphological identification and
comparisons followed the original description (Orlov ef al. 2008).

RESULTS
REDESCRIPTION OF RHACOPHORUS CHUYANGSINENSIS ORLOV, NGUYEN & HO, 2008

In the following we provide a detailed morphological description of R. chu -
vangsinensis based on 15 adult males and two adult females from Lam Dong and
Khanh Hoa Provinces. southern Vietnam (for measurements see Table 1).

Head approximately as long as wide; snout slightly pointed in dorsal view,
pointed and slightly exceeding to mouth in profile: nostril round, and closer to tip of
snout than to eye: canthus rostralis distinct; eye diameter 0.7-0.9 times of snout length:
interorbital region flat, wider than internarial distance and as large as or little bit larger
than width of upper eyelid; tympanum round, not raised above temporal region, with a
slightly elevated rim; diameter of tympanum 0.4-0.5 times of eye diameter, distance
between eye and tympanum 0.3-0 4 times the tympanum diameter; supratympanic fold
distinct, from behind of eye to beyond level of axilla; choanae small, round; vomerine
teeth grouped in two oblique rows, beginning nearby anterior edges of the choanaes,
closer to choanae than to each other: tongue bifid at rear.

Limbs slender:; relative lengths of fingers I<II<IV<IIT:; tips of fingers flat, en-
larged into round discs with circummarginal grooves: disc of third finger as large as or
little larger than the tympanum diameter: fingers incompletely webbed, formula I(1)
IHi(1)e(0.5) Ii(1.5)e(1) IV(1); narrow, smooth flap of skin present along outside of
fourth finger and lower arm, ending at elbow; subarticular tubercles on fingers and toes
round, prominent and conspicuous; nuptial pad distinct in males, located on lateral and
dorsal aspect of first finger, from near its base to proximal end of penultimate phalanx:
length of nuptial pad slightly larger than that of the first finger:; discs of toes round with
circummarginal grooves, smaller than those of fingers; relative lengths of toes
IKII<V<III<IV; toes broadly webbed, formula I(0) Ili(1)e(0.5) ITHi(1)e(0)
[Vi(0.5)e(0.5) V(0.5); dermal fringe along outside of fifth toe and foot narrower than
that along outside of fourth finger and lower arm, and ending at tibiotarsal articulation
with a long, pointed projection; inner metatarsal tubercle flat, oval, its length about
one-third to half of that of first toe; outer metatarsal tubercle absent: heels overlapped
when legs are held at right angles to body; tibiotarsal articulation extending to anterior
edge of eye or between eye and tip of snout.

Skin smooth on dorsal body, head, and limbs; throat, chest, and lower part of
flank slightly granular; belly and ventral surface of thigh coarsely granular: posterior
cloacal appendix present.

Coloration in preservative. Dorsal surfaces of body and limbs ground yellow to
dark brown; dorsal pattern which is yellow in life becomes creamish-white in preser-
vative; nuptial pad white; surfaces of belly, limbs, discs and webbings whitish-cream:
the blue color surrounding the black patches on flanks and upper arms is only slightly
discernible if at all.

Coloration in life. Back and upper surfaces of limbs pale green or dark brown,
with many small white or yellow spots; ventral surface bright yellow without small

REDESCRIPTION OF RHACOPHORUS CHUYANGSINENSIS 417

TABLE 1. Measurements (mean +standard deviation, followed by minimum and maximum in
parentheses; in mm) of Rhacophorus chuyangsinensis from Bi Doup-Nui Ba National Park, Lam
Dong Province and Hon Ba Nature Reserve, Khanh Hoa Province, Vietnam; n: number of
specimens. See the methods for abbreviations.

Males (n = 15)

Females (n 2)

SVL 37.9 +2.6 (34.8-43.8) 59.1 +1.6 (58.0—-60.2)
HL 14.7 +0.7 (13.5-16.1) 215209 012-219)
HW 15.0 +0.6 (14.17-16.3) 22.2 +0.4 (21.5-22.9)
SNL 6.0 +0.7 (3.7-6.5) 7.3 +1.6 (6.1-8.4)
NS 2.3 +0.3 (1.8-2.8) 3.4 +0.3 (3.2-3.6)

IN 59202 (29-35) 4.6 +0.7 (4.1-S.1)
ED 4.9 +04 (4-54) 5.9 +0.1 (5.9-6.0)
UEW 3.8 +0.6 (2.4-4.6) 3.3 +1.7 (2.145)

10 4.5 +0.8 (3.8-7..9) 8.5 +3.5 (60-110)
DFE 8.3 +0.6 (7.4-9.3) 11.7 +0.8 (11.1-12.3)
DBE 12.9 +1.3 (9.7-14.3) 17.0 +0.7 (16.5-17.5)
TD 2.4 +0.5 (20-42) 4.1 +0.9 (3.548)
E-T 1.0 +0.2 (0.6-1.3) 1.9 +0.3 (1.8-2.1)
A-G 20.9 +1.3 (18.5-23.2) 33.5 +1.0 (32.8- 34.2)
UAL 6.9 +0.4 (6.4—7.8) 11.8 +0.6 (11.3-12.2)
LAL 7.2 +0.6 (6.5-8.2) 11.0 +0.1 (10.9-11.1)
HAL 12.1 +0.8 (110-134) 17.4 +0.2 (17.3-17.5)
TFPW 1.8 +0.3 (2.0-2.8) 3.3 +0.1 (3.3-3.4)
THL 18.9 +1.3 (16.7-20.7) 28.2 +1.2 (27.3-290)
TBL 20.1 +1.0 (19.0-21.7) 29.9 +0.5 (29.5-30.3)
FOL 17.1 +2.6 (13.9-24 8) 24.2 +0.1 (24.1-243)
TFOL 27.0 +1.4 (24.3-29.2) 39.3 +0.8 (38.7-39.9)
IMTL 1.3 +0.2 (0.9-1.7) 1.6 +0.5 (1.27-1.5)
FTL 3.0 +0.5 (2.243) 5.6 +2.3 (40-72)
NPL 3.3 +03 (2.8-3.7) -

spots; thin, light stripes present along canthus rostralis, from tip of snout to middle of
the eye: flanks, anterior and posterior surfaces of limbs yellow to orange; axilla, groin,
anterior surface of upper arm and thigh, and posterior surfaces of thigh and tibia
usually with large, black patches that vary in size and shape; these patches are some-
times surrounded by bright blue color and are more obvious in females (more details
are given in the discussion of the sexual dimorphism); webbings on fingers and toes
yellow or orange, sometimes with black pattern at base; discs of fingers and toes
yellow to orange; posterior cloacal appendix white; long, pointed projection at the heel
of the same color as dorsal surface of thigh or yellow; pupil horizontal, black; iris
brown, fading into red-orange at the upper and lower parts and surrounded by an inner
black and outer blue circle (Fig. 2). Dorsum of some males (UNS00506-00508;
ZFMK91517-91519) reddish brown, with two yellow dorsolateral stripes stretching
from posterior corner of eye to groin and a medium third yellow stripe extending from
behind the middle of the back towards cloaca; in such coloured specimens, also the
light snout stripes are more distinctly developed, and sometimes a light stripe between
eyes 1s present, forming a triangle on the dorsal surface of head (UNS0O0506 &
ZFMK91519) (Fig. 2 E & F).

Sexual dimorphism. Female size on average 1s 1.6 times larger than that of
males. The females also differ from males by their flank pattern, which consists of the

418 D. T. A. TRAN ET AL

FIG. 2

Different color patterns of Rhacophorus chuyangsinensis in life. (A-B) female (VNMN965) at
day and night time. (C-F) males (C: UNS00500, D: UNSO00508, E: ZFMK91518, and F:
UNSO00506).

same colour than the dorsal pattern, viz. dark reddish brown or ground brown, with
many small whitish spots (versus bright yellow flanks in males without small whitish
spots). In females the bright bluish color on axilla, groin and front of the forearm 1s
also more obvious and furthermore may even extend towards the middle of the flanks

REDESCRIPTION OF RHACOPHORUS CHUYANGSINENSIS 419

and cover nearly the whole anterior surface of the forearm (Figs 2A, 2B). In addition,
the posterior surface of the thigh and tibia of females is covered with large black, elon-
gated patches.

EXTENDED DIAGNOSIS (AFTER ORLOV ET AL. 2008, COMBINED WITH OUR NEW DATA)

A small rhacophorid species with 35.1-44.15 mm SVL in adult males, and
58.0-60.2 mm in adult females; body depressed; head wide and flat, approximately as
long as wide; snout somewhat pointed; diameter of the eye 0.7-0.9 times of snout
length; iris brown to red-orange at the upper and lower parts, surrounded by inner black
and outer blue circle; pupil black, horizontal; tympanum round, small but clearly visi-
ble; supratympanic fold extending just beyond level of axilla; back and dorsal surfaces
of limbs smooth; throat slightly granulated, belly and ventral surface of thigh coarsely
granular; vomerine teeth in two oblique ridges that reach the upper part of the round-
ed choanas; discs of fingers and toes flat, large, round, with circummarginal grooves;
discs on fingers larger than toe discs; fingers incompletely webbed (formula: I(1)
IHi(1)e(0.5) Ii(1.5)e(1) IV(1)); toes extensively webbed (formula: I(0) IHi(1)e(0.5)
IHi(1)e(0) IV1(0.5)e(0.5) V(0.5)); dermal fringe along the outside of the fourth finger
and lower arm present; similar ridge of skin present along the outside of the fifth toe
and foot, ending at heel, which bears a long, pointed projection; back and dorsal sur-
faces of limbs pale green to dark green or dark brown, with many small white or yel-
low spots; belly yellow without spots; webbings on fingers and toes yellow to orange;
posterior cloacal appendix white.

NATURAL HISTORY

The new records of Rhacophorus chuyangsinensis from Bi Doup-Nui Ba
National Park (Lam Dong Province) and Hon Ba Nature Reserve (Khanh Hoa
Province) were made nearby or within rocky streams in evergreen forests at elevations
between 1,320-1 ,600 m a.s.1. The frogs were observed during evening and night time
sitting on high branches of trees along the streams, approximately 1.5-2 m above the
ground. Sometimes these trees were up to 5 m distant from water sources (Fig. 3). On
22 May 2010, we measured environmental temperatures of 19.2—-19.5°C, and humi-
dities of 92.4-93.6% at the sites where the frogs were found. One female (UNS00509),
which was collected during dry season on 18 March 2010, was gravid and contained
large eggs up to 3 mm diameter.

DISCUSSION

Our new rhacophorid records from Bi Doup-Nui Ba National Park (Lam Dong
Province) and Hon Ba Nature Reserve (Khanh Hoa Province) were morphologically
well assignable to the recently described species Rhacophorus chuyangsinensis (Orlov
et al. 2008). But we also found some differences between the new collections and the
type series of R. chuyangsinensis. Our new records from Lam Dong and Khanh Hoa
provinces had the interorbital distance as large as or slightly larger than the width of
the upper eyelid (in contrast to the condition described for two of the three males by
Orlov ef al. 2008); the nuptial pad length was 1.0-1.3 times the first finger length in

420 D. T. A. TRAN ET AL

FIG. 3

Habitat of Rhacophorus chuyangsinensis in (A) Bi Doup-Nui Ba National Park, Lam Dong
Province; and (B) Hon Ba Nature Reserve, Khanh Hoa Province, Vietnam.

our new series (versus only half of finger’s length according to Orlov ef al. 2008); the
tympanum diameter was nearly half (0.4-0.5) of the eye diameter in the new series
(versus tympanum diameter being 0.6-0.7 of eye diameter according to Orlov ef al.
2008). Based on our new records of R. chuyangsinensis we could also show that adult
males have a wider range in snout-vent length than it was noted previously (only
42.9-44.2 mm after Orlov ef al. 2008 versus 35.1-43.8 mm in the new series from Lam
Dong and Khanh Hoa). Besides providing first data on the size and the so far unknown
colour pattern of the females of this recently described species, the first record of a so
far unknown, both triangle-shaped and striped light dorsal colour pattern in male À.
chuyangsinensis are particularly noteworthy. Because these unusually patterned males
occurred in the same microhabitat and because we did not find significant morpho-
logical differences compared to normal patterned congeners, we evaluate them as
representing the same species. However, future bioacoustic and molecular approaches,
which are lacking at time, must confirm the conspecific status of the different colour
morphs. Orlov ef al. (2008) could not document male advertisement calls of À.
chuyangsinensis during October 2007. Thus, these authors assumed that the species
probably does not reproduce at that time of the year. Our first record of a gravid female
collected at the end of March 2010 indicates that reproduction at least takes place in
the dry season. Beyond an extended distribution range of the species we also could
show that this species can also be found at lower altitudes, viz. between 1,320-1 ,600
m a.s.l.

ACKNOWLEDGEMENTS

The first author thanks the staff of Bi Doup-Nui Ba National Park, especially
the staff of Hon Giao and Giang Ly stations for their support. Phung My Trung thanks
Mr. Le Van Nong and the staff of the Hon Ba Nature Reserve for their support during
field work. We also thank Ta Van Thuc, Da Du Ha Tien, Nguyen Van Cuong, Le Thi
Thuy Duong, Le Thi Thanh Ngan, Nguyen Dinh Hien and Le Thanh Hung for their
great assistance in the field. We acknowledge Nguyen Quang Truong and two ano -

REDESCRIPTION OF RHACOPHORUS CHUYANGSINENSIS 421

nymous reviewers for commenting on the manuscript. The surveys have been
conducted under permission of 1430/GT-BNN-KL. This study was partially supported
by the Ministry of Education and Training of Vietnam (MOET, Project 322), the
Alexander Koenig Gesellschaft (AKG), the Idea Wild, and the German Academic
Exchange Service (DAAD).

REFERENCES

GLAW, F. & VENCES, M. 2007. A field guide to the amphibians & reptiles of Madagascar. Third
edition. Cologne, Vences and Glaw Publishers, 496 pp.

NGUYEN, V.S., HO, T. C. & NGUYEN, Q. T. 2009. Herpetofauna of Vietnam. Edition Chimaira,
Frankfurt am Main, 768 pp.

ORLOV, N. L., NGUYEN, N.S. & HO, T. C. 2008. Description of a new species and new records
of Rhacophorus genus (Amphibia: Anura: Rhacophoridae) with the review of amphib-
ians and reptiles diversity of Chu Yang Sin National Park (Dac Lac Province, Vietnam).
Russian Journal of Herpetology 15 (1): 67-84.

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REVUE SUISSE DE ZOOLOGIE 118 (3): 423-449; septembre 2011

The Psocoptera (Insecta: Psocodea) of St Helena and Ascension
Island (South Atlantic) with a new record from South Africa

Charles LIENHARD! & N. Philip ASHMOLE?

l Muséum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland.
E-mail: charleslienhard@bluewin.ch

2 Kidston Mill, Peebles, EH45 8PH, UK.

The Psocoptera (Insecta: Psocodea) of St Helena and Ascension Island
(South Atlantic) with à new record from South Africa. - Four new
species are described: Cerobasis atlantica Lienhard sp. n. (Trogiidae) from
St Helena, Sphaeropsocopsis insularum Lienhard sp. n. (Sphaeropsocidae)
from St Helena and Ascension Island, /ndiopsocus mendeli Lienhard sp. n.
(Psocidae) from Ascension Island and Blaste helenae Lienhard sp. n. (Pso-
cidae) from St Helena. The latter is closely related to the St Helena endemic
Blaste basilewskyi Badonnel; this could be an example of sympatric spe-
ciation. Helenatropos abrupta Lienhard, formerly supposed to be a St
Helena endemic, is for the first time recorded from South Africa and its
male 1s described; it may have been introduced to St Helena. The recently
published doubtful record of the Mexican species Cerobasis maya Garcia
Aldrete from Ascension Island is confirmed. The male of the blind cave-
dwelling St Helena endemic Sphaeropsocopsis myrtleae Lienhard &
Ashmole is described for the first time; its genital morphology indicates a
close relationship to the African Sphaeropsocopsis reisi Badonnel. Several
other species are recorded for the first time from one or both of these
islands. The number of species recorded from St Helena is raised to 23 (6
endemics), that from Ascension Island to 13 (2 endemics). À checklist of the
27 psocid species recorded from these islands 1s presented and a brief bio-
geographical analysis is provided.

Keywords: Trogiidae - Sphaeropsocidae - Psocidae - new species - new
records - cave fauna - blind psocid - island endemics - island biogeography.

INTRODUCTION

St Helena and Ascension Island, 1300 km apart and respectively 1800 and 1500
km from continental land, are among the most isolated islands in the world. Similar in
size (respectively 122 and 97 km?) and in their origin as volcanic oceanic islands
formed near the Mid-Atlantic Ridge, they differ dramatically in their age: St Helena is
about 14 million years old but Ascension probably emerged only about one million
years ago. Study of the biology of this pair of islands offers snapshots of two very
different stages in the development of ecosystems in situations so remote that natural

Manuscript accepted 20.04.2011

424 C. LIENHARD & N. P. ASHMOLE

colonization by plants and animals is a rare event and evolution can proceed in iso -
lation. On St Helena, now geologically stable but with heavily eroded coasts and land-
scapes, processes of phyletic change, the splitting of lineages and interactions among
species have created a mature and diverse biota (Ashmole & Ashmole, 1997, 2000a).
Ascension, an order of magnitude younger and still with raw volcanic terrain, shows
early stages in these same processes, while the paucity of native plants ensures that the
low diversity invertebrate fauna is dominated by scavengers and predators.

Knowledge of the psocids of St Helena and Ascension Island has developed
mainly as a result of a series of collecting opportunities in the past two decades. In
1995 Philip and Myrtle Ashmole spent five months in field work on St Helena, prima-
rily attempting to find invertebrate species adapted to subterranean life, but with
limited success (Ashmole & Ashmole, 2000a: 131-132). More diverse collections of
psocids were obtained in 2003, during a survey of invertebrates of Prosperous Bay
Plain in connection with the proposed airport (Ashmole & Ashmole, 2004a, 2004b). In
2006 a survey of the invertebrate communities of the Central Peaks of St Helena
commissioned by the St Helena National Trust (Mendel ef al., 2008) showed that a few
psocids occur in the cloud forest, while collecting away from the Peaks at this time
provided new data on previously recorded species and led to discovery of one new
endemic species.

In 1990 and 1995 Philip and Myrtle Ashmole investigated the invertebrate
fauna of lava flows, pyroclastics and volcanic caves on Ascension Island (Ashmole &
Ashmole, 1997, 2000a, 2000b). Although psocids had not been found previously on the
island, the use of specialised trapping techniques showed that they were actually
widespread and diverse in the most barren habitats and led to discovery of a generic
endemic species in subterranean habitats. Howard Mendel collected invertebrates on
and around Green Mountain in 2003, adding several species to the known fauna, and a
little further collecting was done there by the Ashmoles in 2005/06.

Previous knowledge of the psocid fauna of St Helena and Ascension Island has
been summarized by Badonnel (1976), Ashmole & Ashmole (1997, 2000a) and
Lienhard & Smithers (2002; for corresponding online species lists see Lienhard,
2004b).

Mendel ef al. (2008) reviewed the data for St Helena, indicating that 22 species
of Psocoptera had been recorded up to January 2008, including two unidentified
species of the genus Liposcelis Motschulsky and three species probably new to science
in the genera Cerobasis Kolbe, Sphaeropsocopsis Badonnel and Blaste Kolbe. These
new species are described and illustrated in the following. However, the two unidenti-
fied species of Liposcelis (material deposited in the MHNG) are only mentioned 1in the
checklist (see Appendix) but not treated in this paper. We also publish here for the first
time the St Helena records of Lepinotus inguilinus and Liposcelis entomophila, already
listed in the above mentioned report (Mendel et al., 2008), and one recent record of
Stenocaecilius caboverdensis, a species also known from Ascension Island, raising the
final number of species known from St Helena to 23 (see checklist in Appendix).

The psocid fauna of Ascension Island is apparently poorer, but also less inves-
tigated, than that of St Helena. The eight species treated by Ashmole & Ashmole
(1997) are also listed by Lienhard & Smithers (2002). One of them is an unidentified

PSOCOPTERA OF ST HELENA AND ASCENSION 425

Liposcelis species, which is mentioned in the checklist (see Appendix) but not treated
in the following (material deposited in the MHNG). The Sphaeropsocopsis species
listed by Ashmole & Ashmole (1997, 2000a) as cf. microps is here described as a new
species, based on material from St Helena and Ascension Island. The record of
Cerobasis maya from Ascension Island can now be confirmed; it was previously men-
tioned as Cerobasis cf. maya by Ashmole & Ashmole (1997, 2000aà). By adding a new
species of the genus /ndiopsocus Mockford, described below, and records of Cerobasis
guestfalica, Stenocaecilius caboverdensis, Peripsocus leleupi and Peripsocus pauliani
we raise the total number of species known from Ascension Island to 13 (see checklist
in Appendix).

In the following we present the descriptions of the new species and the new
records, together with the first South African record of Helenatropos abrupta. Habitus
figures are given for the most interesting endemics, the subterranean and troglo-
morphic Troglotroctes ashmoleorum (Ascension Island, Fig. 4a) and Sphaeropsocopsis
myrtleae (St Helena, Fig. 6a). The 27 psocid species known from St Helena and
Ascension Island are listed in the Appendix and treated in a brief biogeographical
discussion.

MATERIAL AND METHODS

Dissection and slide-mounting followed the methods described by Lienhard
(1998). The material examined has been deposited in the following institutions:
BMNH = The Natural History Museum, London, UK; MHNG = Muséum d'histoire
naturelle, Geneva, Switzerland; NMSE = National Museum of Scotland, Edinburgh,
UK; SEHU = Systematic Entomology, Hokkaido University, Sapporo, Japan; UNAM
= Universidad Nacional Auténoma de México, México City.

The following abbreviations are used in the descriptions: BL = body length (in
alcohol); F = hindfemur (length); F+tr = hindfemur and trochanter (length); FW =
forewing (length); I0/D = shortest distance between compound eyes divided by ante-
roposterior diameter of compound eye in dorsal view of head; PI1-P4 = articles of
maxillary palp; T = hindtibia (length); t1, t2, t3 = tarsomeres of hindtarsus (length,
measured from condyle to condyle); V = width of head capsule on vertex.

Bibliographical references of original taxa descriptions not given in this paper
can be found in Lienhard & Smithers (2002).

TAXONOMIC TREATMENT
TROGIIDAE

Helenatropos abrupta Lienhard, 2005 Fig. 1
Helenatropos abrupta Lienhard, 2005a: 691; description of female from St Helena.

TYPE MATERIAL: MHNG and BMNH, 39 (holotype and two paratypes), mentioned by
Lienhard (2005).

NEW MATERIAL: MHNG and BMNH, 8, 11%, South Africa, Cape Town, Table
Mountain National Park, mostly from pine plantations, sometimes from "Fynbos" shrubland,
collected by leaf litter extraction (12 individuals), pitfall traps (3 ind.), sugar-baited ant traps (3
ind.) and on Protea log (1 ind.), leg. C. Uys or C. Uys & M. Picker, May, October, November
2008 and January 2009.

426 C. LIENHARD & N. P. ASHMOLE

EG:
Helenatropos abrupta Lienhard, male: (a) Hypandrium. (b) Phallosome. (c) Right paraproct.

DESCRIPTION OF MALE: Colouration and general morphology as described for
the female by Lienhard (2005a). Sclerotized metanotal winglet-like lobes of males
usually almost touching each other medially as in most females from South Africa and
from St Helena [Note: In the holotype figured by Lienhard (2005a: figs 1, 6) these
lobes are exceptionally well-separated medially]. Epiproct simple, paraproct nearly
triangular (Fig. 1c), its dorsal part sclerotized, its membranous ventral part strongly
shortened. Hypandrium heavily sclerotized, basally with a well-developed hypandrial
brush, consisting of about 20 acuminate setae (Fig. 1a). Phallosome simple (Fig. 1b),
lacking conspicuous internal sclerotizations, parameres distally bifurcate, their
posterior part with an anteriorly curved tip. Measurements (4 MNHG 7987, um): BL
= 1700; F= 350: T=436:H=130 04719 506.

DISCUSSION: This species was tentatively considered as a St Helena endemic by
Lienhard (2005a). However, in the discussion of the original description it 1s men-
tioned that soil-dwelling psocids of the African continent, at present rather poorly

PSOCOPTERA OF ST HELENA AND ASCENSION 427

investigated, could represent a more or less recent source of colonization for the island
of St Helena.

During an ecological research project in the Table Mountain National Park
numerous specimens of H. abrupta Were recently collected in South Africa (see
material mentioned above). The South African females are identical to the St Helena
specimens, therefore both populations have to be assigned to the same species. The
existence of this species on the island of St Helena is probably due to introduction from
South Africa, probably along with the creeping plant Carpobrotus edulis (Aïzoaceae)
which was brought to St Helena from South Africa in the 19h century. The type
material was collected on St Helena in pitfall traps set among mats of this creeper.

The monotypic genus Helenatropos Lienhard is characterized by a series of stri-
king autapomorphies (see Lienhard, 2005a: 695). The presence of a forked sensillum
on P4 in both sexes (see Lienhard, 2005a: fig. 4, 1998: fig. 261), of a well-developed
hypandrial brush (Fig. la) and of a simple phallosome with distally bifurcate para-
meres (Fig. 1b) could indicate a relatively close relationship to the genus Lepinotus
Heyden. However, the phylogenetic position of Helenatropos Within the Trogiidae
could only be elucidated by careful analysis of the 9 other genera of this family, which
is not the purpose of this paper; these genera are listed by Lienhard & Smithers (2002)
and Li Fasheng (2002).

Cerobasis atlantica Lienhard sp. n. Fig. 2

HOLOTYPE: MHNG, G , St Helena, Earwig Gully, Prosperous Bay Plain, S15°57.459'
W5°39.059', ca 290-310m, a gully southeast of the Central Basin, 27.1x.2003, leg. P. & M.
Ashmole (site PBP4, sample 79).

PARATYPE: MHNG, ©? (allotype), St Helena, Cliff Top, S15°57.318' W5°39.873', ca 290-
310m, part of the cliff edge east of Prosperous Bay Plain, 27.1x.2003, leg. P. & M. Ashmole (site
PBP3, sample 61).

DESCRIPTION: Colouration: Body light yellowish white with a small dark brown
median patch on pronotum and some tiny brown spots on abdominal tergites, arranged
in segmental transversal rows. Compound eyes black, labrum dark brown, flagello -
meres apically brown, in distal half of antenna almost completely brown. Winglets
unpigmented, no brown transversal tibial rings recognizable on legs.

Morphology: Maxillary palp with P4 much enlarged and slightly shorter than P2
(Fig. 2c), lacking forked sensillum. Lacinial tip with three relatively shallow tines (Fig.
2b). Forewing reduced to an oblong winglet, bearing a longitudinal row of 3-5 stout
truncate setae in addition to the normal pilosity (Fig. 2a), hindwing absent. Winglets
laterally clearly protruding from mesothorax in dorsal view, not covering lateral parts
of metanotum as in many other Cerobasis species (see Lienhard, 1998: fig. 24b).
Therefore dorsal pilosity not only well-developed in middle but also on each lateral 1/3
of metanotum. Mesonotum relatively long (length almost equal to half width of
vertex), its hindmargin slightly indented laterally in dorsal view. Pearman's organ of
hindcoxa well-developed, hindtibia with 4 terminal spurs and 3 internal spurs in apical
half (one hindtibia of the female with 4 internal spurs of normal size and one additional
short internal spur close to the apical spurs). Pretarsal claws lacking preapical tooth,
with basal appendix and slightly enlarged membranous pulvillus.

428 C. LIENHARD & N. P. ASHMOLE

FIG. 2

Cerobasis atlantica Lienhard sp. n., male holotype (a-d), female allotype (e-g): (a) Right meso-
thoracic winglet. (b) Tip of lacinia. (c) Maxillary palp (pilosity not shown, except for spur
sensillum of P2). (d) Phallosome. (e) Gonapophyses. (f) Spermathecal parietal gland. (eg)
Spermapore region.

Hypandrial brush with about 40 acuminate setae. Phallosome as in Fig. 2d,
lacking strongly sclerotized internal structures near apex of parameres, mushroom-
shaped apodemes well-developed. Female gonapophyses as in Fig. 2e, dorsal valvula

PSOCOPTERA OF ST HELENA AND ASCENSION 429

reduced to a short rudiment, external valvula suboval, relatively broad, setose, with
two stouter setae about in middle and relatively short apical hairs. Region of sperma-
pore characteristic (Fig. 2g). Both spermathecal parietal glands similar in size, with
numerous pores and a central rosette of papillae (Fig. 2f). Spermatheca containing one
spermatophore with a very long channel, similar to that of Cerobasis annulata figured
by Lienhard (1998: fig. 21f).

MEASUREMENTS (y m): Male holotype: BL = 1370; FW = 180; F = 300; T = 460;
ti= 168; t2 = 52; t3 = 60; length of phallosome = 250. — Female allotype: BL = 1510;
RU = 390 = 560: t1= 180; 2 = 56; 13. = 65.

ETYMOLOGY: The specific epithet refers to the distribution of the species on the
Atlantic island of St Helena.

DISCUSSION: Cerobasis atlantica seems to be endemic to the island of St Helena
and has so far been recorded only from Prosperous Bay Plain, an arid habitat. It
belongs to a group of species close to the widespread Cerobasis annulata, most of
which are endemics of one or several Macaronesian islands (see Lienhard, 1984, 1998,
2004b and Lienhard & Smithers, 2002). The new species is easy to distinguish from
C. annulata, which has also been recorded from St Helena (Badonnel, 1976), by its
reduced pigmentation, its more elongate winglets, the presence of 4 terminal spurs on
hindtibia (3 in C. annulata), the absence of a sclerotized longitudinal internal structure
near the apex of the paramere, the structure of the spermapore region and the broad
suboval external valvula (for C. annulata see description by Badonnel, 1976 and
Lienhard, 1998). The structure of the phallosome of C. atlantica 1s similar to that of the
Macaronesian species C. harteni Lienhard, 1984 (see Lienhard, 1984: fig. 19), known
from Cabo Verde and the Azores (Lienhard & Smithers, 2002; Lienhard, 2004b).
However, C. harteni is completely apterous, lacking Pearman's organ on hindcoxa,
with only two apical and two internal spurs on hindtibia and with more elongate
external valvulae and a very characteristic spermapore region (see Lienhard, 1984: figs
7 and 18).

Cerobasis guestfalica (Kolbe, 1880)

MATERIAL EXAMINED: BMNH, 1% , Ascension Island, Devil's Cauldron, S7°56' W14°19,
12.vi11.2003, leg. H. Mendel (off Juniperus).

COMMENT: This widespread and sometimes also domestic species is here
recorded for the first time from Ascension Island; it is also known from St Helena
(Badonnel, 1976) and from several other Atlantic islands, as Canaries, Azores,
Bermudas (see Lienhard & Smithers, 2002).

Cerobasis maya Garcia Aldrete, 1991 Fig. 3
Cerobasis maya Garcia Aldrete, 1991: 324; description of male from Mexico.
Cerobasis cf. maya Garcia Aldrete, 1991. — Ashmole & Ashmole, 1997, 2000a. — Lienhard &
Smithers, 2002. — Lienhard, 2004b.
HOLOTYPE (examined): UNAM, & , Mexico, Yucatan Peninsula, Quintana Roo, Cancun,
2.x1.1971, on dead hanging fronds of coconut palm, leg. A. N. Garcia Aldrete.
NEW MATERIAL: MHNG, 16 , Ascension Island, Command Hüll, 17.-21.111.1990, leg. N.
P. & M. J. Ashmole, trapping on lava (grass cover low, small quantities of several different

430 C. LIENHARD & N. P. ASHMOLE

FIG. 3

Cerobasis maya Garcia Aldrete, male holotype (a), males from St Helena (b-g): (a) Phallosome,
apical part. (b) Tip of left lacinia. (c) Tip of right lacinia (same specimen). (d) Maxillary palp
(pilosity not shown, except for spur sensillum of P2). (e) Right mesothoracic winglet with
insertion points of setae. (f) Phallosome (closed position). (g) Half of phallosome (open position,
axis of symmetry indicated by broken line).

lichens) (sample 0008 Asc). - MHNG, 24, Ascension Island, South Gannet Flow, 23.-
27.11.1990, leg. N. P. & M. J. Ashmole, trapping off-lava (thistles, no moss or lichen) (sample
0818 Asc).-—BMNH, 14 , Ascension Island, South Gannet Hill, S7°58' W14°23', 4.viii.2003, leg.
H. Mendel (litter, extracted by Winkler apparatus).

PSOCOPTERA OF ST HELENA AND ASCENSION 43]

DESCRIPTION OF MALE FROM ASCENSION ISLAND: Colouration: Not very well
preserved. Body yellowish, frons medially with an approximately anchor-shaped
brown patch, sometimes subdivided into smaller spots, several other brown spots on
head, thorax and abdomen, laterally often fused to form larger patches or bands,
winglets hyaline or with some brown pigment (as figured by Garcia Aldrete, 1991:
fig. 2). Compound eyes black, basal flagellomeres apically brown (distal half of
antennae lost in all specimens examined). Femora with some redbrown hypodermal
pigment towards apex, tibiae with two transversal rings of redbrown hypodermal
pigment.

Morphology: Maxillary palp as in Fig. 3d, P4 lacking forked sensillum. Lacinial
tip as shown in Fig. 3b, c (usually both laciniae of same shape, corresponding to Fig.
3c). Forewing reduced to a short winglet bearing 2-3 stout setae in addition to the
normal pilosity (Fig. 3e), hindwing absent. Winglets laterally clearly protruding from
mesothorax in dorsal view (as figured for the holotype by Garcia Aldrete, 1991: fig. 2),
covering only the lateral corners of metanotum, therefore pilosity covering almost all
the metanotum, except for 1/6 of its dorsal surface near lateral margin (Note: in many
other Cerobasis species lateral parts of metanotum extensively covered by winglets
and each lateral 1/3 of its dorsal surface lacking pilosity; see Lienhard, 1998: fig. 24c).
Mesonotum relatively long (its length about equal to half width of vertex), its hind-
margin almost straight in dorsal view. Pearman's organ of hindcoxa well-developed,
hindtibia with 4 terminal spurs and 2 internal spurs in apical half (3 internal spurs in
one of 8 hindtibiae examined). Pretarsal claws lacking preapical tooth, with basal
appendix and slightly enlarged membranous pulvillus. Hypandrial brush with about
60-90 acuminate or slightly truncate setae. Phallosome as in Fig. 3f and Fig. 3g, weakly
sclerotized, mushroom-shaped apodemes well-developed, parameres internally with a
short pointed process; shape of this process different in closed (Fig. 3f) and open
(Fig. 3g) position of the phallosome (see Discussion below).

MEASUREMENTS (um): Male holotype (data from Garcia Aldrete, 1991, except
for length of phallosome): FW = 113; F = 287; T = 486; t1= 184; t2 = 50; t3 = 58;
length of phallosome = 180. — Male from Ascension Island (MHANG 7146): BL = 1260;
EN RE UF — 2460 11= 172; 2 = 47; 55 = 57; lensth of phallosome = 165:

DISCUSSION: The above mentioned material collected by Philip and Myrtle
Ashmole has already been mentioned by these authors as Cerobasis cf. maya (Ashmole
& Ashmole, 1997, 2000; see also Lienhard & Smithers, 2002 and Lienhard, 2004b).
AII males from Ascension Island are so similar to the only previously known specimen
of C. maya, its holotype, that there is no reason to consider them as belonging to a
different species or subspecies in spite of some slight differences concerning phallo -
some morphology. The whole phallosome of the holotype 1s figured by Garcia Aldrete
(1991: fig. 4), details of its distal part are also represented in Fig. 3a (mushroom-
shaped apodemes of the parameres not shown in this figure). The comparison with Fig.
3f, g, representing the phallosome of two males from Ascension Island in closed and
open position, shows the variable aspects of the apical structures depending on its
position after slide-mounting. The position of these parts in Fig. 3f 1s rather similar to
that observed in the slide of the holotype (Fig. 3a). A careful analysis of these stuctures

432 C. LIENHARD & N. P. ASHMOLE

in all available males showed that the differences between Fig. 3a and Fig. 3f are
largely due to slightly different positions after slide-mounting. The only significant
difference between the holotype and the males from Ascension Island is the presence,
in the holotype, of a small field of scale-like sculpture on the parameres near the base
of the internal process (Fig. 3a and Garcfa Aldrete, 1991: fig. 4); this sculpture is absent
or only very weakly developed in the males from Ascension Island (Fig. 3f, g).
However, compared to usual interspecific differences in phallosome morphology in the
genus Cerobasis, this extremely slight difference does not justify any taxonomic
decision about specific or subspecific separation of the Ascension population,
especially in view of the low numbers of individuals available at present.

According to Garcia Aldrete (1991) this species seems to be more closely
related to some Macaronesian species than to the other known Mexican species of the
genus Cerobasis; he mentions the possibility that C. maya may have been introduced
to Yucatan Pensinsula from the Caribbean. Ashmole & Ashmole (1997) tentatively
suggested that this species was native to Ascension, but pointed out that it provides one
of the rare examples of apparent New World affinities in the Ascension arthropod fauna
(see also Biogeographical discussion, below).

Lepinotus inquilinus Heyden, 1850

MATERIAL EXAMINED: MHNG, 29, St Helena, Woodcot, S15°57.2' W5°42.7', ca 489m,
18.11.2006, leg. P. & M. Ashmole, off laboratory table (sample 2619).

COMMENT: This cosmopolitan and usually domestic species (see Lienhard &
Smithers, 2002) is here recorded for the first time from St Helena. As in the case of
Liposcelis entomophila, this species is undoubtedly introduced to this island (Mendel
et al., 2008).

LIPOSCELIDIDAE

Liposcelis entomophila (Enderlein, 1907)

MATERIAL EXAMINED: MHNG, 2%, St Helena, Rupert's Battery Cave, ca SOm,
ra leg. P. & M. Ashmole (sample 4444) and 4.x1.2006, leg. E. Thorpe (sample

COMMENT: This cosmopolitan and often domestic species (see Lienhard &
Smithers, 2002) is here recorded for the first time from St Helena. As Lepinotus inqui-
linus (see Mendel ef al., 2008) and Liposcelis bostrychophila (Fig. 4b) it is undoub-
tedly introduced to this island. The latter species was erroneously listed as a St Helena
endemic by Mendel ef al. (2008).

Troglotroctes ashmoleorum Lienhard, 1996 Fig. 4a
Troglotroctes ashmoleorum Lienhard, 1996: 118; description of both sexes from Ascension
Island.

TYPE MATERIAL: MHNG and NMSE (see Lienhard, 1996).

NEW MATERIAL: MHNG and BMNH, 14, 21% (most of them heavily damaged),
Ascension Island, South Gannet Hill, 17.-30.v.1995, leg. N. P. & M. J. Ashmole, pipe trap
inserted vertically ca 2m into lava rubble (sample 0620). See photographs of trap and biotope in
Asmole & Ashmole (2000b: figs 14.5 and 14.6).

PSOCOPTERA OF ST HELENA AND ASCENSION 433

FIG. 4

Troglotroctes ashmoleorum Lienhard (a) and Liposcelis bostrychophila Badonnel (b), females,
to same scale (scale bar: 0.5 mm). The comparison of 7. ashmoleorum With the habitus of this
typical Liposcelis species makes evident the troglomorphic habitus of the former (1. e. relatively
long legs and antennae, strongly reduced eyes and weakly developed pigmentation).

DISCUSSION: Ashmole & Ashmole (1997, 2002b) already incidentally men -
tioned this new record, without giving detailed collecting data. This troglomorphic
species (Fig. 4a) was originally found in caves (Lienhard, 1996), but the above
mentioned individuals were caught in a pipe trap in barren volcanic rubble, suggesting
that it is widespread underground. However, a single paratype female was also trapped

434 C. LIENHARD & N. P. ASHMOLE

in a crevice on barren lava in 1990, so that individuals must sometimes come to the
surface, probably at night (Asmole & Ashmole, 2000b).

T. ashmoleorum 1s closely related to the genus Liposcelis (see Lienhard, 1996
and Fig. 4a, b), probably phylogenetically embedded within this large genus (Grimaldi
& Engel, 2006); therefore the validity of the monotypic genus Troglotroctes Lienhard
has only provisionally been maintained by Yoshizawa & Lienhard (2010).

SPHAEROPSOCIDAE

Sphaeropsocopsis insularum Lienhard sp. n. Fig. 5

Sphaeropsocopsis cf. microps Badonnel, 1963 (one damaged female from Ascension Island, see
paratype below). — Ashmole & Ashmole, 1997, 2002a. — Lienhard & Smithers, 2002;
Lienhard, 2004b.

HOLOTYPE: MHNG, ©, St Helena, Rupert's Battery Cave, ca 50m, 25.xi-10.xi.2003,
leg. N. P. & M. J. Ashmole, modified pitfall trap ("boot trap") (sample 1835).

PARATYPES: BMNH, 19 , St Helena, Rupert's Battery Cave, ca 50m, 16.ii.2006, leg. N.
P. & M. J. Ashmole, modified pitfall trap ("boot trap") (sample 2805).- MHNG, 19 , Ascension
Island, Lower Valley Crater, near Northeast Bay, 15-23.v.1995, leg. N. P. & M. J. Ashmole, pipe
trap inserted vertically ca 2m into cinders (sample 1107). See photograph of pipe trap in
Ashmole & Ashmole (2002b: fig. 14.5).

DESCRIPTION OF FEMALE (male unknown): Head light brown, rest of body
whitish brown to yellowish. Hindwings absent, forewings lost in all specimens, their
lunulate insertion points visible dorso-laterally near posterior margin of mesothorax
(Fig. 5d). Eyes with 3 ommatidia, ocelli absent, frontal suture not visible, vertical
suture well-developed, vertex clearly notched in middle (Fig. 5a). Sculpture of vertex
consisting of small simple tubercles, mostly arranged into clearly delimited polygonal
or scale-shaped areoles (diameter of tubercles slightly smaller than diameter of the
alveoli of the small vertical hairs). Antenna with 15 articles (mostly damaged, flagellar
sense clubs not observed). Maxillary palp as in Fig. 5a, P4 elongate fusiform, its sub-
apical sensory field with 2 long proximal setiform sensilla and 3 similar but somewhat
shorter distal sensilla, the latter surrounding 3 club sensilla, two of them slender, the
proximal one short and thick, almost spherical (Fig. 5b, c). Labial palp with 3 thin-
walled sensilla, the lateral one differentiated as a thick short sense club (Fig. 5i).
Lacinial tip bifurcate, inner tine shorter than outer tine, the latter with two small
secondary denticles on 1ts inner side (similar to S. myrtleae, see Lienhard & Ashmole,
1999: fig. 5). Sculpture of thoracic tergites similar to that of vertex but areoles absent
or indistinct. Mesonotum not subdivided into lobes (Fig. Sd). Legs slender but not
particularly long (index T/V = 1.2), no coxal organ present, tibiae with two apical
spurs, pretarsal claws with a minute preapical denticle and some ventral microtrichia.
Epiproct and paraprocts simple (Fig. 5h). Subgenital plate broader than long, flattened
on distal margin, T-shaped sclerite well-developed (Fig. Se). Gonapophyses typical for
the family (see Mockford, 2009), apex of external valvula broadly rounded, not bilobed
(Fig. 5g). Region of spermapore with a weakly developed field of small microtrichia
(Fig. 5f), spermathecal duct and sac not observed. Measurements (holotype, mm): BL
OSDIV = 601 tre 25 = MMA DES 265;

ETYMOLOGY: The specific epithet, a feminine noun in apposition, refers to the
distribution on two South Atlantic islands (island = lat. insula; genitive plural: insu -
larum).

PSOCOPTERA OF ST HELENA AND ASCENSION 435

né nee
NS NA à
AT EE de z si

Li Fos

FIG. 5

Sphaeropsocopsis insularum Lienhard sp. n., female holotype (c-d, g-1), female paratypes (a-b,
from Ascension Island; e-f, from St Helena): (a) Head, frontal view (pilosity of antennae and
maxillary palps not shown). (b) P4 of maxillary palp (pilosity not shown, except for subapical
sensory field of thin-walled sensilla). (c) Ditto (other specimen and different position). (d)
Dorsal view of mesothorax (with insertion points of forewings) and anterior part of wingless
metathorax. (e) Subgenital plate. (f) Spermapore region. (g) Gonapophyses. (h) Epiproct and
right paraproct. (i) Labial palp (pilosity not shown, except for thin-walled sensilla).

436 C. LIENHARD & N. P. ASHMOLE

DISCUSSION: The species was previously mentioned from Ascension Island by
Ashmole & Ashmole (1997, 2002a) as Sphaeropsocopsis cf. microps, based on the
tentative identification of the damaged paratype female (see above).This specimen had
eyes with three ommatidia, a character only known before from the Chilean species S.
microps. However, this female and the two new females from St Helena, undoubtedly
belonging to the same species, clearly differ from the female of S. microps by the
cuticular sculpture on vertex. In the latter species 1t consists of large irregularly lobate
tubercles which are not arranged into areoles (Badonnel, 1963). Unfortunately fore-
wings are broken and lost in all available specimens of S. insularum; however, the
insertion point of the forewing (Fig. Sd) is similar to that figured by Badonnel (1963:
fig. 59) for S. chilensis Badonnel. Thus, the new species is not apterous but has pro-
bably elytriform forewings similar to those of S. microps and S. chilensis (see
Badonnel, 1963). The presence of a nearly spherical club sensillum in the subapical P4
sensory field seems to be characteristic of S. insularum; the corresponding sensillum
of the only known African species of the genus, S. reisi Badonnel, is also thick, but
much longer (Badonnel, 1971: fig. 5). Without information on morphology of forewing
and male genitalia it 1s impossible to decide if S. insularum is more closely related to
this African species, as 1s S. myrtleae, the second St Helena species of the genus (see
below), or to S. microps, which 1s only known from natural edaphic habitats in Chile
(see Badonnel, 1963, 1967). S. reisi has eyes with 9 ommatidia (see Badonnel, 1971).
However, eye reduction observed in S. insularum (3 ommatidia) and S. microps (3-4
ommatidia, occasionally 5; see Badonnel, 1963, 1967 and Mockford, 2009) is likely to
be due to convergence.

Sphaeropsocopsis myrtleae Lienhard & Ashmole, 1999 Fig. 6
Sphaeropsocopsis myrtleae Lienhard & Ashmole, 1999: 907: description of female from St
Helena.

HOLOTYPE: MHNG, ? , St Helena, Rupert's Battery Cave, 13-17.ii1.1995, leg. N. P. & M.
J. Ashmole, modified pitfall trap ("boot trap") (sample 680 SH).

NEW MATERIAL: MANG, 16 19 , and BMNH, 1 ©, St Helena, Rupert's Battery Cave, ca
50m, 25.x1-10.x11.2003, leg. N. P. & M. J. Ashmole, modified pitfall trap ("boot trap") (sample
1835).

DESCRIPTION OF MALE: Body and appendages white to yellowish, head capsule
very light brown, only sclerotized parts of mandibles dark brown. General morphology
as in female (see Lienhard & Ashmole, 1999 and Discussion below) but almost
apterous, only small lobes of rudimentary forewings postero-laterally on mesothorax
(Fig. 6f), hindwings absent. Maxillary palps lacking (broken), both antennae damaged.
Large lateral sense club on labial palp as in female (see Fig. 6d). Legs relatively long
(index T/V = 1.5). Epiproct, paraprocts and hypandrium simple. Phallosome as shown
in Fig. 6e. Measurements (4 MHNG 7626, um): BL = 1220; V = 250; F+tr = 304;
LEe385;11=150 2 =39 62-56.

DISCUSSION: Based on two of the three females known at present, the figure of
the head (Fig. 6b) could be completed (see Lienhard & Ashmole, 1999: fig. 1, llacking
maxillary palps and some setae) and figures of the sensilla of labial and maxillary palps
could be made (Fig. 6c, d). Initially, the absence of subdivision of mesonotum into
lobes was the main reason to assign the species to the genus Sphaeropsocopsis

PSOCOPTERA OF ST HELENA AND ASCENSION 431

FIG. 6

Sphaeropsocopsis myrtleae Lienhard & Ashmole: (a) Habitus of female, dorsal view, antennae
incomplete (scale bar: 0.5 mm). (b) Head, frontal view (reconstruction based on two slightly
damaged females; pilosity of antennae and maxillary palps not shown). (c) P4 of maxillary palp,
female (pilosity not shown, except for subapical sensory field of thin-walled sensilla). (d) Labial
palp, female (pilosity not shown, except for thin-walled sensilla). (e) Phallosome. (f) Dorsal
view of mesothorax with forewing rudiments, male.

438 C. LIENHARD & N. P. ASHMOLE

Badonnel and not to Badonnelia Pearman (see Lienhard & Ashmole, 1999; erroneously
mentioned as "mesothoracic sternites" in the Discussion on p. 909). The presence of an
elongate fusiform P4 now confirms this assignment (P4 subcylindrical in Badonnelia:;
see Badonnel, 1963). The newly discovered male also confirms the initial generic
assignment and allows a better understanding of the origin of this island endemic. The
morphology of the phallosome of S. myrtleae is very similar to that of the African
species S. reisi Badonnel, known from Angola (see Badonnel, 1971: fig. 2). However,
in S. reisi both sexes have well-developed pigmentation and hemispherically pro -
minent eyes of 9 ommatidia. The female of S. reisi has elytriform, vaulted forewings
reaching the tip of the abdomen and slightly enveloping it laterally (Badonnel, 1971);
each forewing bears four longitudinal veins. This type of elytriform forewing is
characteristic for the family Sphaeropsocidae (Mockford, 2009). In S. myrtleae the
forewings are reduced to short narrow membranous flaps, bearing only two longitu -
dinal veins (Fig. 6a). The cave-dwelling S. myrtleae is the only blind (anophthalmic)
psocid species known at present. Even in forms with reduced compound eyes, as
certain species of Liposcelis, at least two ommatidia are always present (Lienhard,
1998), except for the troglobitic Speleopsocus chimanta Lienhard (Prionoglarididae),
recently discovered in a Venezuelan cave, which has only one minute ommatidium on
each side of the head (Lienhard ef al., 2010). The epigaeic African ancestor of S.
myrtleae probably colonised St Helena by air and became adapted to subterranean life
after reaching the island (Lienhard & Ashmole, 1999: see also Biogeographical
discussion, below).

CAECILIUSIDAE

Stenocaecilius caboverdensis (Meinander, 1966)

MATERIAL EXAMINED: BMNH, 1 % , St Helena, Cuckhold's Point, S15°58' W5°42',771m,
x11.2005-1.2006, leg. H. Mendel (Malaise trap). - BMNH, 1% (damaged, lacking abdomen),
Ascension Island, Grazing Valley, S7°57' W14°21', vii.2003, leg. H. Mendel (pitfall trap). —
BMNH,19 ,Ascension Island, Green Mt., S7°57' W14°21", 6.viii.2003, leg. H. Mendel (vacuum
sampler).

COMMENT: This atlanto-mediterranean species is here recorded for the first time
from St Helena and Ascension Island. According to Lienhard & Smithers (2002) it is
already known from three Macaronesian archipelagoes (Cabo Verde, Azores, Madeira)
and from several mediterranean countries (Cyprus, Greece, Israel, Portugal and
Tunisia).

PERIPSOCIDAE

Peripsocus leleupi Badonnel, 1976

MATERIAL EXAMINED: BMNH and MHNG, 59, Ascension Island, Devil's Cauldron,
S7°56 W14°19", 12.vi1.2003, leg. H. Mendel (off Juniperus).

COMMENT: This species was previously considered as a St Helena endemic
(Ashmole & Ashmole, 2000); it is closely related to the African species Peripsocus
ghesquierei Badonnel (Badonnel, 1976; for distribution see Lienhard & Smithers,
2002).

PSOCOPTERA OF ST HELENA AND ASCENSION 439

Peripsocus pauliani Badonnel, 1949

MATERIAL EXAMINED: BMNH and MHNG, 6%, Ascension Island, Devil's Cauldron,
S7°56' W14°19', 12.vin1.2003, leg. H. Mendel (off Juniperus).

COMMENT: This widespread pan-tropical waïf (see Lienhard & Smithers, 2002)
is here recorded for the first time from Ascension Island; it is also known from St
Helena (Badonnel, 1976).

PSOCIDAE

Blaste helenae Lienhard sp. n. Fig. 7a-f

HOLOTYPE: MHNG, à , St Helena, Lot Summit, 454m, off St Helena rosemary Phylica
polifolia, 29 1.2006, leg. P. & M. Ashmole (sample 4165).

PARATYPES: BMNH 16 ,MHNG 1% (allotype) and 1 nymph, same data as for holotype.

DESCRIPTION: General colouration and morphology of both sexes very similar
to that described by Badonnel (1976) for Blaste basilewskyi, except for less extensive
forewing markings (Fig. 7a) and the following characteristics of genital morphology.
Hypandrium with a median pair of short but slender terminal processes (Fig. 7d);
phallosome on each side with a short outwards-curved hook (Fig. 7c). Subgenital plate
of female with a widely opened V-shaped sclerotization, arms of the V relatively
slender (Fig. 7f); sclerotization of spermapore region as in Fig. 7e.

MEASUREMENTS: Male holotype: BL = 2.2 mm, IO/D = 2.1; FW = 2.5 mm; F =
440 yum; T = 830 pm; t1= 240 pm; t2 = 150 pm. — Female allotype: BL = 2.4 mm;
IO/D = 2.6; FW = 2.5 mm, F = 410 pm; T = 780 pm; t1= 210 pm; t2 = 140 pm.

ETYMOLOGY: The specific epithet refers to the island of St Helena, a British
overseas territory in the South Atlantic Ocean, which is named after Saint Helena of
Constantinople.

DISCUSSION: This new species is closely related to Blaste basilewskyi, the
second species of this genus known from St Helena (junior synonym: Blaste atlantica
New, 1977: 255; see Lienhard & Smithers, 2002: 379). Both species can be assigned
to the subgenus Euclismia Enderlein (see Badonnel, 1976). B. helenae clearly differs
from B. basilewskyi by less extensive forewing markings in both sexes (see figures of
B. basilewskyi given by Badonnel, 1976), by smaller hooks of the phallosome (see Fig.
7h and Badonnel, 1976: fig. 173) and by slightly longer and more slender median pair
of terminal processes of hypandrium (see Fig. 7g and Badonnel, 1976: fig. 172).
Female genitalia of both species are almost identical, except for slight differences
concerning the sclerotized area surrounding the spermapore (see Fig. 71 and Badonnel,
1976: figs 174, 175). Measurements of B. basilewskyi are rather variable (Badonnel,
1976), those given here for B. helenae are close to the lowermost values observed by
this author for B. basilewskyi, sometimes even lower.

The unique systematic position of Blaste basilewskyi within the genus (see
Badonnel, 1976; New, 1977) gives no hint on the possible origin of this species, which
is common on St Helena and is associated mainly with the endemic gumwoods
Commidendrum spp., although it has also been found on other plants (Badonnel, 1976
and personal observations). The discovery of Blaste helenae on the endemic St Helena
rosemary Phylica polifolia suggests a niche separation between the two species. B.

440 C. LIENHARD & N. P. ASHMOLE

FIG. 7

Blaste helenae Lienhard sp. n. (a-d, male holotype; e-f, female allotype): (a) Forewing. (b)
Hindwing. (c) Phallosome. (d) Hypandrium, ventral view. (e) Spermapore region. (f) Subgenital

plate. — Blaste basilewkyi Badonnel (g-i): (g) Hypandrium, ventral view. (h) Phallosome. (i)
Spermapore region.

PSOCOPTERA OF ST HELENA AND ASCENSION 441]

helenae Was beaten off this plant during the first collecting of invertebrates on the
summit of Lot, a massive intrusion of phonolitic rock, exposed by erosion, which
forms a striking feature of the landscape of Sandy Bay. Its nearly vertical sides are
almost devoid of vegetation but the summit — less than one tenth of a hectare in extent
— provides a refuge for a few specimens of Phylica polifolia. The rosemary was
formerly widespread in dry places in the west of the island but has now been almost
entirely lost as a wild tree (Cronk, 2000). Further investigation is needed to determine
whether B. helenae is also present on any of the other surviving specimens. Badonnel
(1976) suggested that Blaste basilewskyi may be derived from one of the earliest
colonizers of St Helena. Therefore the existence of the apparently much rarer but
extremely closely related sister-species Blaste helenae on the same island is here
tentatively interpreted as a result of sympatric speciation, possibly resulting from adap-
tation to life on different endemic plants, rather than of double invasion.

Indiopsocus mendeli Lienhard sp. n. Figs 8-9

HOLOTYPE: MHNG, GG, Ascension Island, Devil's Cauldron, S7°56' W14°19",
12.viii.2003, leg. H. Mendel (off Juniperus bermudiana).

PARATYPES: BMNH, MHNG and SEHU, 194, 109 (one of them allotype, MHNG
8018), same data as for holotype. - BMNH and MHNG, 116,89, Ascension Island, Mt Red
Hill, S7°58' W14°21', 14.viii.2003, leg. H. Mendel (off Zecoma stans).-— BMNH, 14 , Ascension
Island, Mt Red Hill, S7°58' W14°21', 14.viii.2003, leg. H. Mendel. - BMNH, 19, Ascension
Island, Devil's Ashpit, S7°57' W14°13", 2.vi1.2003, leg. H. Mendel (vacuum sampler).

DESCRIPTION: Colouration: Head and thorax pale brown, with dark brown
markings, postclypeus with brown longitudinal stripes, antenna brown, compound eye
black, legs yellowish to medium brown. Maxillary palp with PI and P2 very light
brown, P3 and P4 darker brown, apical half of P4 blackish brown. Forewing pattern
shightly sexually dimorphic, with more extensive brown markings in female (Fig. 8f)
than in male (Fig. 8a), especially in basal half of the wing. Abdomen whitish, with
some red-brown hypodermal pigment, in particular laterally, terminalia dark brown. In
male, membranous zone anteriorly to hypandrium on each side with a brown sclero-
tized patch (Fig. 9a), sometimes not very distinct.

Morphology: Compound eyes very large and prominent in male, distinctly
smaller in female (see I0/D values, below), ocelli well-developed. Wing venation as in
Fig. 8a, b, f; posterior apex of pterostigma with a very short and sometimes weakly
developed spur vein.

Male terminalia (Fig. 9): Posterior margin of clunium medially slhightly pro-
minent and overlapping base of epiproct (Fig. 9d), the latter hemicircular, basally on
each side with a well-sclerotized swelling (Fig. 9a, d). Paraproct with a short lateral
protuberance, a pointed apical process and a small setose protuberance basally of the
latter; trichobothria forming an arched, posteriorly open sense cushion (Fig. 9a, d).
Medio-distal protuberance of hypandrium slightly asymmetrical, subdivided into a few
pustulate or denticulate lobes, partly weakly but mostly heavily sclerotized (Fig. 9a, c).
Phallosome basally broad and truncate, distally with a pair of lateral lobiform and
weakly sclerotized processes and three well-sclerotized subacute apical processes of
almost equal length, separated by two narrow V-shaped indentations (Fig. 9b).

Female terminalia: Posterior margin of clunium straight, epiproct and para-
procts simple, as usual in the family. Subgenital plate as in Fig. 8d, basal sclerotization

442 C. LIENHARD & N. P. ASHMOLE

FIG. 8

Indiopsocus mendeli Lienhard sp. n.: (a) Forewing, male (scale bar: 1 mm). (b) Hindwing, male
(to same scale). (c) Gonapophyses, female. (d) Subgenital plate, female. (e) Spermapore region,
female. (f) Forewing, female (to same scale as Fig. 8a, b).

V-shaped, median zone of apical lobe particularly well-sclerotized on each side.
Gonapophyses and region of spermapore as in Fig. 8c, e; posterior lobe of external
valvula inconspicuous.

PSOCOPTERA OF ST HELENA AND ASCENSION 443

F1G. 9

Indiopsocus mendeli Lienhard sp. n., male: (a) Abdominal apex, lateral view (pilosity not shown,
except for paraproctal trichobothria). (b) Phallosome. (c) Hypandrium, posterior view (slightly
squashed). (d) Clunium, epiproct and paraprocts (dorsal view, pilosity partially omitted, para-
procts in different position).

444 C. LIENHARD & N. P. ASHMOLE

MEASUREMENTS: Male holotype: BL = 2.0 mm; IO/D = 0.9; FW = 2.75 mm; F
= 590 um; T = 1200 y m; t1= 436 pm; t2 = 143 pm. — Female allotype: BL = 2.1 mm;
IO/D = 1.7; FW = 2.95 mm; F = 590 pm; T = 1240 y m; t1= 414 pm; t2 = 144 ym.

ETYMOLOGY: The specific epithet refers to the collector of the type material,
Howard Mendel (BMNH), in recognition of his important contributions to scientific
study of St Helena and Ascension Island fauna.

DISCUSSION: The new species is closely related to /ndiopsocus dentatus
(Thornton & Woo, 1973), which 1s only known from the Galapagos Islands (Thornton
& Woo, 1973; Lienhard & Smithers, 2002). The forewing pattern of the female of
I. dentatus (see Thornton & Woo, 1973: fig. 77) 1s very similar to that figured here for
the male of /. mendeli (Fig. 8a), while in the female of the latter some additional dark
markings are usually visible in the basal half of the wing (Fig. 8f). Female genitalic
characters of both species are very similar (see Thornton & Woo, 1973: figs 78, 79; the
spermapore region of 1. dentatus is not known). However, these species are easy to dis-
tinguish by the different shape of the apical lobes of the hypandrium and especially of
the terminal processes of the phallosome (see Thornton & Woo, 1973: figs 80-82); in
particular, the two deep indentations between the three mediodistal processes of the
phallosome are broadly U-shaped in /. dentatus.

After the above treated Cerobasis maya this 1s the second example of apparent
New World affinities in the Psocoptera fauna of Ascension Island (see also Biogeo-
graphical discussion).

BIOGEOGRAPHICAL DISCUSSION

The psocopteran fauna of St Helena and Ascension Island is now fairly well
known (St Helena: 23 spp.; Ascension: 13 spp.; see checklist in Appendix and
comments in Introduction). Detailed data (up to the year 2000) on the general distri-
bution of the non-endemic species can be found in Lienhard & Smithers (2002) and
more recent additional data in Lienhard (2003-2011, in Psocid News).

Several species have probably been introduced by human activities, such as the
following widespread (in some cases cosmopolitan) and often domestic species of the
suborders Trogiomorpha and Troctomorpha: Cerobasis annulata (St Helena), C. guest-
Jalica (St Helena, Ascension), Lepinotus inquilinus (St Helena), Psocathropos lachlani
(Ascension), Psyllipsocus ramburii (St Helena, Ascension), Liposcelis bostrychophila
(St Helena, Ascension), L. entomophila (St Helena). The cosmopolitan and sometimes
domestic Psocomorpha species Ectopsocus briggsi (St Helena) belongs probably also
to this category. Furthermore, the recently discovered species Helenatropos abrupta,
initially described as endemic to St Helena but later also found in Table Mountain
National Park, Cape Town, was probably also introduced to the island from South
Africa (see Taxonomic treatment).

One widespread tropical waif, Peripsocus pauliani (St Helena, Ascension), and
the following predominantly Western Palaearctic species with atlanto-mediterranean
distribution, known also from Macaronesian archipelagoes, have probably reached
these South Atlantic islands by natural dispersal: Stenocaecilius caboverdensis (St
Helena, Ascension), Ectopsocus strauchi (St Helena, Ascension), Trichopsocus clarus

PSOCOPTERA OF ST HELENA AND ASCENSION 445

(St Helena), Myopsocus eatoni (St Helena). The case of the Mexican species Cerobasis
maya, at present only known from Ascension Island and the Yucatan Peninsula, is more
puzzling, in view of the unfavourable conditions for trans-Atlantic dispersal of insects
from west to east (Ashmole & Ashmole, 1997); anthropogenic distribution due to intro-
duced plants or phoretic dispersal by seabirds cannot be excluded (some Cerobasis
species are known to live occasionally in birds' nests, e. g. C. guestfalica, see Lienhard,
1986).

Two species occur on both islands, St Helena and Ascension, without being
known from elsewhere: Sphaeropsocopsis insularum Lienhard sp. n. and Peripsocus
leleupi. This could be due to independent invasion from the same origin (Africa for P.
leleupi, unknown for S. insularum; see Taxonomic treatment) or to natural dispersal or
human-assisted transfer from one island to the other.

The following six species can be considered as St Helena endemics (see
Ashmole & Ashmole, 2000a and Mendel et al., 2008, but note that the cosmopolitan
and often domestic species Liposcelis bostrychophila Was accidentally listed as an
endemic in the 2008 report, and that the species Peripsocus leleupi has now also been
recorded on Ascension Island, see above): Cerobasis atlantica Lienhard sp. n.,
Sphaeropsocopsis myrtleae, Stenocaecilius benoiti, Peripsocus decellei, Blaste basi-
lewskyi and Blaste helenae Lienhard sp. n. Cerobasis atlantica belongs to a species
group containing several Macaronesian endemics (see description, above), while
Sphaeropsocopsis myrtleae, Stenocaecilius benoiti and Peripsocus decellei are related
to African species (see Badonnel, 1976 and Taxonomic treatment, above). The exis-
tence of two very closely related endemic sister-species of unknown origin, Blaste
basilewskyi and B. helenae, is here tentatively interpreted as a result of sympatric
speciation (see description of B. helenae, above).

Two species can be considered as endemics of Ascension Island, 7roglotroctes
ashmoleorum and Indiopsocus mendeli Lienhard sp. n. The latter is closely related to
I. dentatus, only known from Galapagos islands (see description, above); together with
Cerobasis maya it provides one of the few examples of apparent New World affinities
in the Ascension and St Helena arthropod fauna (Ashmole & Ashmole, 1997). As
mentioned above for C. maya, an anthropogenic introduction of 7. mendeli cannot be
excluded. The plants on which the species has been found on Ascension Island
(Juniperus bermudiana and Tecoma stans) are widely planted and sometimes invasive
in many parts of the Pacific. Thus /. mendeli is clearly a species associated with intro-
duced plants.

The subterranean and troglomorphic species Troglotroctes ashmoleorum
(Ascension, F1g. 4a) and Sphaeropsocopsis myrtleae (St Helena, Fig. 6a) are among the
most interesting endemic arthropods of these South Atlantic islands, being the only
known members of the suborder Troctomorpha with clear morphological adaptations
to subterranean life. The adaptations presumably evolved after arrival of the ancestral
forms respectively on Ascension Island and St Helena. The case of 7roglotroctes 1s
especially interesting in view of the relatively recent origin of Ascension Island
(around one million years ago). This species is one of a small but taxonomically
diverse group of arthropods on that island which demonstrate the relatively rapid evo-
lution of troglomorphic characteristics (Ashmole & Ashmole, 1997: 570).

446 C. LIENHARD & N. P. ASHMOLE

Sphaeropsocopsis myrtleae, from the much older island of St Helena, may be
derived from a lineage that reached it in the distant past. This species is especially
significant since it provides almost the only piece of evidence of the existence on this
island of a highly adapted subterranean fauna (Lienhard & Ashmole, 1999). Ancient
volcanic terrain tends to lack specialised subterranean fauna because weathering of
volcanic habitats over long periods leads to the silting up of cracks and superficial
underground spaces, preventing the inflow of nutrients to deeper layers and ultimately
sealing caves, so that they may become sterile. Nearer the surface, the formation of soil
and growth of vegetation leads to faunal succession (Ashmole ef al., 1992) and any
caves where life might persist tend to be inaccessible. On St Helena this process has
led to dominance of introduced species in the soil and the remaining subsurface spaces
(Ashmole & Ashmole, 2000a: 131-132). Rupert’s Battery Cave, where S. myrtleae was
found, was the only lava tube to which we could gain access sufficient to sample even
the most superficial parts of a cave environment.

ACKNOWLEDGEMENTS

We are grateful to Howard Mendel (BMNH), Mike Picker and Charmaine Uys
(University of Cape Town, South Africa) for entrusting their specimens to C. Lienhard
and for depositing many of them in the Psocoptera collections of the MHNG. Howard
Mendel and Alfonso Garcia Aldrete (UNAM) are acknowledged for reading the
manuscript and making valuable suggestions, the latter also for the loan of the type of
Cerobasis maya. Field work by N. P. & M. J. Ashmole on St Helena and Ascension
Island was supported by the Carnegie Trust for the Universities of Scotland, the British
Ecological Society, the U.K. Foreign and Commonwealth Office and the Overseas
Territories Environment Programme. Grateful thanks are also due to all those people
who helped to make research on the islands so productive and enjoyable.

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LIENHARD, C. & SMITHERS, C. N. 2002. Psocoptera (Insecta): World catalogue and bibliography.
Instrumenta Biodiversitatis 5: xli+745 pp. Muséum d'histoire naturelle, Genève.
LIENHARD, C., HOLUSA, O. & GRAFITTI, G. 2010. Two new cave-dwelling Prionoglarididae from
Venezuela and Namibia (Psocodea: ‘Psocoptera': Trogiomorpha). Revue suisse de

Zoologie 117(2): 185-197.

MENDEL, H., ASHMOLE, P. & ASHMOLE, M. 2008. Invertebrates of the Central Peaks and Peak
Dale, St Helena (with additional records from other sites). Based on a survey commis-
sioned by the St Helena National Trust, 122 pp. (Unpublished report).

MOCKFORD, E. L. 2009. Systematics of North American species of Sphaeropsocidae
(Psocoptera). Proceedings of the Entomological Society of Washington 111(3): 666-685.

NEW, T. R. 1977. À new species of Blaste (Psocoptera, Psocidae) from St. Helena. Nouvelle
Revue d'Entomologie 7(3): 253-256.

THORNTON, I. W.B. & Woo, A. K.T. 1973. Psocoptera of the Galapagos Islands. Pacific Insects
15(1): 1-58.

YOSHIZAWA, K. & LIENHARD, C. 2010. In search of the sister group of the true lice: A systematic

review of booklice and their relatives, with an updated checklist of Liposcelididae
(Insecta: Psocodea). Arthropod Systematics & Phylogeny 68(2): 181-195.

APPENDIX: Checklist of Psocoptera species known from St Helena and Ascension Island (arran-
gement of suborders and families according to Lienhard & Smithers, 2002; for species groups of
Liposcelis see Lienhard, 1998; * = island endemic)

ST HELENA ASCENSION ISLAND
TROGIOMORPHA
Trogiidae Cerobasis annulata (Hagen, 1865)
*Cerobasis atlantica Lienhard sp. n.
Cerobasis guestfalica (Kolbe, 1880) Cerobasis guestfalica
Cerobasis maya Garcia Aldrete, 1991
Helenatropos abrupta Lienhard, 2005
Lepinotus inguilinus Heyden, 1850
Psyllipsocidae Psocathropos lachlani Ribaga, 1899
Psyllipsocus ramburii Selys- Psyllipsocus ramburii
Longchamps, 1872
TROCTOMORPHA
Liposcelididae Liposcelis bostrychophila Badonnel, Liposcelis bostrychophila
1931
Liposcelis entomophila (Enderlein,
1907)
Liposcelis spec. (species group A)
Liposcelis spec. (species group ©) Liposcelis spec. (species group C)
*Troglotroctes ashmoleorum Lienh.,
1996
Sphaeropsocidae Sphaeropsocopsis insularum Lienhard Sphaeropsocopsis insularum
Sp. n.

*Sphaeropsocopsis myrtleae Lienh.
& Ashm., 1999

PSOCOMORPHA
Caeciliusidae

Ectopsocidae

Peripsocidae

Trichopsocidae
Psocidae

Myopsocidae

PSOCOPTERA OF ST HELENA AND ASCENSION 449

*Stenocaecilius benoiti (Badonnel,
1976)

Stenocaecilius caboverdensis
(Meinander, 1966)

Ectopsocus briggsi McLachlan, 1899
Ectopsocus strauchi Enderlein, 1906
*Peripsocus decellei Badonnel, 1976
Peripsocus leleupi Badonnel, 1976
Peripsocus pauliani Badonnel, 1949
Trichopsocus clarus (Banks, 1908)
*Blaste basilewskyi Badonnel, 1976
*Blaste helenae Lienhard sp. n.

Myopsocus eatoni McLachlan, 1880

Stenocaecilius caboverdensis
Ectopsocus strauchi

Peripsocus leleupi
Peripsocus pauliani

*Indiopsocus mendeli Lienhard sp. n.

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REVUE SUISSE DE ZOOLOGIE 118 (3): 451-466; septembre 2011

Redescription of the genus Marcenendius Navas
(Psocodea: ‘Psocoptera’: Amphientomidae)
with a key to western Palaearctic amphientomids

Charles LIENHARD! & Arturo BAZ?

l Muséum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland.
E-mail: charleslienhard@bluewin.ch

2 Universidad de Alcalä de Henares, Departamento de Zoologfa y Antropologfa
Fisica, E-28871 Alcalä de Henares (Madrid), Spain.

Redescription of the genus Marcenendius Naväs (Psocodea: ‘Psocop-
tera’: Amphientomidae) with a key to western Palaearctic amphien-
tomids. - Based on recently collected specimens from continental Spain and
Mallorca island the type species of the genus Marcenendius Naväs, 1913,
M. nostras Naväs, 1913, 1s redescribed and tentatively synonymized with
the second known species of this genus, M. illustris Naväs, 1923. AI these
taxa were considered as enigmatic since their original description, almost
one hundred years ago. Marcenendius 1s redefined to contain also the
Macaronesian species M. fortunatus (Naväs, 1917) comb. nov. and the
African species M. angolensis (Badonnel, 1955) comb. nov., both formerly
assigned to VNephax Pearman, 1935. The diagnosis of the latter genus 1s
revised and for the species N. nepalensis (New, 1973) the original combi-
nation Seopsis nepalensis New comb. rev. is reinstated. An identification
key to the four amphientomid species known from the western Palaearctic
is presented: Nephax sofadanus Pearman, 1935, N. postalatus Lienhard,
2009, Marcenendius nostras, M. fortunatus. Nymphs of the latter two
species are characterized by the presence of characteristically curled
"corkscrew" hairs on dorsal side of thorax and abdomen, a kind of setae
previously unknown in Psocoptera, which are probably responsible for
nymphal camouflage due to adherent dust particles.

Keywords: Nephax Pearman - Spain - Mallorca - Macaronesia - nymphal
camouflage - cave fauna - soil fauna.

INTRODUCTION

Since its description by Naväs (1913) from south-eastern Spain (Alicante
Province), almost one hundred years ago, Marcenendius Naväs was one of the most
enigmatic genera of European Psocoptera. Its author assigned it to the family
Amphientomidae (suborder Troctomorpha, infraorder Amphientometae) and pointed
out that these predominantly tropical scaly-winged psocids were not known previously
from Europe. Ten years later, Naväs (1923) described a second species of this genus
from north-eastern Spain (Tarragona Province). Unfortunately the type material of
these species could not be found by Meinander (1979) when revising the European

Manuscript accepted03.06.2011

452 C. LIENHARD & A. BAZ

Psocoptera recorded by Naväs. Due to the very superficial original descriptions and to
the lack of new material these taxa were usually considered as nomina dubia and there-
fore not keyed in comprehensive works by subsequent authors (Badonnel, 1943;
Roesler, 1944: Smithers, 1990; Lienhard, 1998), though regularly cited in checklists of
Spanish Psocoptera or in papers on local faunistics, based on Naväs' original records
(Acon Remacha, 1980; Baz, 1989, 2007).

Pearman (1935) described a second genus of Amphientomidae from Palestine
(present day Israel), Nephax Pearman, the type-species of which, N. sofadanus
Pearman, was redescribed by Lienhard (1988). À second European species, initially
described in the family Lepidopsocidae from the Canary islands (Tenerife) by Naväs
(1917), was tentatively assigned to Nephax by Meinander (1973). Both species are
brachypterous and very similar in habitus, but they differ by several significant mor-
phological characters (see Lienhard, 1988, 1998). A third species, Nephax postalatus
Lienhard, macropterous but morphologically closely related to the type species, was
recently described from the United Arab Emirates (Lienhard, 2009). Some similarities
between this macropterous species and the two fully-winged species of Marcenendius
suggested that this genus could perhaps be a senior synonym of Nephax (see Lienhard,
2009). At the same time, some striking morphological differences between the Macaro-
nesian species N. fortunatus (Naväs) and the two closely related species N. sofadanus
and N. postalatus suggested that it might be justified assigning them to two different
genera.

The recent discovery of a fully winged amphientomid species in the southeast
of the Iberian peninsula and on the island of Mallorca enables us to resolve both prob-
lems: the apparent heterogeneity of the genus Nephax, as it was defined by Lienhard
(1988, 1998), and the enigmatic status of the genus Marcenendius. Based on this new
material we redescribe the type species M. nostras. The second species described by
Naväs, M. illustris, is here tentatively considered as a junior synonym of M. nostras.
The Macaronesian species Nephax fortunatus 1s closely related to M. nostras and has
to be transferred to Marcenendius.The genus Nephax is redefined and an identification
key to the four presently known western Palaearctic amphientomids 1s given.

MATERIAL AND METHODS

Dissection and slide-mounting followed the methods described by Lienhard
(1998). The material examined has been deposited in the following institutions:
MHNG = Muséum d'histoire naturelle, Geneva, Switzerland; UAH = Universidad de
Alcalä de Henares, Spain.

The following abbreviations are used in the descriptions: BL = body length (in
alcohol); F = hindfemur (length); FW = forewing (length); H = height of head capsule
from top of the vertex to anterior labral margin; HW = hindwing (length); IO/D =
shortest distance between compound eyes divided by anteroposterior diameter of
compound eye in dorsal view of head; L = length of head capsule from the hind margin
of the gena, near compound eye, to maximal postclypeal bulge, in lateral view: P2 =
second article of maxillary palp; T = hindtibia (length); t1, t2, t3 = tarsomeres of hind-
tarsus (length, measured from condyle to condyle). Abbreviations of Wing veins are
used according to Yoshizawa (2005).

REDESCRIPTION OF MARCENENDIUS NAVAS 453

TAXONOMIC TREATMENT

Marcenendius Naväs, 1913
Marcenendius Naväs, 1913: 334. Type species (by original designation): Marcenendius nostras

Naväs, 1913: 334.

REVISED DIAGNOSIS: Belonging to Amphientomidae. Forewings densely
covered with scales (Figs 1, Sc). Median ocellus close to postclypeus, smaller than
lateral ocelli, always present in fully-winged forms, occasionally strongly reduced or
absent in brachypterous forms; lateral ocelli widely separated, close to compound eyes.
Antenna with 14 articles (1. e. 12 flagellomeres), rarely the two apicalmost flagello-
meres fused. P2 only with simple, short hairs, lacking spur sensillum and macrochaetae
(Fig. 6c). Mandibles of normal shape (Fig. 6a). Distal half of labrum with a pair of
well-sclerotized longitudinal labral rods, originating from labral nodes (Keler, 1966:
nodus labralis) on internal face of labrum and running parallel to each other in about
1/3 of its width from lateral margin (Fig. 2e). Phallosome Y-shaped, with a slhightly
sclerotized longitudinal internal zone on each side in apical half (Fig. 3d). Internal
T-shaped sclerite of female subgenital plate present but occasionally strongly reduced
(Figs 2d, 3a, 6d). Nymphs with dorsal side of abdomen and thorax (including wing-
pads) densely covered by characteristically curled "corkscrew" hairs (Fig. 6b).

SPECIES ASSIGNED TO MARCENENDIUS: Marcenendius nostras Naväs, 1913;
Marcenendius fortunatus (Naväs, 1917) comb. nov. from Nephax; Marcenendius
angolensis (Badonnel, 1955) comb. nov. from Nephax.

DISCUSSION: Naväs' descriptions concerning his genus Marcenendius are rather
superficial (Naväs, 1913, 1923), but he mentions one character that 1s really diagnostic
within western Palaearctic amphientomids, 1. e. the presence of three widely separated
ocelli (see Naväs, 1923). In the only other amphientomid genus of this region, Nephax
Pearman, the median ocellus is absent. Several macropterous amphientomids recently
collected in continental Spain and on the island of Mallorca correspond to Naväs’
description with respect to this ocellar character. In the following we explain briefly
why we tentatively assign these specimens to the genus Marcenendius and in particular
to 1ts type species Marcenendius nostras.

The venation of forewing and hindwing figured by Naväs (1913, 1923)
corresponds to macropterous individuals and is very similar to that observed in the new
material, except for the presence of only one anal vein in the figure given by Naväs
(1913) for M. nostras. We interpret this as an inaccuracy of Naväs' figure (in Amphi-
entomidae two anal veins are always present; see Smithers, 1972). The shape of the
apex of forewing and hindwing seems to be somewhat variable in Marcenendius,
broadly rounded (Naväs, 1913: hindwing; Navas, 1923: forewing) or slightly acu -
minate (Naväs, 1913: forewing; Naväs, 1923: hindwing). The wings of the new
material are about intermediate in shape (Fig. 2a, b). In Naväs' figures of forewings the
veins CuP and Al reach the Wing margin separately (nodulus absent). This condition
is rather unusual in Amphientomidae where CuP and Al generally end together on
wing margin, forming the nodulus (see Smithers, 1972). But absence of nodulus has
also been observed in the brachypterous and macropterous amphientomids previously
known from western Palaearctic (see Lienhard, 1998, 2009). In the new material from

454 C. LIENHARD & A. BAZ

continental Spain and Mallorca the distal part of AI is usually very faint and curved
towards CuP near wing margin, meeting the latter on the margin (nodulus); this
situation could be confirmed on several slide-mounted forewings (Fig. 2a). However,
at low magnification AI seems to end in the membrane, almost parallel to CuP. Thus,
the absence of a nodulus in Naväs' figures 1s probably due to inaccurate drawing (1. e.
incorrect extrapolation of the faint distal part of Al).

The presence in Spain (Alicante, Almeria, Malaga and Mallorca) of an am-
phientomid species not belonging to the genus Nephax and showing some characters
mentioned by Naväs (1913, 1923) for his genus Marcenendius is here considered as
sufficient for redescribing the genus based on this new material, which is tentatively
assigned to the type species M. nostras. Marcenendius is Well-defined by a striking
autapomorphy of nymphs, the presence of "corkscrew" hairs on dorsal side of thorax
and abdomen (Fig. 6b), a kind of setae previously unknown in Psocoptera. These hairs
were present in the nymphs from Alicante, Almeria and Mallorca (M. nostras) and also
in nymphs from different localities on Canary islands which belong to a second species
of Marcenendius, M. fortunatus (Naväs) comb. nov. (see also discussion of this
species, below).

Meinander (1973) already considered the African species Nephax angolensis
Badonnel as "evidently congeneric" with M. fortunatus (Naväs) when redescribing the
latter as Nephax fortunatus. Because there 1s no doubt that this Macaronesian species
belongs to the newly defined genus Marcenendius, We formally transfer also the
African species to this genus, M. angolensis (Badonnel) comb. nov. According to the
description by Badonnel (1955) it differs from both western Palaearctic species by the
absence of a stigmapophysis in forewing and by the particular shape of the T-shaped
sclerite of the female subgenital plate and of the lacimial tip.

For discussion of differences in forewing venation between macropterous and
brachypterous amphientomids, see genus discussion of Nephax Pearman, below.

Marcenendius nostras Naväs, 1913 Figs 1-4; 5a, b

Marcenendius nostras Naväs, 1913: 334 (2 syntypes of unknown sex, depository not known,
probably lost).

Marcenendius illustris Naväs, 1923: 11 (holotype of unknown sex, depository not known, prob-
ably lost). Syn. nov.

MATERIAL EXAMINED: Spain (Alicante): MHNG, 19 , Xixona, Peñas Roset, under stones
in a ravine (dry river bed), 15.vi.2003, leg. V. M. Ortuño; MHNG, 1 nymph, same locality, under
stones, 23.v.2004, leg. V. M. Ortuño; UAH, 9 nymphs, same locality, under stones, 29.x.2009,
leg. V. M. Ortuño. — Spain (Almeria): UAH, 19% ,5 nymphs, Maria, Umbria de la Virgen, under
stones, 17.xii.2002, leg. V. M. Ortuño. — Spain (Malaga): UAH, 14, Velez-Malaga, Finca La
Alegria, on flight interception trap, 15.x.2008, leg J. M. Vela. — Spain (Mallorca island): MEHNG,
36,19,2 nymphs, Calviä, cave "Avenc de sa Finestreta", about 8m from the entrance inside the
cave, on the wall, 1.v.2010, leg. M. Vadell; UAH, 1 nymph, Calvié, cave "Avenc des Eriço",
about 15m from the entrance inside the cave, 27.1v.2007, leg. M. Vadell: UAH, 8 nymphs,
Calvi, cave "Cova de Na Boira", about 12m from the entrance inside the cave, 24.11.2007, leg.
M. Vadell; UAH, 7 nymphs, Calvi, cave "Cova de s'Estora", about 7m from the entrance inside
the cave, 28.11.2010, leg. M. Vadell.

REDESCRIPTION: Colouration: Body yellowish to brown, specimens from
Mallorca particularly dark coloured. Compound eyes black; vertex yellowish with
some light brown patches, vertical suture distinct, frontal sutures faint or not visible;

REDESCRIPTION OF MARCENENDIUS NAVAS

FIG. 1

Marcenendius nostras Naväs, from Mallorca, on wall of cave (sex unknown, body length about
3 mm): (a) Habitus in dorsolateral view. (b) Habitus in dorsal view. Photographs: M. Vadell.

456 C. LIENHARD & A. BAZ

postclypeus, frons and genae brown, anteclypeus yellow; basal two thirds of labrum
brown, apical part yellowish white. Thorax laterally with a longitudinal zone of reddish
brown hypodermal pigment, legs yellowish to medium brown, no transversal annu-
lation visible on tibiae. Forewing membrane tinged with brown in basal two thirds,
apical one third very light brown or almost hyaline; pattern due to scales somewhat
variable (see Fig. 1), usually not well-preserved in alcohol specimens. Hindwing mem-
brane light brown, with few scales. Membranous parts of abdomen yellowish with
some hypodermal pigment, in particular laterally; terminalia light to dark brown; in
male the three sternites preceding hypandrium medially with a brown sclerotized
transversal band.

Morphology: See genus diagnosis (above) and identification key (below), with
the following additions. Vertex abruptly rounded but not clearly sharp-edged, occiput
slightly concave. Compound eyes of about same size in both sexes (10/D 1.9-2,2). Tip
of lacinia as in Figs 2c and 3e, inner tine relatively close to lacinial apex (1. e. outer tine
not much longer than its basal width), shape of distal margin of outer tine somewhat
variable. Pretarsal claws with a distinct preapical tooth and a row of microtrichia on
ventral margin, femora of forelegs antero-ventrally with a longitudinal row of small
denticles (see Lienhard, 1998: Fig. 491, j), Pearman's organ of hindcoxa incomplete
(tympanum present, rasp absent). Wing venation as in Fig. 2a, b; forewing densely
covered with scales, hindwing with scales only in apical and marginal parts; scales
longitudinally striate, with truncate or slightly concave apex. — Female terminalia:
Epiproct and paraproct simple. Gonapophyses as in Figs 2h and 3b, usually bare (in the
female from Mallorca with a very fine subbasal hair on the right ventral valvula and
one/two such hairs on external valvulae). Subgenital plate heavily pilose, dorsally on
each side near lateral margin of apical part with a more or less sclerotized area (this
area particularly well-sclerotized in the female from Mallorca, shown by interrupted

, lines in Fig. 3a), T-shaped sclerite reduced to a small tubercle (Figs 2d and 3a). Sper-

mapore region with an oval or almost circular plate bearing the spermapore and a trian -
gular appendix (Figs 2g, 3c). Spermathecal duct long and spirally coiled; duct/sac in-
terface as in Fig. 2f, wall of sac near origin of duct with a transversal row of 8 (Alicante
2) or 9 (Mallorca ©) denticles, some of them minute; canal of spermatophore bent but
not S-shaped (Fig. 2f).- Male terminalia: Epiproct, paraproct and hypandrium simple,
the latter comprising two sclerotized sternites. Phallosome as in Fig. 3d.

MEASUREMENTS: Female (® Alicante/$ Mallorca): BL = 2.7/3.2 mm; FW =
2.813.2 mm; HW =2.3/26 mm F-10775 mA 1825/1570 pa = 605 Re
t2 = 84/103 pm; t3 = 130/132 pm; IO/D = 1.9/2.2. - Male (Mallorca MHNG 8060):
BL = 2.9 mm; FW =33 mm; EW=2.7mm; F=990 4m L="1580 2m otre
2= 100 40 = 155 mn MIO DE

DISTRIBUTION (see also Fig. 4): Spain: Alicante (Orihuela, S of Alicante, type
locality of M. nostras; Xixona, Peñas Roset, see above), Tarragona (Cabaces, a village
in the hills near Montsant, region near Reus, type locality of M. illustris), Almeria
(Maria, see above), Malaga (Velez-Malaga, see above), Mallorca (Calviä, see above).

DISCUSSION: This macropterous species is characterized by the strong reduction
of the T-shaped sclerite of the female subgenital plate (Figs 2d, 3a). No significant

REDESCRIPTION OF MARCENENDIUS NAVAS 457

FIG. 2

Marcenendius nostras Naväs, from Alicante, female: (a) Forewing (pigmentation of membrane
not shown). (b) Hindwing (ditto). (c) Lacinial tip. (d) Hindmargin of subgenital plate with
rudiment of T-shaped sclerite, pilosity not shown. (e) Labrum, pilosity not shown. (f)
Spermatheca, sac near duct, with one spermatophore. (g) Spermapore region. (h) Gonapophyses.

458 C. LIENHARD & A. BAZ

FIG. 3

Marcenendius nostras Naväs., from Mallorca, female (a-c) and male (d-e): (a) Hindmargin of
subgenital plate with rudiment of T-shaped sclerite and dorsolateral sclerotized areas (interrupted
lines), pilosity not shown. (b) Gonapophyses. (c) Spermapore region. (d) Phallosome. (e)
Lacinial tip.

diagnostic characters separating M. nostras from M. illustris are indicated by Naväs
(1913, 1923). In view of the new material from south-eastern Spain and from Mallorca
(see Fig. 4), belonging to one relatively widely distributed and rather variable species
(the Mallorca specimens are somewhat darker and larger than the specimens from
continental Spain), we see no reason to think that Naväs' specimen from Tarragona
might represent another species of this genus. Thus, we tentatively propose the above
mentioned synonymy. The species has been found under stones in open places
(Alicante and Almeria specimens) but also in caves, relatively near to their entrance
(Mallorca specimens). Nymphal camouflage has not been directly observed in this
species. The presence of corkscrew hairs (see genus discussion) suggests a similar
biology as described below for M. fortunatus.

REDESCRIPTION OF MARCENENDIUS NAVAS 459

M. illustris locus typicus

+

Spain

M. nostras locus typicus 4

Mallorca

FIG. 4
Marcenendius nostras Naväs, map of distribution.

Marcenendius fortunatus (Naväs, 1917) comb. nov. Figs 6a-d; Sc, d
Perientomum fortunatum Naväs, 1917: 20.
Nephax fortunatus (Naväs, 1913): Meinander, 1973: 143 (redescription based on holotype, male,

figs);, Lienhard, 1988: 369 (further description, male and female, figs); Lienhard, 1998:

141 (diagnosis, figs).

DISTRIBUTION: M. fortunatus is a Macaronesian endemic known from the
following islands. Canary Islands: Tenerife (Naväs, 1917; Meinander, 1973: Lienhard,
1988; Baz & Zurita, 2001, 2004); La Palma (Domingo-Quero ef al., 2003); un -
published records from Tenerife, La Palma, La Gomera, El Hierro, Gran Canaria,
Fuerteventura, Lanzarote (leg. C. Lienhard, material in the MHNG). — Cape Verde
Islands: island of Santiago (Lienhard, 1988: Lienhard & Garcia, 2005).

DISCUSSION: Based on general morphology of adults and nymphs, this brachy -
pterous species is closely related to M. nostras (see key, below). Macropterous
specimens of M. fortunatus have never been observed, but the most significant mor-
phological difference between this species and M. nostras is the strong reduction, in
the latter, of the T-shaped sclerite of the female subgenital plate.

M. fortunatus lives usually under stones at places incompletely covered by xe-
rophytic shrub vegetation, often also in stone walls built by humans (Fig. 5d). Adults
are rather well camouflaged by their mottled wing pattern due to scales (Fig. 5c) and

460 C. LIENHARD & A. BAZ

FIG. 5. Marcenendius nostras Naväs: (a) Biotope (dry river bed) of the Alicante specimens
(Peñas Roset). (b) General view of Peñas Roset.— Marcenendius fortunatus (Naväs): (c) Female
from Fuerteventura island (Canary Islands), near Lajares, 15.xi.1998, habitus in dorsolateral
view, body length 2.5 mm. (d) Biotope of this specimen (stone wall built by humans).
Photographs: V. M. Ortuño (a, b), C. Lienhard (c, d).

REDESCRIPTION OF MARCENENDIUS NAVAS 461

FIG. 6

Marcenendius fortunatus (Naväs), female (a-d): (a) Mandibles. (b) Three pronotal "corkscrew"
hairs. (c) Maxillary palp. (d) Hindmargin of subgenital plate with T-shaped sclerite, pilosity not
shown. — Nephax sofadanus Pearman, female (e-h): (e) Hindmargin of subgenital plate with
T-shaped sclerite, pilosity not shown. (f) Labrum, pilosity not shown. (g) Maxillary palp, with
spur sensillum and macrochaetae shown in black. (h) Mandibles.

nymphs usually carry some loosely arranged dust particles on dorsal side of thorax
(including wing pads) and of abdomen (personal observation by C. Lienhard). This
nymphal camouflage is probably due to the presence of corkscrew hairs on these parts

462 C. LIENHARD & A. BAZ

of the body (see genus discussion). Environmental dust particles probably adhere much
more easily to such curled hairs than to straight hairs. However, the effectiveness of
these hairs seems to be less than in the case of glandular hairs, where covering with
dust particles is more complete and nymphal camouflage much more impressive (e. g.
nymphs with glandular haïrs of Protroctopsocidae or Psocidae; see Lienhard, 1998).

Nephax Pearman, 1935
Nephax Pearman, 1935: 134. Type species (by original designation): Nephax sofadanus

Pearman, 1935: 134.

REVISED DIAGNOSIS: Belonging to Amphientomidae. Forewings densely
covered with scales. Median ocellus absent (even in fully-winged forms); lateral ocelli
widely separated, close to compound eyes. Antenna with 13 articles (1. e. 11 flagel-
lomeres). P2 with spur sensillum on inner side and some external macrochaetae in
addition to the simple and relatively short general pilosity (Fig. 6g). Mandibles with
shortened apical part (Fig. 6h). Labrum lacking pair of parallel longitudinal labral rods;
weakly developed sclerites originating from labral nodes diverging and running almost
parallel to antero-lateral margin of labrum (Fig. 6f). Phallosome V-shaped, lacking
internal sclerotizations. Internal T-shaped sclerite of female subgenital plate well-
developed (Fig. 6e). Nymphs with simple straight hairs on dorsal side of thorax and
abdomen.

SPECIES ASSIGNED TO NEPHAX: Nephax sofadanus Pearman, 1935; Nephax
capensis Pearman, 1935; Nephax postalatus Lienhard, 2009.

DISCUSSION: The genus Nephax is well-defined by the autapomorphic shor -
tening of the mandibles, which has been observed in the type species and in N. posfa-
latus. The South African Nephax capensis Pearman probably does not belong to this
genus, as it is defined here; the species has two ocelli placed close together and
mandibles of normal shape (see Pearman, 1935). However, we provisionally retain the
original combination, because no assignment to another amphientomid genus 1s
evident at present. The species Seopsis nepalensis New, 1973, has been tentativley
transferred to Nephax by Li Fasheng (1993) (see also Lienhard & Smithers, 2002).
Based on the original description by New (1973) it is clear that this species does not
belong to this genus as it is defined here (ocelli close together, phallosome with internal
sclerites, spermapore plate large and associated with gonapophyses; see New, 1973:
figs 5 and 7). Thus the original combination Seopsis nepalensis New comb. rev. is here
reinstated.

The striking differences in forewing venation between the macropterous species
N. postalatus and the brachypterous species N. sofadanus are probably related to wing
reduction, exactly the same differences have been observed in the genus
Marcenendius, between the macropterours M. nostras and the brachypterous M. fortu-
natus. In macropterous forms Rs is connected to M by a short crossvein and the
stigmapophysis is situated distally of the RI-RSs bifurcation (this corresponds to the
typical venation of Amphientomidae; see Smithers, 1972); in brachypterous forms Rs
and M are fused for a length and the stigmapophysis is situated on R stem, basally of
the RI-Rs bifurcation. We consider the striking similarity in forewing venation

REDESCRIPTION OF MARCENENDIUS NAVAS 463

between these brachypterous forms not as a synapomorphy but as a case of conver-
gence due to wing reduction.

Nephax sofadanus Pearman, 1935 Fig. 6e-h

Nephax sofadanus Pearman, 1935: 134 (syntypes, male, figs).

Nephax sofadanus Pearman, 1935: Lienhard, 1988: 369 (further description, male and female,
designation of lectotype, figs); Lienhard, 1998: 139 (diagnosis, figs).

DISTRIBUTION: N. sofadanus is an eastern Mediterranean species known from
the following countries. Israel (Pearman, 1935; Lienhard, 1988). Cyprus (Lienhard,
1998). Greece: Cyclades islands of Iraklia and Paros (Lienhard, 1988); several un-
published records (leg. K. Thaler and C. Lienhard, material in the MHNG) from Crete
and Dodecanese islands (Rhodes and Karpathos). Croatia: several localities near Split
(leg. T. Radja, material in the MHNG).

DISCUSSION: AIT specimens of N. sofadanus known at present are brachypter-
ous, they usually live under stones in open places incompletely covered by xerophytic
shrub vegetation (e. g. Greek “"phrygana" vegétation). Adults are rather well camou-
flaged by their mottled wing pattern due to scales, while nymphs are always "naked"
(Lienhard, 1998: plate 9h). A nymphal camouflage by dust particles has never been
observed. The dorsal pilosity on thorax and abdomen is normal, neither corkscrew
hairs as in Marcenendius nor glandular hairs as in nymphs of some other psocids are
present (see discussion of Marcenendius fortunatus, above).

Nephax postalatus Lienhard, 2009
Nephax postalatus Lienhard, 2009: 50 (male and female, figs).

DISTRIBUTION: United Arab Emirates: Hatta and Wadi Maidaq (Lienhard,
2009).

DISCUSSION: No details are known about the biology of this macropterous
species, because all available specimens were trapped (light trap, Malaise trap, water
trap); the nymph remains unknown (see Lienhard, 2009).

KEY TO THE WESTERN PALAEARCTIC AMPHIENTOMIDS
NOTE: For additional figures see Lienhard (1988, 1998, 2009) and Meinander (1973).

1 Antenna with 13 articles (1. e. 11 flagellomeres). Median ocellus absent
(even in fully-winged forms). P2 with spur sensillum on inner side and
some external macrochaetae in addition to the simple and relatively short
general pilosity (Fig. 6g). Mandibles with shortened apical part (Fig. 6h).
Labrum lacking pair of parallel longitudinal labral rods (Fig. 6f).
Nymphs with simple straight hairs on dorsal side of thorax and abdomen
pee CR TERRE PRE PR EEE Nephax Pearman...…. 2

— Antenna with 14 articles (1. e. 12 flagellomeres), rarely the two apical-
most flagellomeres fused. Median ocellus usually present, especially in
fully-winged forms. P2 only with simple, short hairs, lacking spur sen-
sillum and macrochaetae (Fig. 6c). Mandibles of normal shape (Fig. 6a).
Distal half of labrum with a pair of well-sclerotized parallel longitudinal

464 C. LIENHARD & A. BAZ

labral rods (Fig. 2e). Nymphs with characteristically curled corkscrew

hairs on dorsal side of thorax and abdomen (Fig. 6b) Marcenendius Naväs…... 3
> Both sexes macropterous, hindwing well-developed, with complete

venation, all wings clearly projecting over abdominal apex. In forewing

Rs connected to M by a short crossvein. Short inflated area of R

(stigmapophysis) situated distally of R1-Rs bifurcation. Head capsule

slightly flattened (1. e. H/L>2). Spermapore plate with a forceps-like pair

OLSHOrT DrOCESSEST EL ENORME ANS ARE Nephax postalatus Lienhard
- Both sexes brachypterous, forewing only slightly projecting over

abdominal apex, hindwing reduced to a minute veinless rudiment. In

forewing Rs and M fused for a length. Stigmapophysis situated on R

stem, basally of R1-RSs bifurcation. Head capsule almost semicircular in

profile (1. e. H/L<2). Spermapore plate with a short median appendix . .

RAP deers rcmoncrots M na ES EU EE Nephax sofadanus Pearman
3 Both sexes macropterous (Figs 1; 2a, b), forewing apically rounded or

slightly acuminate, hindwing well-developed, with complete venation,

all wings clearly projecting over abdominal apex. In forewing Rs

connected to M by a short crossvein; short inflated area of R

(stigmapophysis) situated distally of RI1-Rs bifurcation (Fig. 2a). T-

shaped sclerite of female subgenital plate strongly reduced (Figs 2d, 3a)

sa Sears eee ee NOR EC ENE ÉÉRCCTE Marcenendius nostras Navâs
- Both sexes brachypterous (Fig. 5c), forewing apically strongly acu-

minate, only slightly projecting over abdominal apex, hindwing reduced

to a minute veinless rudiment. In forewing Rs and M fused for a length;

stigmapophysis situated on R stem, basally of RI1-Rs bifurcation. T-

shaped sclerite of female subgenital plate well-developed (Fig. 6d) ....

sa nt rat ns ERP RER RL RE Marcenendius fortunatus (Navâs)

ACKNOWLEDGEMENTS

We are grateful to V. M. Ortuño, M. Vadell, J. M. Vela, T. Radja and K. Thaler
for entrusting their specimens of Marcenendius nostras and Nephax sofadanus to A.
Baz and C. Lienhard respectively. V. M. Ortuño and M. Vadell also provided excellent
biotope or habitus photographs. E. L. Mockford (Normal, Illinois, USA) and J. Hollier
(MHNOG) are acknowledged for reading the manuscript and making valuable sugges-
tions and F. Marteau (MHNOG) for technical assistance.

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REVUE SUISSE DE ZOOLOGIE 118 (3): 467-484; septembre 2011

An annotated list of the parasitic nematodes (Nematoda)
of freshwater fishes from Paraguay deposited in the
Museum of Natural History of Geneva

David GONZAÂLEZ-SOLIS! & Jean MARIAUX?2

l El Colegio de la Frontera Sur (ECOSUR), Unidad Chetumal. Av. Centenario Km. 5.5,
C.P. 77900, Chetumal, Quintana Roo, Mexico. Email: dgonzale@ecosur.mx

2 Dpt des Invertébrés, Muséum d’histoire naturelle, CP 6434, CH-1211 Genève 6,
Switzerland.

An annotated list of the parasitic nematodes (Nematoda) of freshwater
fishes from Paraguay deposited in the Museum of Natural History of
Geneva. - The list comprises the evaluation of the material deposited in the
Museum d'Histoire naturelle, Geneva, whose taxonomic examination
revealed the presence of 78 taxa of parasitic nematodes of freshwater fishes
from Paraguay. Out of these, 43 were specifically identified, while 35 larval
or subadult forms were only determined at the generic of familial level
because of their developmental status. A total of 150 new hosts and 44 geo-
graphical records were reported in this survey, thus increasing the number
of known nematodes infecting fishes in Paraguay from 28 to 87. Rondonia
rondoni and Procamallanus (Spirocamallanus) inopinatus Were the most
frequent nematodes both in number of localities and hosts. Camallanidae
was the best represented nematode family with 10 species, followed by
Anisakidae (9), Pharyngodonidae (7) and Cucullanidae (5). The nematode
fauna of Paraguyan fishes is quite similar to that of fishes from Brazil and
Argentina.

Keywords: Nematoda - Paraguay - Freshwater - Fishes.

INTRODUCTION

The nematode fauna parasitizing freshwater fishes from the Neotropical region
is poorly known and there still exist vast territories where parasitological surveys are
lacking (Moravec, 1998). Paraguay represents one of these scarcely studied areas,
since only few reports on fish parasites, based on a limited number of hosts and from
very specific geographical areas, have been published so far (Masi Pallarés ef al., 1973;
Petter 1984, 1989, 1990, 1994, 1995ab; Petter and Cassone, 1984; Petter and Dlouhy,
1985; Petter and Morand, 1988). From these works, to date 28 nematodes have been
reported from Paraguay, particularly from localities of the Paranä and Paraguay Rivers
and provinces bordering or close to Brazil (Concepcion, Alto Paranä and San Pedro)
and Argentina (Misiones, Neembucu, Itapua). This paper aims to contribute to the
knowledge of the nematode fauna in freshwater fishes from various Paraguayan
provinces, especially those for which no previous information was reported, on the
basis of the examination of material deposited in the Muséum d’histoire naturelle in
Geneva (MHNG).

Manuscript accepted 25.03.2011

468 D. GONZÂLES-SOLIS & J. MARIAUX

MATERIALS AND METHODS

The nematode specimens used for this publication were recovered during the
various expeditions of the MHNG in Paraguay (EMGP) between 1987 and 1996.
Fishes belonging to 63 genera and 92 species, were collected during February-
November 1987, October-November 1988, 1989, 1991, 1996, and August 1994 from
20 localities mostly located in the southeastern region of the country (Central,
Neembucu, and Concepcion Provinces), the vicinity of Asuncion City and along the
Rio Pilcomayo. Only two or three localities were located in the Paraguayan Alto
Paranä, which represents the most nematologically studied region in South America
(see Moravec, 1998).

Nematodes were cleared through the evaporation of water of the glycerine-
water mixture (ratio 1:10-1:2) (Moravec, 1998). The following list mentions the
nematodes recovered from 151 fish specimens belonging to 92 species. Taxonomic
references for hosts and parasites are Froese and Pauly (2011) and Moravec (1998)
respectively. Each identified taxon 1s listed together with its host names, abbreviated
localities and sampling date. Host species with an asterisk (*) represent new records.
A list of localities 1s provided in the appendix.

RESULTS

ORDER ENOPLIDA
FAMILY CAPILLARIIDAE RAILLIET, 1915
Capillariidae gen. sp.

SPECIMENS: MHNG INVE 73740, 73501, 73531, 73509, 74454.

HOSTS: Astyanax asuncionensis Géry, 1972 (EP 01/11/87). Cynopotamus
argenteus (Valenciennes, 1836) (VI 13/11/87). Roeboides microlepis (Steindachner,
1879)* (VI 13/11/87). Pygocentrus nattereri Steindachner, 1908 (= Serrasalmus
nattereri)* (VI 13/11/87, PC 05/01/96).

REMARKS: These specimens were young females or males whose specific or
generic identication Was impossible. Capillariids from À. asuncionensis and C. argen-
teus might belong to Capillostrongyloides sentinosa or Paracapillaria piscicola,
respectively. This is the first record of capillariids in R. microlepis and P. nattereri.

Capillostrongyloides sentinosa (Travassos, 1927)
SPECIMENS: MHNG INVE 74278.

HOST: Hoplias malabaricus (Bloch, 1794) (ED 01/08/94).

REMARKS: This is a new geographical record since this taxon was originally
reported from Brazil (Travassos, 1927).

ORDER OXYURIDA CHABAUD, 1974
FAMILY PHARYNGODONIDAE TRAVASSOS, 1919
Pharyngodonidae gen. sp.

SPECIMENS: MHNG INVE 73325, 73345.

HOST: Trachelyopterus galeatus (Linnaeus, 1766) (= Parauchenipterus
galeatus) (AY 14/05/87, CA 02/06/87).

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 469

REMARKS: Larvae poorly developed.

Brasilnema sp.
SPECIMEN: MHNG INVE 73783.

HOST: Hoplerythrinus unitaeniatus (Spix & Agassiz, 1829)* (ES 04/10/89).

REMARKS: Only one young female available that represents the first report of
the genus in this fish host.

Brasilnema pimelodellae Moravec, Kohn et Fernandes, 1992
SPECIMENS: MHNG INVE 73513.

HOST: Pimelodella sp. (VI 13/11/87).

REMARKS: New geographical record, previously reported from Brazil (Moravec
et al. 1992)

Cosmoxynema sp.
SPECIMENS: MHNG INVE 73511, 73621.

HOST: Psectrogaster curviventris Eigenmann & Kennedy, 1903* (VI 13/11/87,
GN 05/11/91).

REMARKS: These young females could belong to the species C. vianai, which
was found in a host of the same family (Pseudocurimata gilberti).

Cosmoxynema vianai Travassos, 1949
SPECIMENS: MHNG INVE 73664, 73675, 73678, 73682, 73690, 74215.

HOST: Cyphocharax modestus (Fernändez-Yépez, 1948) (= Curimata cf.
modesta)* (JL 18/10/87). Cyphocharax voga (Hensel, 1870) (= Curimata cf. voga)*
(AT 23/10/89).

REMARKS: Both are new host and geographical records.

Cosmoxynemoides aguirrei Travassos, 1949
SPECIMENS: MHNG INVE 73671,74186, 74216.

HOSTS: Curimata sp. (GN 19/10/89). Cyphocharax modestus* (JL 18/10/87).
Cyphocharax voga* (AT 23/10/89).

REMARKS: New host and geographical records.

Ichthyouris sp.
SPECIMEN: MHNG INVE 73691.

HOST: Cyphocharax modestus* (JL 18/10/87).

REMARKS: Only one male available that could be a new species since the genus
has never been found in this fish family (Curimatidae).

Ichthyouris brasiliensis Moravec, Kohn et Fernandes, 1992
SPECIMENS: MHNG INVE 73402, 73415, 73426.

HOST: Anadoras weddellii (Castelnau, 1855)* (EG 18-19/11/87).

470 D. GONZALES-SOLIS & J. MARIAUX

REMARKS: First record of this species in a Doradidae, formerly described from
Loricariidae (Moravec et al. 1992)

Parasynodontisia petterae Moravec, Kohn et Fernandes, 1992
SPECIMENS: MHNG INVE 73348, 73382, 73385, 73387, 13388.

HOST: Rhinelepis aspera Spix & Agassiz, 1829 (CA 02/06/87, VI 14/11/87).
REMARKS: New locality record.
Spinoxyuris Sp.
SPECIMEN: MENG INVE 73685.
HOST: Cyphocharax modestus* (JL 18/10/87).

REMARKS: First record of this genus in a curimatid species.

Spinoxyuris oxydoras Petter, 1994
SPECIMENS: MHING INVE 73592, 73766, 73767.

HOSTS: Anadoras weddellii (Castelnau, 1855)* (AM 01/11/89). Doradidae gen.
sp. (RS 08/10/91).

REMARKS: New host and geographical records.

Travnema araujoi Fernandes, Campos et Artigas, 1983
SPECIMENS: MHNG INVE 73660, 73669, 73670, 73674, 73677, 73680, 73684, 73689.

HOST: Cyphocharax modestus* (JL 18/10/87).
REMARKS: New host and geographical records, previously only found in Brazil.

ORDER ASCARIDIDA SKRJABIN ET SCHULZ, 1940
FAMILY ATRACTIDAE RAILLIET, 1917

” Klossinemella theringi (Travassos, Artigas et Pereira, 1928)
SPECIMENS: MENG INVE 73326.

HOST: Schizodon dissimilis (Garman, 1890)* (AY 14/05/87).

REMARKS: New host and geographical records, previously reported from Brazil
and Argentina (Moravec, 1998).

Rondonia sp.
SPECIMENS: MHNG INVE 73354, 73633.

HOSTS: Pimelodus maculatus Lacepède, 1803 (PE 12/08/87). Platydoras cos -
tatus (Linnaeus, 1758) (GN 07/11/91).

REMARKS: These larval nematodes probably belong to the species R. rondoni
which parasitizes congeneric fishes. Unfortunately, their poor development prevents a
specific identification.

Rondonia rondoni Travassos, 1920

SPECIMENS: MHNG INVE 74303, 74354, 74358, 74383, 74384,74385, 74437, 74471,
73308,73371,73854,73856, 74307, 74318, 74320, 74325, 74398, 74309, 73544, 74287, 74388,
74648,73302,73307,73312,73314,73317,73323,73343,73344,73351, 73590, 73653, 73654,
74205, 74335, 74338, 74403, 74221.

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY

HOSTS: Ageneiosus inermis Valenciennes, 1840 (= À. brevifilis)* (SA 06/11/95).
Doradidae gen. sp. (ED 26/06/96, 17/10/96). Piaractus mesopotamicus (Holmberg,
1887) (EO 12/05/87, VI 14/11/87, SA 14/10/89). Pimelodus sp. (R6 09/10/88, ED
01/08/94, 26/06/96, 26/09/96). Pimelodus cf. albicans (Valenciennes, 1840)* (ED
28/09/96). Pimelodus argenteus Perugia, 1891* (SA 02/11/95, 03/11/95). Pimelodus
maculatus* (SA 02/11/95). Oxydoras kneri Bleeker, 1862* (PI 05/01/96). Prerodoras
granulosus (Valenciennes, 1821) (PV 15/02/87, EO 12/05/87, YA, AG 12-14/05/87,
CA 02/06/87, PL 09/06/87, RS 08/10/91, AP 19/03/89, GN 19/10/89, SA 03/11/95).
Serrasalmus nattereri* (ED 02/10/96). Sorubim lima (Bloch & Schneider, 1801)* (SA

21/11/93).

REMARKS: Some new host and geographical records (Central Province).

FAMILY COSMOCERCIDAE RAILLIET, 1916

Raillietnema sp.
SPECIMEN: MHNG INVE 73702.

HOST: Hoplias malabaricus* (ES 26/10/87).

REMARKS: First record of the genus in this fish species and geographical area.

FAMILY KATHLANIIDAE LANE, 1914

Chabaudinema americanum Diaz-Hungria, 1968
SPECIMEN: MHNG INVE 73855.

HOST: Piaractus mesopotamicus* (SA 14/10/89).
REMARKS!: First record in this fish species and geographical area.

Spectatus spectatus Travassos, 1923
SPECIMEN: MANG INVE 73372.

HOST: Piaractus mesopotamicus* (VI 14/11/87).
REMARKS: First record in this fish species and geographical area.

FAMILY QUIMPERIIDAE GENDRE, 1928

Quimperiidae gen. sp.
SPECIMEN: MANG INVE 73553.
HOST: Brycon orbignyanus (Valenciennes, 1850) (AA 07/10/89).
REMARKS!: Poorly developed larva.

Paraseuratum sp.
SPECIMENS: MHNG INVE 73484, 73485.
HOST: Hoplias malabaricus (ES 04/10/89).
REMARKS: A small larva that probably belongs to the species P. soaresi.

Paraseuratum soaresi Fâbio, 1982
SPECIMENS: MHNG INVE 73708, 73731, 74638.

472 D. GONZALES-SOLIS & J. MARIAUX

HOST: Hoplias malabaricus (ES 26/10/87, 29/10/87, EM 08/08/96).
REMARKS: New geographical record.

Touzeta ecuadoris Petter, 1987
SPECIMEN: MHNG INVE 73710.

HOST: Cichlasoma dimerus (Heckel, 1840)* (ES 26/10/87).
REMARKS: New host and geographical records.

FAMILY CUCULLANIDAE MÜLLER, 1777

Cucullanidae gen. sp.
SPECIMENS: MHNG INVE 73365. 73484. 73637. 73658.

HOSTS: Brycon hilarii Perugia, 1897 (AA 03/11/87). Pimelodus sp. (ED
01/08/94). Plagioscion ternetzi Daneri, 1954 (CA 23/08/87). Rhamdia quelen (Quoy &
Gaimard, 1824) (JL 8/10/87).

REMARKS: These specimenes were small and poorly developed larvae that
could belong to the genus Cucullanus.

Cucullanus sp.

SPECIMENS: MHNG INVE 73328, 73347, 73483, 73486, 73545, 73608, 73611, 73757,
73762, 73765, 74226, 74227, 74228, 74229, 74297, 74365.

HOSTS: Ageneiosus inermis* (SA 29/11/93). Brycon hilarii (AA 03-04/11/87).
Luciopimelodus pati (Valenciennes, 1835) (AT 28/10/89). Megalonema platanum
(Günther, 1880) (CA 29/05/87). Pimelodus sp. (R6 09/10/88). Pimelodus cf. albicans
(ED 03/08/94). Pimelodus maculatus (CA 02/06/87). Pimelodus ornatus Kner. 1858*
(AT 28/10/89). Pinirampus pirinampu (Spix & Agassiz, 1829)* (SA 27/11/93,
" 29/11/93). Plagioscion ternetzi (RS 09/10/91). Rhamdia quelen* (AT 25/10/89).
Zungaro zungaro (Steindachner, 1877) (= Paulicea luetkeni) (SA 27/11/93).

REMARKS: Larvae and young females that could be of the species Cucullanus
pinnai pinnai Or C. p. pterodorasi.

Cucullanus (Cucullanus) pinnai pinnai Travassos, Artigas et Pereira, 1928

SPECIMENS: MHNG INVE 73356, 73357, 73546, 73626, 74308, 74314, 74321, 74329,
74333, 74341, 74387, 74393, 74397, 74399, 74413, 74418, 74422, 74424,

HOSTS: Ageneiosus inermis* (SA 04/11/95). Pimelodus sp. (R6 09/10/88, GN
07/11/91, ED 26/06/96). Pimelodus cf. albicans (ED 28/09/96, 02/10/96). Pimelodus
argenteus* (SA 02/11/95, 03/11/95). Pimelodus maculatus (PE 12/08/87, SA
03/11/95). Sorubim lima* (ED 02/10/96).

REMARKS: Some new host and geographical records for those localities out of
the Paranä River basin.

Cucullanus (Cucullanus) pinnai pterodorasi Moravec, Kohn et Fernandes, 1997
SPECIMENS: MHNG INVE 73319, 73583.

HOST: Plagioscion ternetzi* (PI 08/10/91). Prerodoras granulosus (AY
14/05/87).

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 473

REMARKS: New host and geographical records.

Cucullanus (Cucullanus) pseudoplatystomae Moravec, Kohn et Fernandes, 1993
SPECIMENS: MHNG INVE 73309, 73350, 73360.

HOST: Pseudoplatystoma corruscans (Spix & Agassiz, 1829) (EO 12/05/87, PL
09/06/87, AY 14/08/87).

REMARKS: New geographical record.

Dichelyne sp.
SPECIMENS: MHNG INVE 73640, 73642.
HOST: Pimelodus cf. albicans (ED 03/08/94).
REMARKS: These two young females might belong to D. pimelodi.

Dichelyne pimelodi Moravec. Kohn et Fernandes, 1997

SPECIMENS: MHNG INVE 73367,74171, 74317, 74343.

HOSTS: Pimelodus sp. (GN 18/10/89). Pimelodus argenteus* (SA 03/11/95,
04/11/95). Pimelodus maculatus* (CA 23/08/87).

REMARKS: New host and geographical records.

Neocucullanus neocucullanus Petter, 1989
(Syn. N. multipapillatus)
SPECIMENS: MHNG INVE 73479, 73482, 73485, 73487.

HOSTS: Brycon sp. (EP 31/10/87). Brycon hilarii* (AA 31/10/87,04/11/87).

REMARKS: Saraiva et al. (2006) considered N. multipapillatus as a junior
synonym of this species. New locality record, because it was reported from the same
basin (Paranä River) but from different province (Itaupu).

FAMILY ACANTHOCHEILIDAE WÜLKER, 1929

Brevimulticaecum sp.
SPECIMEN: MHNG INVE 74349.

HOST: Hoplosternum sp. (PO 15/08/96).
REMARKS: New host and geographical records.

FAMILY ANISAKIDAE RAILLIET ET HENRY, 1912
Anisakidae gen. sp.

SPECIMEN: MANG INVE 74214.

HOSTS: Eigenmannia virescens (Valenciennes, 1836)* (AH 22/10/89). Hoplias
malabaricus* (ED 01/08/94).

REMARKS: Very small larvae with poorly developed organs. This 1s the first ani-
sakid form occurring in these fish species.

Contracaecum sp. type 1

SPECIMENS: MHNG INVE 73352,73362, 73364, 73492, 73539, 73552, 73557, 73577,
73599, 73619, 73622, 73665, 73667, 73673,73676,73679,73683,73687,73717,73718,73761,

474 D. GONZAÂLES-SOLIS & J. MARIAUX

713763,73773,73782,73786, 73840, 73842,73848, 73850, 73852, 73863, 73866,73875,73880,
73885,73890, 74217, 74218, 74222, 714295, 74312, 74322, 74331, 74334, 74392, 74394, 74417,
74430, 74436, 74446, 74460, 74461, 74469, 74628, 74630, 74637.

HOSTS: Acestrorhynchus altus Menezes, 1969* (ES 05/10/89, SN 17/10/89).
Ageneiosus inermis* (AT 28/10/89). Ancistrus sp.* (AG 25/10/91). Astyanax sp.
(AT 23/10/89). Brycon sp.* (AA 09/10/89). Callichthys callichthys (Linnaeus, 1758)*
(ES 02/10/89). Cichlidae gen. sp. (AT 25/10/87). Crenicichla lepidota Heckel, 1840
(AT 25/10/89, 24/10/89). Curimatella immaculata (Fernändez-Yépez, 1948) (= Curi -
mata bimaculata)* (GN 05/11/91). Cyphocharax modestus* (JL 18/10/87). Gymnotus
carapo Linnaeus, 1758 (ES 27/10/87). Hoplerythrinus unitaeniatus* (ES 4-6/10/89).
Hoplias malabaricus (ES 04/10/89). Hoplosternum littorale (Hancock, 1828)* (SN
17/10/89). Leporinus sp. (ED 03/08/94). Leporinus lacustris Amaral Campos, 1945
(SN 17/10/89). Leporinus obtusidens (Valenciennes, 1837) (ED 26/06/96, PI 05/01/96,
EF 15/02/96). Loricariichthys labialis (Boulenger, 1895)* (EF 15/02/96). Pimelodus
cf. albicans* (ED 28/09/96). Pimelodus argenteus* (SA 03/11/95). Pimelodus macu-
latus (PE 11/08/87, SA 2-3/11/95). Plagioscion ternetzi* (CA 23/08/87).
Pseudoplatystoma corruscans (AY 14/08/87). Pygocentrus nattereri* (SA 04/10/96,
PC 05/01/96, EF 15/02/96). Rhamdia quelen* (ES 27/10/87, 05/10/89, SN 05/10/89,
16-17/10/89, ED 17/10/96). Rhaphiodon vulpinus Agassiz, 1829 (RS 09/10/91).
Salminus brasiliensis Valenciennes, 1850 (= Salminus maxillosus)* (GN 18/10/89).
Serrasalmus marginatus Valenciennes, 1837* (EF 15/02/96). Serrasalmus spilopleura
Kner, 1858* (PO 07/12/96). Sorubim lima* (SA 27/11/93, ED 02/10/96). Triportheus
paranensis (Günther, 1874)* (VI 13/11/87).

REMARKS: New host and geographical records. These larvae have been reported
as Contracaecum Sp. in many papers (see Moravec, 1998).

. Contracaecum sp. type 2

SPECIMENS: MHNG INVE 73871, 73874,73450, 73414, 73441, 74657, 73719, 73397,
73416,73733,73759,73715,73777, 713389, 73392, 73394, 73408, 73473, 13623, 13771, 73780,
73857,73858, 73864, 74439, 74642, 74400, 74339, 73342, 74342, 74459, 73655, 73860, 73520,
73602,74617,74634,73369,73889, 74429, 74434, 74408, 74445, 74415, 74421, 74649, 74346,
73643.

HOSTS: Acestrorhynchus altus* (EG 20/11/87, SN 17/10/89). Anadoras wed-
dellii* (EG 18/11/87, 19/11/87). Brycon hilarii* (AA 03/11/87). Gymnotus carapo (ES
27/10/87). Hoplerythrinus unitaeniatus (EG 18/11/87, Santa Sofia 29/10/87, AT
25/10/89, SU 03/10/89). Hoplias malabaricus (EG 18/11/87, 20/11/87, GN 05/11/91,
PC 28/09/89, SU 03/10/89, SN 16/10/89, EM 06/12/96, 08/08/96). Pimelodus cf. albi-
cans* (ED 28/09/96). Pinirampus pirinampu* (SA 04/11/95). Pseudoplatystoma
corruscans (CA 02/06/87, SA 04/11/95). Pygocentrus nattereri* (SA 04/10/96, PC
05/01/96). Rhamdia quelen* (JL 18/10/87, SN 16/10/89). Rhaphiodon vulpinus
(VI 13/11/87, RS 09/10/91, EF 15/02/96). Salminus brasiliensis* (CA 23/08/87,
GN 18/10/89). Serrasalmus spilopleura* (ED 02/10/96, PO 07/12/96). Sorubim lima*
(ED 02/10/96). Synbranchus marmoratus Bloch, 1795* (PO 09/08/96).
Trachelyopterus sp.* (ED 17/10/96).

REMARKS: New host and geographical records. These larvae have been reported
as Contracaecum Sp. in many papers (see Moravec, 1998), and differ from

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 475

Contracaecum sp. type 1 in the length ratio of the ventricular appendix and intestinal
caecum.

Hysterothylacium sp.

SPECIMENS: MHNG INVE 73333, 73349, 73355, 73358, 73359, 73366, 73368, 73370,
73391,73393, 73395, 73399, 73400,73411,73417, 73425, 73442, 73443, 73444, 73449, 73453,
73456, 73467, 73470, 73474, 73477, 73489, 73490, 73493, 73499, 73502, 73503, 73508, 73517,
D 00221227,15325,13512,13587, 73600, 73603, 73605, 73607, 73610, 73618, 73620,
73862, 73886, 74658.

HOSTS: Acestrorhynchus altus* (EG 20/11/87, VI 13/11/87, AT 24/10/89).
Anadoras weddellii* (EG 19/11/87). Brycon hilarii* (AA 03/11/87). Cichlasoma
dimerus* (EG 18/11/87). Cynopotamus argenteus* (VI 13/11/87). Cynopotamus kin-
caidi (Schultz, 1950)* (PE 12/08/87). Hoplias malabaricus* (EG 18/11/87, 20/11/87).
Hoplerythrinus unitaeniatus* (EG 18/11/87). Hoplosternum sp.* (EG 19/11/87).
Hoplosternum littorale* (EG 19/11/87, SN 17/10/89). Loricaria sp.* (VI 13/11/87).
Loricariichthys platymetopon Isbrücker & Nijssen, 1979* (EG 20/11/87). Lycengraulis
grossidens (Eigenmann, 1907)* (PE 13/08/87). Markiana nigripinnis (Perugia, 1891 )*
(EG 20/11/87). Megalechis thoracata (Valenciennes, 1840) (= Hoplosternum thora -
cata)* (PZ 16/10/91). Pellona flavipinnis (Valenciennes, 1837)* (VI 13/11/87).
Pimelodella sp.* (VI 13/11/87). Pimelodus maculatus* (CA 23/08/87). Plagioscion
ternetzi* (CA 30/05/87, 23/08/87, GN 05/11/91, PI 08/10/91, RS 09/10/91).
Pygocentrus nattereri* (VI 13/11/87). Rhamdia quelen* (SN 16/10/89). Rhaphiodon
vulpinus (VI 13/11/87, RS 09/10/91). Rhinelepis aspera* (CA 02/06/87). Roeboides
microlepis* (VI 13/11/87). Salminus brasiliensis* (CA 23/08/87). Triportheus para-
nensis* (EG 20/11/87, VI 13/11/87).

REMARKS: New host and geographical records. These larvae could belong to
different species, although their morphology is quite similar, except for larvae found in
Rhamdia quelen that could belong to H. rhamdiae.

Hysterothylacium rhamdiae Brizzola et Tanzola, 1995

SPECIMENS: MHNG INVE 73716, 73728, 73839, 73841, 73844, 73845, 73853, 73865,
73878.

HOST: Rhamdia quelen* (ES 27/10/87 29/10/87, 05/10/89, 06/10/89, SN
17/10/89).

REMARKS: New host and geographical records.

Raphidascaris sp.

SPECIMENS: MHNG INVE 73374, 73375, 73378, 13379, 73381, 73494, 74472.

HOSTS: Doradidae gen. sp. (EF 15/02/06), Hypostomus sp. (VI 14/11/87).
Loricaria Sp. (VI 13/11/87).

REMARKS: Small larvae that could belong to R. (S.) hypostomi for Hypostomus
sp. and R. (S.) manherti for Loricaria Sp.

Raphidascaris (Sprentascaris) hypostomi (Petter et Cassone, 1984)

SPECIMENS: MHNG INVE 73373, 73693, 73697, 73698, 73699, 73701.

HOSTS: Hypostomus dlouhy* (JL 18/10/87). Sturisoma robustum* (VI
14/11/87).

REMARKS: New host and locality records.

476 D. GONZALES-SOLIS & J. MARIAUX

Raphidascaris (Sprentascaris) mahnerti (Petter et Cassone, 1984)

SPECIMENS: MHNG INVE 73353, 73476, 73735, 74206, 74208, 74212, 74463, 74620,
74623.

HOSTS: Geophagus sp.* (PI 05/01/06). Loricariichthys sp. (GN 19/10/89).
Loricariichthys labialis (EF 15/02/96). Loricariichthys platymetopon (EG 20/11/87,
ES 29/10/87). Ricola macrops Regan, 1904* (PE 12/08/87).

REMARKS: Two new host records.

Raphidascaroides sp.
SPECIMEN: MHNG INVE 74181.

HOST: Pachyurus sp. (GN 19/10/89).
REMARKS: Small and weakly developed larva.

Raphidascaroides brasiliensis Moravec et Thatcher, 1997

SPECIMENS: MHNG INVE 733625, 73629, 73631, 73632, 73634, 73635, 74173, 74174,
74175,74176, 74177, 74188, 74271, 74274.

HOSTS: Platydoras armatulus (Valenciennes, 1840)* (GN 18/10/89, 19/10/89).
Platydoras costatus* (GN 05/11/91,07/11/91,GY 15/09/94).

REMARKS: New host and geographical records.

Goezia brasiliensis Moravec, Kohn et Fernandes, 1994
SPECIMEN: MANG INVE 73361.

HOST: Pseudoplatystoma corruscans (AY 14/08/87).
REMARKS: New geographical record.

- Porrocaecum sp.
SPECIMENS: MHNG INVE 73327, 73495, 73630.

HOSTS: Crenicichla niederleinii (Holmberg, 1891)* (VI 13/11/87). Platydoras
costatus* (GN 07/11/91). Schizodon dissimilis* (AY 14/05/87).

REMARKS: First record of this larva in these fish hosts and in South America.

ORDER SPIRURIDA CHITWOOD, 1933
FAMILIA SPIRURIDAE

Spiruridae gen. sp.
SPECIMEN: MENG INVE 73721.
HOST: Hoplerythrinus unitaeniatus (ES 27/10/87).
REMARKS: Very small and poorly developed larvae.

FAMILY CAMALLANIDAE RAILLIET ET HENRY, 1915

Camallanidae gen. sp.
SPECIMEN: MHNG INVE 73488.
HOST: Brycon hilarii (AA 04/11/87).
REMARKS: Small camallanid larva that might belong to P. (S.) inopinatus.

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 477

Procamallanus sp.
SPECIMENS: MHNG INVE 73627, 73872.
HOSTS: Acestrorhynchus altus (SN 17/10/89). Pimelodus sp. (GN 07/11/91).

REMARKS: Small larvae without striations on the buccal capsule.

Procamallanus (Procamallanus) sp.
SPECIMEN: MHNG INVE 74269.
HOSTS: Ageneiosus inermis* (SA 29/11/93).
REMARKS: New host record.

Procamallanus (Procamallanus) annipetterae Kohn et Fernandes, 1988
SPECIMENS: MHNG INVE 73341,73377, 73380.

HOST: Hypostomus sp. (VI 14/11/87). Hypostomus piratatu Weber, 1986* (CA
01/06/87).

REMARKS: New host and locality records (Central Province).

Procamallanus (Procamallanus) peraccuratus Pinto, Fâbio, Noronha et Rolas,

1976
SPECIMENS: MHNG INVE 73480, 73481, 73547, 73568, 73575, 73582, 73768, 73774,
73846, 73859.

HOSTS: Crenicichla lepidota (EP 01/11/87, AA 03/11/87, AT 23-25/10/89,
ES 05/10/89, SN 16/10/89). Hoplias malabaricus (SU 02/10/89). Rhamdia quelen*
(SP 22/09/89).

REMARKS: One new host record (R. quelen), although this fish in fact could
serve as paratenic or paradefinitive host (Moravec, 1998).

Procamallanus (Spirocamallanus) sp.
SPECIMENS: MHNG INVE 73330, 74223, 74224, 74356, 74390, 74395, 74423.
HOSTS: Loricariichthys sp. (ED 26/06/96). Loricariidae gen. sp. (ED 26/06/96).
Pimelodus cf. albicans (ED 28/09/96). Plagioscion ternetzi* (CA 30/05/87). Sorubim
lima (SA 27/11/93, ED 02/10/96).

REMARKS: Young females that might belong to P. (S.) pimelodi for pimelodid
hosts, or P. (S.) cervicalatus for loricartids.

Procamallanus (Spirocamallanus) cervicalatus (Petter, 1990)
SPECIMENS: MHNG INVE 73376, 74624.

HOSTS: Loricaria sp. (VI 14/11/87). Loricariichthys labialis* (EF 15/02/96).
REMARKS: New host and locality records (Central Province).

Procamallanus (Spirocamallanus) hilarii Vaz et Pereira, 1934

SPECIMENS: MHNG INVE 73615, 73726, 73849, 74213.

HOSTS: Astyanax sp. (AH 22/10/89). Rhamdia quelen (AR 13/10/91, ES
29/10/87, SN 05/10/89).

REMARKS: New geographical record.

478 D. GONZAÂLES-SOLIS & J. MARIAUX

Procamallanus (Spirocamallanus) inopinatus Travassos, Artigas et Pereira, 1928

SPECIMENS: MHNG INVE 73451, 73455, 73462, 73466, 73469, 73472, 73475, 713478,
73496, 73505, 73525, 73529, 73534, 73560, 73601, 73867, 73868, 73869, 73877, 73879, 74283,
74290, 74293, 74313, 74350, 74351, 7352, 74391, 74401, 74405, 74425, 74433, 74443, 74450,
74467, 74625, 74635, 74644.

HOSTS: Acestrorhynchus altus* (SN 17/10/89). Brycon sp. (SA 03/11/95).
Brycon hilarii (EP 31/10/87). Hoplias malabaricus (EG 20/11/87). Leporinus sp. (ED
03/08/94). Leporinus lacustris (SN 17/10/89). Leporinus obtusidens (ED 26/06/96, PI
05/01/96, EF15/02/96). Markiana nigripinnis* (EG 20/11/87, EM 06/12/96). Pellona
flavipinnis* (VI 13/11/87). Pygocentrus nattereri (VI 13/11/87, SA 04/10/96, PC
05/01/96, EF 15/02/96). Rhaphiodon vulpinus* (RS 09/10/91). Roeboides microlepis*
(VI 13/11/87). Serrasalmus sp. (ED 01/08/94). Serrasalmus nattereri (ED 02/10/96,
SA 04/10/96, EF 15/02/06). Serrasalmus spilopleura (ED 03/08/94, 02/10/96).
Trachydoras paraguayensis (Eigenmann & Ward, 1907) (VI 13/11/87). Triportheus
paranensis (EG 20/11/87).

REMARKS: Despite the wide host range of this species, five new host records are
reported herein.

Procamallanus (Spirocamallanus) krameri (Petter, 1974)
SPECIMEN: MENG INVE 73760.
HOST: Hoplerythrinus unitaeniatus (AT 25/10/89).
REMARKS: New geographical record.

Procamallanus (Spirocamallanus) paraensis Pinto et Noronha, 1976
SPECIMEN: MEING INVE 73604.
HOST: Rhaphiodon vulpinus* (RS 09/10/91).

REMARKS: New host and geographical records.

Procamallanus (Spirocamallanus) paraguayensis (Petter, 1990)

SPECIMENS: MHNG INVE 738838, 73891, 73892, 74447.

HOST: Hemiodus orthonops Eigenmann & Kennedy, 1903 (SN 17/10/89, GN
18/10/89, PC 04/01/96).

REMARKS: New locality record (Neembucu Province).

Procamallanus (Spirocamallanus) pimelodus Pinto, F4âbio, Noronha et Rolas, 1974

SPECIMENS: MHNG INVE 73346, 73498, 73609, 73612, 73636, 73638, 73639, 73641,
74285, 74301, 74386, 74396, 74647.

HOSTS: Pimelodella sp. (VI 13/11/87). Pimelodella griffini Eigenmann, 1917*
(AR 13/10/91). Pimelodus sp. (ED 01/08/94, 26/06/96, 26/09/96). Pimelodus cf. albi-
cans* (ED 03/08/94, ED 28/09/96). Pimelodus maculatus* (CA 02/06/87). Pla -
gioscion ternetzi* (RS 09/10/91).

REMARKS!: New host and geographical records.

Procamallanus (Spirocamallanus) rarus Travassos, Artigas et Pereira, 1928
SPECIMENS: MANG INVE 73306, 73616.
HOSTS: Jheringichthys sp.* (PV 16/02/87). Rhamdia quelen* (AR 13/10/91).
REMARKS: New host records.

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 479

FAMILY HEDRURIDAE NITZSCH, 1821
Hedruris Sp.
SPECIMENS: MHNG INVE 73861.,73876., 73882. 73884. 73881.

HOSTS: Acestrorhynchus altus* (SN 17/10/89). Hoplosternum littorale* (SN
17/10/89). Rhamdia quelen* (SN 16/10/89).

REMARKS: New host records, but probably as facultative hosts.

FAMILY GUYANEMIDAE PETTER, 1974
Guyanema baudi Petter et Dlouhy, 1985
SPECIMENS: MHNG INVE 73335, 729, 73779, 74219, 74347.

Hosts: Hoplias malabaricus (EG 18/11/87, ES 29/10/87, SU 03/10/89, AT
24/10/89, PO 14/08/96). Hypostomus piratatu* (CA 01/06/87).

REMARKS: New host record.

Guyanema seriei paraguayensis Petter et Dlouhy, 1985
SPECIMENS: MANG INVE 73758, 73772, 13776, 73781.

HOST: Hoplerythrinus unitaeniatus (AT 25/10/89, ES 01/10/89, 04/10/89, SU
03/10/89).

REMARKS: New locality (Itapua Province).

Guyanema seriei seriei Petter, 1974
SPECIMEN: MEING INVE 73720.

HOST: Hoplerythrinus unitaeniatus (ES 27/10/87).
REMARKS: New geographical record.

Travassosnema sp.
SPECIMEN: MANG INVE 73571.

HOST: Acestrorhynchus altus (AT 24/10/89).

REMARKS: Young female that probably belongs to the species 7ravassosnema
(ravassosi paranaensis.

Travassosnema travassosi paranaensis Moravec, Kohn et Fernandes, 1993
SPECIMENS: MHNG INVE 73446, 73563, 73570, 73571, 73847, 73870, 73873.
HOST: Acestrorhynchus altus* (EG 20/11/87, AT 23/10/89, ES 05/10/89, SN
17/10/89).

REMARKS: New host and geographical records.

FAMILY PHILOMETRIDAE BAYLIS ET DAUBNEY, 1926
Philometridae gen. sp.
SPECIMENS: MHNG INVE 73566, 73580, 73581.
HOST: Crenicichla lepidota (AT 23-24/10/89).

REMARKS: Males in poor condition, impossible to determine to the generic
level.

480 D. GONZALES-SOLIS & J. MARIAUX

FAMILY GNATHOSTOMATIDAE RAILLIET, 1895

Gnathostoma Sp.
SPECIMEN: MHNG INVE 73706.

HOST: Hoplias malabarius* (ES 26/10/87).
REMARKS: New host and geographical records.

FAMILY RHABDOCHONIDAE TRAVASSOS, ARTIGAS ET PEREIRA, 1928

Rhabdochona sp.

SPECIMENS: MHNG INVE 73537, 73573, 73606, 73614, 73617, 73624, 73778, 73838,
73843,73851,73892, 74179, 74183, 74311, 74360, 74389, 75033.

HOSTS: Auchenipterus nuchalis (Spix & Agassiz, 1829)* (GN 19/10/89).
Doradidae gen. sp.* (ED 26/06/96). Hemiodus orthonops* (GN 18/10/89).
Hoplerythrinus unitaeniatus* (SU 03/10/89, ES 06/10/89). Pimelodella griffini (AR
13/10/91). Pimelodus maculatus (SA 02/11/95). Plagioscion ternetzi* (RS 09/10/91).
Rhamdia quelen (ES 04/10/89, 05/10/89). Rhinodoras dorbignyi (Kner, 1855)* (AR
13/10/91). Trachelyopterus sp.* (AT 24/10/89). Trachelyopterus galeatus* (GN
05/11/91). Triportheus paranensis* (VI 13/11/87).

REMARKS: Small larvae that probably belong to different species. Pimelodid
fishes perhaps acts as accidental hosts (Moravec, 1998). Rhadochona kidderi kidderi
and R. k. texensis have been reported in Rhamdia guatemalensis; however, the larval
rhabdochonid found in R. quelen could represent a different species. This 1s the first
record of these larvae in nine new hosts.

Rhabdochona acuminata (Molin, 1860)
SPECIMENS: MHNG INVE 74187.

HOST: Leporinus friderici (Bloch, 1794) (GN 19/10/89).

REMARKS: This is the second record of R. acuminata in anostomid fishes in
South America and the first for this fish species. New geographical record.

Rhabdochona uruyeni Diaz-Hungria, 1968
SPECIMEN: MHNG INVE 73745.

HOST: Brachychalcinus retrospina Boulenger, 1892* (EP 02/11/87).
REMARKS: New host and geographical records.

FAMILY CYSTIDICOLIDAE SKRJABIN, 1946
Cystidicoloides sp.
SPECIMENS: MHNG INVE 73363, 73459, 73464, 73468, 73471, 73491, 73532
HOSTS: Markiana nigripinnis* (EG 20/11/87). Plagioscion ternetzi* (CA

23/08/87). Roeboides microlepis* (VI 13/11/87). Triportheus paranensis* (EG
20/11/87, VI 13/11/87).

REMARKS: Those larvae occurring in characids might belong to C. dlouhyi. This
is the first record of this genus in the marked fish species.

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 48 1

Spinitectus Sp.

SPECIMEN: MHNG INVE 73613.

HOST: Pimelodella griffini (AR 13/10/91).

REMARKS: Females that could belong to S. multipapillatus.
Spinitectus asperus Travassos, Artigas et Pereira, 1928

SPECIMEN: MHNG INVE 73540.

HOST: Prochilodus lineatus Steindachner, 1881* (VI 13/11/87).

REMARKS: New host and locality records (Central Province).

Spinitectus cf. pachyuri Petter, 1984
SPECIMENS: MHNG INVE 74178, 74182, 74209, 74449.

HOSTS: Auchenipterus nuchalis (GN 19/10/89, PC 04/01/96). Pachyurus sp.
(GN 19/10/89).

REMARKS: New locality record (Neembucu Province).

Spinitectus rodolphiheringi Vaz et Pereira, 1934

SPECIMENS: MHNG INVE 73704, 73737, 73741, 73743, 73744, 74170, 74172, 74180,
74211,74344.

HOSTS: Astyanax asuncionensis* (EP 01/11/87, 02/11/87). Auchenipterus
nuchalis* (GN 18/10/89). Hoplias malabaricus* (ES 26/10/87). Oxydoras kneri* (SA
04/11/95). Pachyurus sp.* (GN 18/10/89, 19/10/89).

REMARKS: All are new host records.

FAMILY PHYSALOPTERIDAE RAILLIET, 1893

Heliconema sp.
SPECIMEN: MHNG INVE 73652.

HOST: Oxyrhopus melanogenis?* (SL 30/07/88).

REMARKS: New host and geographical record.
Pseudoproleptus sp.

SPECIMENS: MHNG INVE 73722, 74431, 74435, 74650.

HOSTS: Hoplerythrinus unitaeniatus* (ES 27/10/87). Oxydoras kneri* (PC
05/01/96). Serrasalmus nattereri* (SA 04/10/96, EF 15/02/96).

REMARKS: This 1s the first finding of these nematodes in Paraguay, therefore all
represent new host and geograpical records.

FAMILY ACUARIIDAE RAILLIET, HENRY ET SISOFF, 1912
Acuariidae gen. sp.
SPECIMENS: MHNG INVE 73742, 74348.

HOST: Astyanax asuncionensis* (EP 01/11/87). Hoplias malabaricus* (PO
15/08/96).

REMARKS: New host and geograpical records.

482 D. GONZALES-SOLIS & J. MARIAUX

CONCLUSIONS

The nematodes reported herein showed different distribution range and hosts
diversity. The species whose adult presented the broadest geographical and host ranges
were À. rondoni (14 localities and 10 hosts) and Procamallanus (Spirocamallanus)
inopinatus (11 and 16). Whereas Rhabdochona sp. (10 and 12), Contracaecum Sp. type
1 (19 and 31), Contracaecum sp. type 2 (19 and 17), and Hysterothylacium sp. (9 and
27) were the larval or juvenile forms occurring in most localities and fishes.

A total of 150 new host and 44 geographical records were reported in this sur-
vey, thus increasing the number of known nematodes infecting fishes in Paraguay from
28 to 87.

Camallanidae was the best represented nematode family with 10 species,
followed by Anisakidae (9), Pharyngodonidae (7) and Cucullanidae (5). The nematode
fauna of the Paranä River in Paraguay is quite similar to that of Brazil and Argentina
because they share a common ichthyofauna, probably represent a single aquatic basin
and most parasitological studies of freshwater fishes have been carried out in the drai-
nage system of this basin (see Moravec, 1998, Vicente and Pinto, 1999, Takemoto er
al. 2009). Other factors contributing to this relatively poor parasite diversity are the
low degree of host specificity of these helminths as well as the phylogenetic rela-
tedness of the ichthyofauna considered in this study.

ACKNOWLEDGEMENTS

We thank the numerous collectors who participated to the EMGP between 1987
and 1996. DGS thanks the Muséum d'Histoire naturelle de Genève for invitation and
support during two stays in 2008 and 2010. This work was also partially supported by
the sabbatical stay of DGS provided by CONACYT and the Institute of Parasitology
of the Academy of Sciences of the Czech Republic (projects Nos. Z60220518 and
LC 5227

REFERENCES

FROESE, R. & PAULY, D. (ED.) 2011. FishBase. World Wide Web electronic publication.
www.fishbase.org, version (02/2011).

MASI-PALLARÉS, R. 1990. Nematodes de los vertebrados del Paraguay y otros paräsitos del
Nuevo Mundo. ECFACIM, Asunciôn, 689 pp.

MASI-PALLARÉS, R., BENITEZ-USHER, C. A. & VERGARA, G. 1973. Helminthes en peces y repti-
les del Paraguay (1era parte). Revista Paraguaya de Microbiologta 8: 67-96.

MORAVEC, F. 1998. Nematodes of freshwater fishes of the Neotropical region. Academia, Praha,
Czech Republic, 464 pp.

MORAVEC, F., KOHN, A. & FERNANDES, B. M. M. 1992. Nematode parasites of fishes of the
Paranä River, Brazil. Part 1. Trichuroidea, Oxyuroidea and Cosmocercoidea. Folia
Parasitologica 39: 327-353.

PETTER, À. J. 1984. Nématodes de poissons du Paraguay. II. Habronematoidea (Spirurida).
Description de 4 espèces nouvelles de la famille des Cystidicolidae. Revue suisse de
Zoologie 91: 935-952.

PETTER, À. J. 1989. Nématodes de poissons du Paraguay. V. Cucullanidae. Description de deux
espèces nouvelles et redéfinition du genre Neocucullanus Travassos ef al. Revue suisse
de Zoologie 96: 591-603.

NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 483

PETTER, À. J. 1990. Nématodes de poissons du Paraguay. VI. Description de deux nouvelles
espèces du genre Spirocamallanus et compléments à la description de Procamallanus
annipetterae Kohn & Fernandes, 1988. Revue suisse de Zoologie 97: 327-338.

PETTER, À. J. 1994. Nématodes de poissons du Paraguay. VII. Oxyuroidea: Spinoxyuris oxydoras
n.£., n. Sp. Revue suisse de Zoologie 101: 761-769.

PETTER, À. J. 1995a. Nématodes de poissons du Paraguay. VIII. Habronematoidea, Dracun-
culoidea et Ascaridoidea. Revue suisse de Zoologie 102: 89-102.

PETTER, À. J. 1995b. Dichelyne moraveci n. sp., parasite de Pseudoplatystoma fasciatum et notes
sur les Cucullanidae du Paraguay. Revue suisse de Zoologie 102: 769-778.

PETTER, À. J. & CASSONE, J. 1984. Nématodes de poissons du Paraguay. I. Ascaridoidea:
Sprentascaris n. gen. Revue suisse de Zoologie 91: 617-634.

PETTER, À. J. & DLOUHY, C. 1985. Nématodes de poissons du Paraguay. III. Camallanina.
Description d’une espèce et d’une sous-espèce nouvelles de la famille des Guyanemidae.
Revue suisse de Zoologie 92: 165-175.

PETTER, À. J. & MORAND, S. 1988. Nématodes de poissons du Paraguay. IV. Redescription de
Spinitectus jamundensis Thatcher et Padilha, 1977 (Cystidicolidae, Nematoda). Revue
suisse de Zoologie 95: 377-384.

SARAIVA, À., DA SILVA, F. À. & SILVA-SOUZA, A. T. 2006. Neocucullanus neocucullanus
Travassos, Artigas et Pereira, 1928 (Nematoda: Cucullanidae) from the Characidae fish,
Brycon hilarii Valenciennes, 1850, from Brazil. Memorias do Instituto Oswaldo Cruz
101(6): 669-672.

TAKEMOTO, R. M., PAVANELLI, G. C., LIZAMA, M. À. P., LACERDA, A. C. F., YAMADA, FE H.,
MOREIRA, L. H. A., CESCHINI, T. L. & BELLAY, S. 2009. Diversity of parasites of fish
from the Upper Paranä River floodplain, Brazil. Brazilian Journal of Biology 69(suppl.
2): 691-705.

TRAVASSOS, L. 1927. Uma nova Capillaria parasita de peixes de âgua doce: Capillaria sentinosa
n. sp. Boletin de Biologia 10: 215-217.

VICENTE, J. J. & PINTO, R. M. 1999. Nematôides do Brasil. Nemat6ides de peixes. Atualizacao:
1985-1999. Revista Brasileira de Zoologia 16(3): 561-610.

APPENDIX: List of localities. The two-lettres code in the first column is the abbreviation used in
the text. Latitudes and longitudes in decimal degrees.

Locality Province Latitude Longitude
AA Arroyo Allegre Concepcion -22.45 -57.60
AG Arroyo Tagatya Mi Concepcion -22.76 -57.59
AH Arroyo Hondo Caaguazu -25.13 -56.35
AM Arroyo Mborevi Presidente Hayes -23.36 -59.07
(Trans Chaco Km 303)
AP Arroyo Piratyi Canindeyu -24.07 -54.30
AR Arroyo Trementina Concepcion -22.82 -56.70
AT Arroyo Tapicuarai San Pedro -24.60 -56.45
AY Arroyo Yabebyry Itapua -27.31 -55.58
CA Campichuelo Itapua -27 43 -55.75
ED Estancia La Dorada Boqueron -22.71 -62.15
EF Estancia Farres (Pilcomayo) Presidente Hayes -25.07 -57.95
EG Estancia (Laguna) General Diaz Alto Paraguay -21.13 -58.50
EM Estancia Las Margaritas (Pilcomayo) Presidente Hayes -23.61 -60.44
EO El Dorado Corrientes -26.40 -54.70
EP Estancia Primavera Concepcion -22.45 -57.63
ES Estancia Santa Sofia Concepcion -22.33 -57.15
GN General Diaz Neembucu -27.27 -57.83
GY Guayrati Central -25.52 -57.50

JL Juan EO Leary Alto Paranä -25.42 -55.38

D. GONZALES-SOLIS & J. MARIAUX

Paso Correa

Puerto Edelira
Piquete-cué
Panchito Lopez
Pozo Arias

Puerto Oro Verde
Punto Zinho

R-Par 638

Rio Salado

San Antonio

San Lorenzo

San Lorenzo (10 Km SE)
Salto Pirareta

San Luis (11Km E)
Villeta

Yabebyry

Central
Itapua
Central
Misiones
Presidente Hayes
Itapua
Concepcion
Concepcion
Central
Central
Central
Neembucu
Cordillera
Concepcion
Central
Misiones

-25.12
-26.90
-25.12
-27.40
-23.63
-26.87
-22.38
-23.83
25 A2
-25.43
-25.30
-26.78
-25.55
-22.43
-25.52
-27.40

-57.45
-55.17
-57.50
512%
-60.32
-55.13
-56.93
-57.27
-57 45
-57.55
-57.50
-57.67
-56.90
-57.34
-57.50
=D el

REVUE SUISSE DE ZOOLOGIE 118 (3): 485-489; septembre 2011

Food habits of escaped Eurasian otters (Lutra lutra) in a suburban
environment in Switzerland

Jean-Marc WEBER, KORA, Thunstrasse 31, 3074 Muri, Switzerland.*
E-mail: imweber@bluewin.ch

Food habits of escaped Eurasian otters (Lutra lutra) in a suburban
environment in Switzerland. - In 2005, a male and a female otters escaped
from the zoo of Bern, and settled in the nearby River Aar. The number of
otters present in the area increased to 5 individuals after the adult pair
reproduced. À monitoring was launched in 2007 in order to examine how
these otters live in this suburban environment. Food habits notably were
investigated. Fish constituted the staple prey (91.5 %) with salmonids being
the most frequently eaten prey category (43.1 %). Seasonal dietary variation
occurred but was not marked. The results and the perspective of a long-term
survival of otters are discussed with regards to the overall decrease in fish
numbers recorded in the Swiss waters.

Keywords: Eurasian otter - Lutra lutra - diet - Switzerland.

INTRODUCTION

The Eurasian otter (Lutra lutra) is considered extinct in Switzerland since 1989
(Weber & Weber, 1991). The idea of reintroducing this species in the country emerged
amongst the nature authorities in the mid-1980s, but was quickly abandoned following
investigations on the feasibility of this project. The extremely high concentration of
PCBs in fish from the Swiss rivers was one factor amongst others that stopped the
reintroduction process (Weber, 1990a, 1992). Nearly 15 years later, in August 2005, a
male and a female captive Eurasian otters escaped from Bern zoological garden fol -
lowing a sudden inundation of their enclosure by the adjacent River Aar, and settled in
the vicinity of the zoo. In the absence of any reintroduction program, the capture of
these individuals was decided by the federal and cantonal authorities. In 2007, the male
was trapped and the female died from injuries of unknown origins. However, several
direct observations of otters and genetic analyses of otter faeces (‘“spraints”) collected
in the otters’ core area confirmed that the escapees reproduced at least once in the
meantime (Weber, 2008). One of the three identified cubs, a male, was also captured
in 2007, leaving a young male and a female in the wild.

An extensive monitoring Was launched to examine how these otters adapt to
their new environment. Their feeding habits were the focus of this study. Indeed,
regarding food availability only, with a total fish biomass of more than 100 Kg/ha, the
River Aar in the neighbourhood of Bern was considered a suitable potential habitat for
otters (Weber, 1997). However, evidences such as a generalized and drastic decline of

Manuscript accepted 10.06.2011
* present address: SFEN, Section Faune, Rue du 1er-Mars 11, 2108 Couvet, Switzerland.

486 J.-M. WEBER

fish catch in Swiss rivers, including the River Aar, since the 1990s suggest a possible
deterioration of the potential trophic conditions for otters (Burkhardt-Holm et al.
2002), but the exact consequences on otter’s diet and more generally on their survival
is unknown. In this context, we aim to sketch a first figure of otter’s feeding habits
following their accidental reintroduction in the wild.

STUDY AREA

Escaped otters were living essentially on an 8 km-long stretch of the River Aar,
3 km south-east of the city of Bern (7°44°94°E; 46°93°33"°N). Three small tributaries,
wetlands and a few ponds adjoin the main river, and an undisrupted riparian forest is
present on both sides of it. More than 40 fish species occur in the region. In the absence
of recent fish biomass counts (1.e. electro-fishing) in the River Aar, the fishing bag was
used to give an index of fish numbers. Thus, barb (Barbus barbus), trout (Salmo trutta
Jario), grayling (Thymallus thymallus), perch (Perca fluviatilis) and pike (Esox lucius),
respectively, are the most frequently angled fish in the area (Fish Management Service
of the Canton Bern, pers. com.). Two species of crayfish, a potential alternative prey to
otters, live also in the area (Stucky & Zaugg, 2005). The river flows in a depression
surrounded by the suburbs of Bern. Therefore, human (e.g. walkers, joggers, swimmers
and bikers) presence 1s important year-round along the stream. À commercial fish farm
raising trout also occurs in the area.

MATERIAL AND METHODS

Known sprainting sites in the study area were checked twice each month, from
November 2007 to May 2010, and all otter faeces were collected. Faeces were
dissected and analysed as described in Jenkins ef al. (1979). Fish and amphibian
remains, mostly scales and bones, were identified according to Conroy ef al. (1993).
The results of the analyses were expressed in relative frequency of occurrence (1.e.
occurrence of a prey item in the spraints/total number of prey items identified in the
spraints x 100) and sorted by season: spring (March — May), summer (June-August),
autumn (September — November) and winter (December — February).

RESULTS

A total of 182 spraints was collected and 246 prey items identified. With 91.5
% of relative frequency of occurrence, fish dominated the diet (Table 1). Amphibians
represented only 4.5 % of the food items, while other prey categories were occasio-
nally taken by otters. Considering fish, salmonids were of prime importance (43.1 %),
followed by cyprinids (21.2 %), bullhead (Cottus gobio; 13 %) and pike (11 %). The
consumption of perch was anecdotal (1.6 %).

Salmonids were otters’ staple prey in every season (Table 2). Their consump-
tion by otters did not show any major seasonal variation (Chi? test = 5.1394, p > 0.05).
Cyprinid exploitation tended to decrease in winter, although the trend was also not
significant (Chi? test = 2.1997, p > 0.05). In contrast, pike occurred more frequently in
the diet during winter than in other seasons (Chi? test = 12.9634, p < 0.01). Predation
on bullhead remained stable along the year.

DIET OF ESCAPED EURASIAN OTTERS 487

TABLE 1. Diet of otters in the River Aar expressed in relative frequency of occurrence (RFO) in
182 scats analysed. N: number of prey items.

N RFO (%) ,

Salmonidae 106 43.1
Undetermined Cyprinidae 42 17.1
Minnow (Phoxinus phoxinus) 10 4.1

Bullhead (Cottus gobio) 32 13.0
Pike (Esox lucius) 24 11.0
Perch (Perca fluviatilis) 4 1.6

Undetermined Fish 4 1.6

Total Fish 223 91.5
Arthropods 2 0.8
Amphibians 11 4.5
Reptiles + 1.6
Birds 2 0.8
Mammals Il 0.4
Undetermined Il 0.4
Total 246 100

TABLE 2. Seasonal variation in the different prey categories (%) found in otters’spraints from the
River Aar. Number of items counted in spraints are in parentheses.

Spring (105) Summer (18) Autumn (90) Winter (33)
Salmonidae 38.1 44 4 3533 30.3
Cyprinidae 19.1 33,2 24.4 124
Cottidae 13,3 LA 13.3 121
Esocidae | 2h92 0 3.4 24.3
Amphibia 6.7 0 2:2 6.0
Other prey 7.6 13 34 552

DISCUSSION

Both habitat type and latitude play a role in the composition of otter’s diet in
Europe (Jedrzejewska ef al., 2001, Clavero et al., 2003). According to these studies, a
fish-dominated diet could be expected in central European waters, what is confirmed
by our results, although Eurasian otters adapt their feeding behaviour to local
conditions, 1.e. according to prey availability and abundance (Kruuk, 2006). Food
composition should, therefore, reflect which prey is readily available to otters in our
study area. Investigations carried out on the feeding habits of reintroduced otters in
1975 in the River Schwarzwasser, a tributary of the River Aar 11 km to the SW of
current study area, gave a figure similar to our findings. Fish clearly dominated the diet
(79.5 % relative frequency of occurrence, N = 127; Weber et al., 1991). Moreover,
otters preyed mainly on salmonids (48 %), while cyprinids occurred in the same
proportion as in the River Aar (23.6 %). The only major differences with our results
were a much higher presence of amphibians (22.8 %), the absence of pike in the food
spectrum and the lack of marked seasonal variation in fish consumption. As in the Aar,
amphibians in the Schwarzwasser area tended to be more preyed upon during winter

488 J.-M. WEBER

(27 %) than in summer (10 %), a trait generally observed where otters exploit this type
of prey (see e.g. Weber, 1990b, Lanzski ef al., 2001, Brzezinski ef al., 2006).

Would the River Aar offer a suitable habitat for otters as suggested by Weber
(1997)? A continued 5-yr presence emphasized by at least two breeding attempts - the
adult female was lactating again when she died (M.-P. Ryser, pers. comm.) - suggests
that otters found an appropriate habitat to survive in that region. However, repro-
duction may not be the sole criterion to ensure a long-term survival, as experienced
with the reintroduced otters of the Schwarzwasser. Indeed, the introduced animals
eventually disappeared from the area approximately 10 years after their release, despite
successful reproduction (Weber ef al., 1991). The suboptimal fish biomass (50 - 100
Kg/ha) recorded in the Schwarzwasser area could explain this failure (Weber, 1990a).
As already mentioned, the River Aar is not an exception to the global decline of fish
catch observed in the Swiss waters (Burkhardt-Holm ef al., 2002). In the study area,
the total weigh of fish catch decreased from 1818 kg in 1996 to 1141 kg in 2006
(- 37.2 %). The bag shrunk from 1569 kg to 609 kg for salmonids only during the same
period (- 61.2 %; Fish Management Service of the Canton Bern, pers. com.). Under
these circumstances, accidentally released otters are possibly confronted to suboptimal
trophic conditions, as far as their main prey, 1.e. salmonids, are concerned. However,
the fish farm present in the area may have served as a secondary food source. Several
observations of intruding otters have been reported and otter sprainting sites were
numerous in the vicinity of the farm (Weber, 2008). However, its role as food supplier
would have remained very local and time-limited, particularly 1f the owners would
prevent the access to the farm to otters. In addition to possible unfavourable local food
conditions, human activities impact considerably the area which may have hampered
future population development (Clavero ef al., 2011). Finally, surveys carried out in
2010 show an important proportion of abandoned sprainting sites in the study area,
suggesting a possible disappearance of one of the remaining individuals (J.-M. Weber,
unpublished observations).

According to these opportunistic observations gathered following the accidental
release of two otters, a perennial presence of otters in the River Aar seems unlikely, as
habitat quality, and more particularly trophic conditions, are probably not optimal
enough to allow a long-term survival of otters. Major improvement of the environment,
and especially of food availability, seem to remain a pre-requisite to any otter release
in the area and more generally in the Swiss waters, as suggested in earlier reports (e.g.
Weber, 1990a).

ACKNOWLEDGEMENTS

I would like to thank the Federal Office of Environment for financial support,
Ueli Iff and Dr Sandra Schorderet Weber for field assistance, Dr Marc Rosset for
documenting the escape of the otters, and two anonymous referees for useful
comments on the manuscript.

DIET OF ESCAPED EURASIAN OTTERS 489

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WEBER, J.-M. & WEBER, D. 1991. The otter (Lutra lutra L.) in western Switzerland, 1989
(pp. 53-55). /n: REUTHER, C. & RÔCHERT, R. (eds), Proceedings of the V. International
Otter Colloquium. Habitat 6, Hankensbüttel, 344 pp.

WEBER, D., WEBER, J.-M. & MÜLLER, H.-U. 1991. Fischotter (Lutra lutra L.) im Schwarz -
wasser-Sense-Gebiet: Dokumentation eines gescheiterten Wiedereinbürgerungsver-
suchs. Mitteilungen der Naturforschenden Gesellschaft in Bern 47: 141-152.

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REVUE SUISSE DE ZOOLOGIE 118 (3): 491-512; septembre 2011

Claude Besuchet, an eminent swiss coleopterists, 80 years old

Peter HLAV AC
Na doline 14, SK-040 14 Koëice, Slovakia. E-mail: phlavac@stonline.sk

INTRODUCTION

As a schoolboy I was, as many others, interested in beetles, I collected large and
atractive beetles such as Cerambycidae, Buprestidae, Carabidae etc. but this was only
for a relatively short period and when I entered grammar school I started to be
interested in many other things. It was almost 20 years later, when, as a student of the
University of Pierre and Marie Currie in Paris, [ visited the Museum of Natural
History in Paris and especially the book shop of this institution. I was browsing shelves
full of very nice books on the nature, birds, mammals when I came to the insect section
and I found the book which immediately fascinated me and attracted my full attention:
Faune de France, Pselaphidae by René Jeannel. I was enchanted by a beauty of this
small beetles. I immediately, although it was at that time horribly expensive for me,
bought it and decided that this will be my hobby and I will start to study the taxonomy
of Pselaphidae. Later I added to my sphere of interest Scydmaenidae and myrme-
cophilous beetles of other families but that is an another story. Since my early begin-
ning it Was just only short period of time until [ made contact with all actively work-
ing Pselaphidologists and started to search for reprints of their papers. Thus in 19941]
wrote to Claude Besuchet hoping he would send some reprints to me, a totally un-
known beginner from small east European country. After two weeks I received a large
package from Geneva. I openned it and it contained almost all Claude Besuchet’s
papers together with nice encouraging letter. “Soyez les bienvenus parmi les entomo-
logistes étudiants les Psélaphides et Scydmaenides” (“Welcome between entomolo-
gists studying Pselaphinae and Scydmaenidae”) these were the first words of the letter
and since then Mr. Besuchet and I have met many times, and for me it is always an
event to discuss on Pselaphines with him and to draw from his never ending knowledge
of this tiny but very beautifull beetles. So, Mr. Besuchet, thanks for the invitation and
the opportunity for me to remind to all entomologists a little about your 80 years of
life of which 63 were very much devoted mainly to your studies of Pselaphines.

BIOGRAPHY

Claude Besuchet was born on 4th July 1930 in Lausanne on the border of Lake
Geneva. The rich and wonderful natural history of this area very soon had attracted
his attention and Claude started to be interested in beetles in 1942 when he was 12
years old. Soon after, on 15th March 1947 he collected his first Pselaphine, 7rimium
brevicorne (Reichenbach). This little event predetermined his further carrier and life
orientation and we can say that rest of his life was devoted to study of these small,
wonderfull creatures. After he had finished his studies at the scientific grammar school
in Lausanne, he naturally continued at the Faculty of Science, University of Lausanne.

Manuscript accepted 25.02.2011

492 P. HLAVAC

In 1954 he graduated with the certificate for Physiology, Bacteriology, Parasitology,
Hygiene and Zoology, consequently he obtained the certificate of pedagogy in 1955
and in 1956 he successfully passed his PhD. Claude started his professional career at
the University of Lausanne where he had been working for short time till 1956. But the
name of Claude Besuchet will be forever connected to the Muséum d'Histoire naturelle
de Genève, where he has been working at the department of entomology since IS!
January 1958 until his retirement at 315$! July 1992. After retiring, Claude remained
active and took the responsibility for compilation of the catalogue of beetles of
Switzerland with the financial help of Fonds national suisse de la Recherche scien-
tifique, Centre suisse de Cartographie de la Faune and Ligue suisse pour la Protection
de la nature. As a consequence of his long service of scientific excellence at Geneva,
in 2005 Claude was appointed "Conservateur honoraire du Muséum d’Histoire
naturelle de Genève”.

Claude Besuchet’s contribution to our knowledge of many groups of beetles
but mainly Pselaphines is monumental, maybe not so much concerning to the quantity
of described taxa but mainly the quality of his work. Claude himself or with his
coauthors published all in all 155 papers, from this 126 purely scientific taxonomic
papers devoted to Staphylinidae (Pselaphinae), Scydmaenidae, Ptuliidae, Leptinidae,
Cerylonidae, Rhipiporidae and Dryopidae. He also published 14 faunistics papers,
mainly on endogean, cavernicolous or rare beetles of Switzerland, and 15 popular
papers about more general aspects of zoology.

He named in total 435 taxa (Table 1), comprising Pselaphinae (360),
Scydmaeninae (30), Ptiliidae (6), Leptinidae (2), Cerylonidae (35), Rhipiporidae (1)
and Dryopidae (1). the majority of these taxa are from the Palaearctic region (350),
followed by the Oriental region (53), Afrotropical region (17), Neotropical region
(10), Australian region (3) and one each from the Nearctic and Oceania.

Table 1. Analysis of Species described by Claude Besuchet
PAL AFR NEA NEO ORT AUS Oceania >

Pselaphidae 592 Il Il 2 14 2 Il 360
Scydmaenidae 10 16 0 0 à Il 30
Ptiliidae 6 6
Leptinidae 2 | 2
Cerylonidae 35 35
Rhipiporidae 1 1
Dryopidae Il 1
TOTAL 350 L7 1 10 5) 3 1 435

Cerylonidae: Interest of Claude on cerylonid beetles started with his splendid work
on the Aculagnathides (1972) and was later developed when Stanislav Adam Slipiñski,
one of most prominent specialists on Cucujoidea, visited Geneva museum a few times.
The result of this fruitfull cooperation was two important revisions, Glyptolopus
Erichson (1987b) and Axiocerylon Grouvelle (1988b).

CLAUDE BESUCHET'S WORK 493

FIG. 1
Claude Besuchet (left) and Ivan LGbl (right) at the Natural History Museum of Geneva.

Dryopidae: Besuchet published only one paper (1978e) on this family and described
a new monotypic terrestrial genus Geoparnus Besuchet.

Rhipiporidae: Besuchet’paper (19564) is still one of the most important studies on the
biology, morphology and taxonomy of Rhipiphoridae although it deals only with the
genus Rhipidius Thunberg. Further, Besuchet published another two papers (1956e,
1957) in which he described one new genus and one new species.

Leptinidae: The revision of the genus Leptinus (Besuchet, 1980e) is so far the most
important taxonomical study of the small sub-familly Platypsilinae (at the time of
publication known as Leptinidae), further Besuchet’s paper on this sub-family records
the presence of Platypsyllus castoris in Switzerland (Besuchet, 1978d).

Ptiliidae: The most important contribution is the key of the central Europen species
(Besuchet & Sundt, 1971c) where six new species and one new genus was also
described (later synonymized with Prinella Motschulsky). Another two species were
described in 1980e and last paper on this family of minute beetles 1s a list of Ptiliidae
from Mongolia collected by Zoltän Kaszab (1969h).

Staphylinidae: Scydmaeninae: The main interest in the subfamily Scydmaeninae of
Claude Besuchet was the tribe Cephennini although he described also other taxa. AI

494 P. HLAVAC

in all in tvelve papers (Besuchet, 1958e, 1959b, 1961h, 1962c, 1971a,b, 1980d, 1981c,
20044, 2004f: Besuchet & Vit, 2000b, 2004b) he named 32 species, from which 31
were new.

Staphylinidae: Pselaphinae: There is no doubt that the name Claude Besuchet will be
forever mainly connected to the subfamily Pselaphinae where he is placed on highest
pedestal together with Achile Raffray, René Jeannel and Orlando Park. Besuchet
named 14 genera (two later synonymyzed ) and 360 species (22 later synonymyzed) of
the subfamily Pselaphinae. But more important than just a number of described taxa is
the revisional approach of Claude to the problem of Pselaphinae. He brought taxo-
nomic order to the subfamily for the fauna of the Palearctic region. Much less attention
he paid to the tropical fauna, only 27 species Were named by him from tropical regions,
single species from the Nearctic region and 332 species from the Palearctic region.

LIST OF TAXA NAMED BY CLAUDE BESUCHET

I. List of tribes and subtribes:

1. Tiracerini Besuchet, 1986a: 263 (Staphylinidae: Pselaphinae)
2. Colilodionini Besuchet, 1991: 514 (Staphylinidae: Pselaphinae)

II. List of Genera:

1. Antrobythus Besuchet, 1985a: 511 (Staphylinidae: Pselaphinae)
Aphiliopsis Besuchet, 1956a: 369 (= Aphilia Reitter) (Staphylinidae:
Pselaphinae)

3 Bathybythus Besuchet, 1974b: 41 (Staphylinidae: Pselaphinae)

4. Cautomus Sbg. Leptoxycheilus Besuchet, 1972: 127 (Cerylonidae)
5. Colilodion Besuchet, 1991: 500 (Staphylinidae: Pselaphinae)
6
fi
8

Se

Couloniella Besuchet, 1983a: 509 (Staphylinidae: Pselaphinae)
Geoparnus Besuchet, 1978: 705 (Dryopidae)
. Nonveillera Paviéevië & Besuchet, 2003a: 279 (Staphylinidae: Pselaphinae)
9. Pachacuti Besuchet, 1987a: 231 (Staphylinidae: Pselaphinae)

10. Paratychus Besuchet, 1960a: 24 (Staphylinidae: Pselaphinae)

11. Plitium Besuchet, 1971c: 329 (= Prinella Motschulsky) (Ptilidae)

12. Pselaphotrichus Besuchet, 1986: 259 (Staphylinidae: Pselaphinae)

13. Pirhidius Besuchet, 1957: 24 (Rhipiphoridae)

14. Tapas Besuchet, 2008a: 74 (Staphylinidae: Pselaphinae)

15. Tasmiger Besuchet, 2008a: 78 (Staphylinidae: Pselaphinae)

16. Thelotia Besuchet, 1999b: 793 (Staphylinidae: Pselaphinae)

17. Tiracerus Besuchet, 1986a: 262 (Staphylinidae: Pselaphinae)

18. Tremissus Besuchet, 1982d: 317 (Staphylinidae: Pselaphinae)

IT. List of species and subspecies of Staphylinidae: Pselaphinae:

1. Acetalius pilosus Besuchet, 1985b: 763
2. Afropselaphus breiti Besuchet, 1961a: 34 (Pselaphogenius)
3. Afropselaphus canariensis Besuchet, 1968: 291

CLAUDE BESUCHET’S WORK 495

Afropselaphus circassicus Besuchet, 1961a: 37 (Pselaphogenius)
Afropselaphus fernandezi Besuchet, 1968: 290

Afropselaphus guanche Besuchet, 1970a: 123

Afropselaphus maroccanus Besuchet, 1963b: 222 (Pselaphogenius)
Afropselaphus spinipalpis Besuchet, 1968: 289

Afropselaphus zacynthius Besuchet, 1961a: 35 (Pselaphogenius)
Amauronyx auberti Besuchet, 1962b: 339

Amauronyx caecus Besuchet, 1962b: 340

Amauronyx caudatus Besuchet, 1999b: 798

Amauronyx cobosi Besuchet, 1959a: 26

Amauronyx franzi Besuchet, 1958e: 908

Amauronyx mussardi Besuchet, 1963c: 229

Amauronyx myops Besuchet, 1962b: 337

Amaurops aubei binaghii Besuchet, 1980b: 615

Antrobythus leclerci Besuchet, 1985a: 512

Antrobythus perplexus Besuchet, 1993b: 223

Bathybythus bleyi Besuchet, 1974b: 43

Batrisodes bifossulatus Besuchet, 1988a: 436

Batrisodes clypeatus Besuchet, 1981a: 290

Batrisodes mitovi Besuchet & Bekchiev, 2007b: 75

Batrisodes rousi Besuchet, 1981a: 292

Batrisodes sulcaticeps Besuchet, 1981a: 289

Batrisodes unisexualis Besuchet, 1988a: 433

Batrisus taurus Besuchet, 2004e: 28 new name for fauricus Besuchet, 1979: 279
Bibloplectus (s.str.) atomus Besuchet, 1958e: 906

Bibloplectus (s.str.) aberrans Besuchet, 1958e: 903

Bibloplectus (s.str.) beaumonti Besuchet, 1955a: 200
Bibloplectus (s.str.) boveyi Besuchet, 1975a: 32

Bibloplectus (s.str.) difficilis Besuchet, 1955a: 177

Bibloplectus (s.str.) elegans Besuchet, 1955a: 183

Bibloplectus elongatus Besuchet, 1953: 231 (= B. strouhali Beier)
Bibloplectus (s.str.) franzi Besuchet, 1964a: 411

Bibloplectus (s.str.) hellenicus Besuchet, 1955a: 194

Bibloplectus (s.str.) hungaricus Besuchet, 1955a: 191
Bibloplectus (s.str.) jeanneli Besuchet, 1955a: 186

Bibloplectus (s.str.) liliputanus Besuchet, 1975a: 33

Bibloplectus linderi Besuchet, 1953: 228 (= tenebrosus Reïtter)
Bibloplectus (s.str.) machulkai Besuchet, 1955a: 187

Bibloplectus minutus Besuchet, 1953: 229 ( = B. obtusus Guillebeau)
Bibloplectus (s.str.) normandi Besuchet, 1955a: 196

Bibloplectus (s.str.) parvulus Besuchet, 1975a: 34

Bibloplectus (s.str.) pauxillus Besuchet, 1975a: 34

Bibloplectus (s.str.) perroti Besuchet, 1955a: 176

Bibloplectus pseudambiguus Besuchet, 1953: 226 (= B. spinosus Raffray)
Bibloplectus (s.str.) subtilis Besuchet, 1975a: 35

496

49.
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31:
32.
53.
54.

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83.
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87.
88.
89.
90.

P. HLAVAC

Bibloplectus (s.str.) tantulus Besuchet, 1975a: 35
Bibloplectus (s.str.) tener Besuchet, 1975a: 33
Bibloplectus therondi Besuchet, 1953: 230 (= B. pusillus Denny)
Bibloporus bicolor franzi Besuchet, 1958e: 903
Bibloporus myops Besuchet, 1970b: 313
Brachygluta abrupta septemtrionalis Besuchet, 1963a: 34
(= B. abrupta Dodero)

Brachygluta alpina Besuchet, 2004a: 109
Brachygluta atlantica Besuchet, 2004a: 154
Brachygluta exigua Besuchet, 1963a: 41
Brachygluta exsculpta Besuchet, 1969e: 405
Brachygluta franciscae Besuchet, 1963a: 35
Brachygluta gnosiaca Besuchet, 2004a: 58
Brachygluta hanseni Besuchet, 1954c: 43 (= B. paludosa Peyron)
Brachygluta hispana Besuchet, 1963a: 40
Brachygluta jordanica Besuchet, 2004a: 78
Brachygluta kurdica Besuchet, 2004a: 144
Brachygluta lefebvrei meridionalis Besuchet, 1962b: 356
(= B. lefebvrei lederi Saulcy)

Brachygluta occidentalis Besuchet, 1963a: 35
Brachygluta perissinottoi Besuchet, 1969e: 402
Brachygluta pusilla Besuchet, 1958d: 335
Brachygluta richteri Besuchet, 1961e: 1

Brachygluta sengleti Besuchet, 1969e: 404
Brachygluta tumidipes Besuchet, 1981b: 243
Brachygluta ultima Besuchet, 2004a: 80
Brachygluta vicaria Besuchet, 1963a: 43

Bryaxis abkhasicus Besuchet & Kurbatov, 2007: 202
Bryaxis adjaricus Besuchet & Kurbatov, 2007: 202
Bryaxis adumbratus Besuchet & Kurbatov,2007: 172
Bryaxis altivagus Besuchet, 1962b: 354

Bryaxis arnoldii Besuchet, 1961£g: 1830

Bryaxis artvinensis Besuchet & Kurbatov, 2007: 179
Bryaxis assingi Besuchet & Kurbatov, 2007: 178
Bryaxis atlanticus Besuchet, 1962b: 353

Bryaxis badius Besuchet, 1961g: 1827

Bryaxis balabanus Besuchet & Kurbatov, 2007: 188
Bryaxis balneator Besuchet & Kurbatov, 2007: 193
Bryaxis bergamascus breiti Besuchet, 1980d: 625

(= B. bergamascus sorinensis Stolz)

Bryaxis borckensis Besuchet & Kurbatov, 2007: 169
Bryaxis brachati Besuchet, 1980b: 624

Bryaxis corsus Besuchet, 1999b: 799

Bryaxis credibilis Besuchet & Kurbatov, 2007: 182
Bryaxis distinguendus Besuchet, 19612: 1830

91.
92.
91.
94.
95.
96.
97.
96.
w.
100.
101.
102.
103.
104.
105.
106.
107.
108.
109.
110.

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113.
114.
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116.
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119.
120.
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122.
23.
124.
123.
126.
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128.
129.
130.
131.
172.
PT3.
134.
135.

CLAUDE BESUCHET’S WORK 497

Bryaxis effeminatus Besuchet, 1983b: 772

Bryaxis egens Besuchet & Kurbatov, 2007: 196
Bryaxis emendatus Besuchet & Kurbatov, 2007: 177
Bryaxis euryscapus Besuchet & Kurbatov, 2007: 164
Bryaxis festivus Besuchet, 1964a: 425

Bryaxis focarilei Besuchet, 1980b: 623

Bryaxis frustratus Besuchet, 1983b: 770

Bryaxis gemellus Besuchet & Kurbatov, 2007: 202
Bryaxis ghilarovi Besuchet, 19612: 1829

Bryaxis halbherri pacei Besuchet, 1983b: 777
Bryaxis herculinus Besuchet, 1962b: 351

Bryaxis immodicus Besuchet & Kurbatov, 2007: 199
Bryaxis ipsimus Besuchet & Kurbatov, 2007: 182
Bryaxis jucundus Besuchet, 19612: 1830

Bryaxis karamane Besuchet, 1958e: 895

Bryaxis khnzoriani Besuchet, 1964a: 423

Bryaxis kovali Besuchet & Kurbatov, 2007: 193
Bryaxis kruegeri Var. machulkai Besuchet, 1955: 277 (= B. kruegeri Machulka)
Bryaxis kurnakovi Besuchet, 1960b: 398

Bryaxis kuzmini Besuchet & Kurbatov, 2007: 194
Bryaxis laevipennis Besuchet & Kurbatov, 2007: 181
Bryaxis lazistanicus Besuchet & Kurbatov, 2007: 198
Bryaxis litoralis Besuchet & Kurbatov, 2007: 177
Bryaxis longifrons Besuchet & Kurbatov, 2007: 173
Bryaxis longulus inflatus Besuchet, 1983b: 778
Bryaxis lurensis Besuchet, 2002: 212

Bryaxis mirificus Besuchet, 1983b: 773

Bryaxis monguzzii Besuchet, 1980b: 622

Bryaxis multiplex Besuchet & Kurbatov, 2007: 199
Bryaxis myops Besuchet & Kurbatov, 2007: 199
Bryaxis nebrodensis Besuchet, 1980b: 619

Bryaxis nitidulus Besuchet, 19612: 1829

Bryaxis nivarius Besuchet & Kurbatov, 2007: 168
Bryaxis obventicius Besuchet & Kurbatov, 2007: 174
Bryaxis orcinus Besuchet & Kurbatov, 2007: 168
Bryaxis osellai Besuchet & Kurbatov, 2007: 190
Bryaxis oseticus Besuchet & Kurbatov, 2007: 196
Bryaxis ossaeus Besuchet, 2008b: 245

Bryaxis pachyscelis Besuchet & Kurbatov, 2007: 170
Bryaxis pedemontanus Besuchet, 1958b: 74

Bryaxis polemon Besuchet & Kurbatov, 2007: 189
Bryaxis ponticus Besuchet & Kurbatov, 2007: 187
Bryaxis porzenna var. ticinensis Besuchet, 1954: 436 (= B. porzenna Reitter)
Bryaxis propinquus Besuchet & Kurbatov, 2007: 186
Bryaxis pulchrotibialis Besuchet & Kurbatov, 2007: 172

498 P. HLAVAC

136. Bryaxis pygmaeus Besuchet & Kurbatov, 2007: 200
137. Bryaxis rifensis Besuchet, 1962b: 351
138. Bryaxis rivularis Besuchet & Kurbatov, 2007: 181
139. Bryaxis rousi Besuchet & Kurbatov, 2007: 200
140. Bryaxis scrutandus Besuchet & Kurbatov, 2007: 176
141. Bryaxis seductus Besuchet & Kurbatov, 2007: 186
142. Bryaxis scherleri Besuchet, 1964a: 421 (= B. judicariensis Dodero)
143. Bryaxis schuelkei Besuchet & Kurbatov, 2007: 203
144. Bryaxis silvicola Besuchet & Kurbatov, 2007: 187
145. Bryaxis solarii Besuchet, 1958c: 8 (= B. rhinophorus W. Blattnÿ & C. BlattnŸ)
146. Bryaxis temporalis Besuchet & Kurbatov, 2007: 197
147. Bryaxis tendensis Besuchet, 2002: 212
148. Bryaxis tenuicornis Besuchet & Kurbatov, 2007: 171
149. Bryaxis tingitanus Besuchet, 1962b: 352
150. Bryaxis transitorius Besuchet & Kurbatov, 2007: 179
151. Bryaxis troglodyÿtes pierottii Besuchet, 1980b: 628
152. Bryaxis tuberculiceps Nonveiller, Paviéevié & Besuchet, 2003b: 287
153. Bryaxis viti Besuchet & Kurbatov, 2007: 163
154. Bryaxis ypsilon Besuchet & Kurbatov, 2007: 169
155. Bythinus confusus Besuchet, 1974c: 337
156. Bythinus hauseri Besuchet, 1978a: 263
157. Bythinus icariensis Besuchet, 1964a: 420
158. Bythinus vicinus Besuchet, 1960a: 21
159. Centrophthalmus klapperichi Besuchet, 1966b: 63
160. Centrophthalmus mesopotamenus Besuchet, 1966b: 64
161. Centrophthalmus sharpi Besuchet, 1966b: 61
162. Centrophthalmus septentrionalis Besuchet, 1960a: 30 (= C. pici Jeannel)
163. Centrotoma kaszabi Besuchet, 1969f: 301
164. Claviger intermedius Besuchet, 1961c: 457
165. Claviger pouzaui validus Besuchet, 1961c: 453
166. Claviger saulcyi lucens Besuchet, 1961c: 455
167. Colilodion incredibilis Besuchet, 1991: 503
168. Colilodion mirus Besuchet, 1991: 506
169. Colilodion inopinatus Besuchet, 1991: 507
170. Colilodion concinnus Besuchet, 1991: 509
171. Couloniella mirabilis Besuchet, 1983a: 510
172. Decatocerus bicornis rotundatus Besuchet, 1961d: 95
(= D. balearicus Jeannel, 1961)
173. Decatocerus catalonicus Besuchet, 1961d: 94
174. Decatocerus pityusensis Besuchet, 1958e: 909
175. Desimia longicornis Besuchet, 1958d: 337 (= D. subtilipalpis Reitter)
176. Dicentrius balcanicus balcanicus Besuchet, 1999c: 229
177. Dicentrius balcanicus pirinensis Besuchet, 1999c: 229
178. Dicentrius behnei Besuchet, 1999c: 228
179. Dicentrius biroi Besuchet, 1999c: 230

180.
181.
182.
183.
184.
185.
186.
187.
188.
189.
190.
191.
192.
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199.
200.
201.
202.
203.
204.
205.
206.
207.
208.
209.
210.
211.
212,
283,
214.
244.
216.
217.
218.
DER.
220.
221.
222.
223.
224.

CLAUDE BESUCHET'S WORK 499

Dicentrius discrepans Besuchet, 1999c: 231
Dicentrius ermischi Besuchet, 1999c: 231
Dicentrius fodori Besuchet, 1999c: 228
Dicentrius rousi Besuchet, 1999c: 231
Dicentrius zerchei Besuchet, 1999c: 228
Euplectus atlanticus Besuchet, 1962b: 346
Euplectus caecus Besuchet, 1990: 161
Euplectus canariensis Besuchet, 1968: 285
Euplectus franzi Besuchet, 1968: 284
Euplectus frater Besuchet, 1964a: 415
Euplectus insignis Besuchet, 1961a: 30
Euplectus kulzeri Besuchet, 1958d: 334
Euplectus micropterus Besuchet, 1970a: 120
Euplectus mussardi Besuchet, 1962b: 344
Euplectus sarawakensis Besuchet, 1956c: 88
Euplectus sexstriatus Besuchet, 1970a: 121
Euplectus sparsus Besuchet, 1964a: 412
Euplectus validus Besuchet, 1958e: 906
Faronus andalusiacus Besuchet, 1969c: 114
Faronus depressus Besuchet, 1960a: 15
Faronus distinctus Besuchet, 1999b: 792
Faronus festivus Besuchet, 1960a: 12
Faronus festivus apterus Besuchet, 1960a: 13 (= F festivus Besuchet)
Faronus gracilis Besuchet, 1969c: 115
Faronus insignis Besuchet, 1958e: 899
Faronus lusitanicus Besuchet, 1969c: 111
Faronus parallelus Besuchet, 1958e: 897
Faronus parnassius Besuchet, 1969e: 397
Faronus rifensis Besuchet, 1963c: 227
Faronus sahlbergi Besuchet, 1960a: 11 (= F. parallelus Besuchet)
Faronus testaceus Besuchet, 1962b: 336
Faronus tingitanus Besuchet, 1962b: 334
Faronus variabilis Besuchet, 1969c: 110
Faronus venustus Besuchet, 1958d: 333
Fustiger appendiculatus Besuchet, 1977b: 261
Fustiger wittmeri Besuchet, 1977b: 264
Geopselaphus affinis Besuchet, 1961f: 259
Geopselaphus alticola Besuchet, 1961f: 260
Geopselaphus balearicus Besuchet, 1969b: 104
Geopselaphus depressus Besuchet, 1961f: 261
Geopselaphus distinguendus Besuchet, 1961f: 254
Geopselaphus espanoli Besuchet, 1969b: 99
Geopselaphus formosus Besuchet, 1961f: 261
Geopselaphus franzi Besuchet, 1961f: 260
Geopselaphus frater Besuchet, 1969b: 100

500

223.
226.
227.
228.
227.
230.
231.
242:
235.
234.
235,
236.
237.
238.
2:
240.
241.
242.
243.
244.
245.
246.
247.
248.
249.
250.
291.
292.
755.
254.
295 -
290:
PAT:

258.
239.
260.
261.
262.
2605.
264.
265:
266.
267.
268.

P. HLAVAC

Geopselaphus jucundus Besuchet, 1961f: 256
Geopselaphus lepidus Besuchet, 1969b: 102
Geopselaphus longipalpis Besuchet, 1969b: 102
Geopselaphus longulus Besuchet, 1961f: 250
Geopselaphus mirandus Besuchet, 1961f: 252
Geopselaphus mussardi Besuchet, 1963c: 231
Geopselaphus nitidus Besuchet, 1969b: 100
Geopselaphus tingitanus Besuchet, 1962b: 364
Glyphobythus fallax Besuchet, 1960c: 403
Glyphobythus hervei Besuchet, 1960c: 404
Halorabyxis gourvesi Besuchet, 1975c:138

Imirus outereloi Besuchet, 1980a: 56

Leptoplectus perraulti Besuchet, 1993a: 340

Linderia picanyolae Besuchet, 1985a: S15

Meliceria (Cyrtoplectus) italica Besuchet, 1966a: 56
Mesoleptochir rougemonti Besuchet, 1974a: 887
Namunia cavernicola Besuchet, 1978b: 131

Namunia lapidicola Besuchet, 1978b: 131

Namunia terricola Besuchet, 1999b: 799
Neopselaphus adisi Besuchet, 1982b: 801
Neopselaphus armatus Besuchet, 1982b: 801
Neopselaphus curtipalpis Besuchet, 1987a: 237
Neopselaphus degalieri Besuchet, 1982b: 806
Neopselaphus filipalpis Besuchet, 1982b: 806
Neopselaphus chalumeaui Besuchet, 1987a: 238
Neopselaphus parki Besuchet, 1982b: 805
Neopselaphus tavakiliani Besuchet, 1982b: 804
Nonveillera lepida Paviéevit & Besuchet, 2003a: 280
Nonveillera romani Paviéevié & Besuchet, 2003a: 282
Octomicrus dentifrons Besuchet, 1999b: 790

Pachacuti huggerti Besuchet, 1987a: 235

Panaphantus afer Besuchet, 1980c: 154

Paramaurops exaratus neapolitanus Besuchet, 1958e: 908
(= P. exaratus Baudi di Selve)

Paratychus minutissimus Besuchet, 1960a: 26
Plectophloeus binaghii Besuchet, 1964a: 417
Plectophloeus erichsoni occidentalis Besuchet, 1969e: 399
Plectophloeus erichsoni orientalis Besuchet, 1969e: 400
Plectophloeus nubigena bosnicus Besuchet, 1964a: 419
Prionobythus genesti Besuchet, 1985a: 509
Pselaphaulax carniolicus Besuchet & Sabella, 2000a: 263
Pselaphaulax siculus Besuchet & Sabella, 1993c: 92
Pselaphogenius laticeps Besuchet, 1961b: 262
Pselaphogenius latinus Besuchet, 1980b: 630
Pselaphogenius lepontinus Besuchet, 1980b: 629

269.
270.
271.
272.
+14.
274.
275.
276.
217.
278.
279.
280.
281.
282.
283.
284.
285.
286.
267.
288.
289.
290.
291,
292.
225.
294.
295.
290.
297:
298.
299.
300.
301.
302.
303.
304.
305.
306.
307.
308.
309.
310.
SIT.
M2,
MS.

CLAUDE BESUCHET’S WORK 501

Pselaphogenius lucanicus Besuchet, 1964a: 435
Pselaphogenius neapolitanus Besuchet, 1964a: 433
Pselaphogenius orientalis Besuchet, 1961a: 39
Pselaphostomus bergamascus Besuchet, 1980b: 628
Pselaphostomus bussacensis estrellensis Besuchet, 1961b: 252
Pselaphostomus franzi Besuchet, 1961b: 246
Pselaphostomus intermedius Besuchet, 1961b: 249
Pselaphostomus lusitanicus Besuchet, 1961b: 244
Pselaphostomus pyrenaeus Besuchet, 1961b: 236
Pselaphostomus stussineri vesulinus Besuchet, 1961a: 33
Pselaphus mysius Besuchet, 1960b: 399

Pselaphus xaymacus Besuchet, 1987a: 236

Pygoxyon bergamascum Besuchet, 1958e: 900 (= P. lombardum Binaghi)
Pygoxyon myops Besuchet, 1958e: 901

Scotoplectus caspicus Besuchet, 1975b: 401
Scotoplectus ponticus Besuchet, 1975b: 400
Scotoplectus weiratheri Besuchet, 1975b: 399
Seracamaurops fritschi Besuchet, 1986c: 4
Syntectodes maldivicus Besuchet, 2008a: 73

Tapas armifer Besuchet, 2008a: 77

Tapas basseti Besuchet, 2008a: 76

Tasmiger strumosus Besuchet, 2008a: 78

Thelotia cebennica Besuchet, 1999b: 795

Tremissus beaumonti Besuchet, 1982d: 320
Tremissus inexspectatus Besuchet, 1982d: 319
Triartiger nomurai Besuchet, 2008a: 74

Tribatus hauseri Besuchet., 1961a: 31

Tribatus lopatini Besuchet, 1964a: 426

Trimium atticum Besuchet, 1969e: 400

Trimium illyricum Besuchet, 1969e: 402

Trimium sardoum Besuchet, 1958c: 7 (= T. amplipenne Reïtter)
Trissemus bellax Besuchet, 1999b: 801

Trissemus holzschuhi Besuchet. 1999b: 805
Trissemus maroccanus wittmeri Besuchet, 1981b: 245
Trissemus micropterus Besuchet, 1970b: 314
Trissemus mundulus Besuchet, 1961e: 2

Trissemus sulcifrons Besuchet, 1999b: 803

Trissemus trilobatus Besuchet, 1999b: 803

Trogaster binaghii Besuchet, 1969d: 214

Trogaster caprai Besuchet, 1969d: 216

Trogaster doderoi Besuchet, 1969d: 213

Trogaster gestroi Besuchet, 1969d: 215

Trogaster solarii Besuchet, 1969d: 216
Trogasteropsis coiffaiti Besuchet, 1977a: 291
Tychobythinus atlanticus Besuchet, 1963b: 218

502

314.
HIS.
316.
FI7.
318.
FT
320.
321.
22
729.
324.
325.
320.
327.
326:
329.
330.
394.
332.
F9
334.
335.
390:
397.
338.
339:
340.
341.
342.
343.
344.
345.
346.
347.
348.
349.
350.
ss.
392.
355$.
354.
335.
356.
SZ.
356.

P. HLAVAC

Tychobythinus brachati Besuchet, 2008b: 246
Tychobythinus curtii Besuchet, 1980b: 615
Tychobythinus escalerai Besuchet, 1962b: 347
Tychobythinus escolai Besuchet, 1974b: 49
Tychobythinus espanoli Besuchet, 1974b: 53
Tychobythinus listai Besuchet, 1985a: 514
Tychobythinus muntani Besuchet, 1974b: 54
Tychobythinus naxius Besuchet, 1993b: 225
Tychobythinus occidentalis Besuchet, 1962b: 349
Tychobythinus rosai Besuchet, 1980b: 616
Tychobythinus strinatii Besuchet, 1982a:50
Tychobythinus urgellesi Besuchet, 1974b: 52
Tychobythinus vignai Besuchet, 1978c: 69
Tychomorphus franzi Besuchet, 1963b: 220
Tychomorphus mussardi Besuchet, 1999a: 56
Tychus affinis Besuchet, 1958e: 911

Tychus altivagus Besuchet, 2011: 30

Tychus anatolicus Besuchet, 1964a: 429

Tychus antalyanus Besuchet & Sabella, 1999e: 250
Tychus asuniensis Besuchet, 1964a: 433

Tychus atlanticus Besuchet & Sabella, 1999e: 239
Tychus brachati Besuchet & Sabella, 1999d: 314
Tychus caspicus Besuchet & Sabella, 1999e: 254
Tychus coifjaiti Besuchet, 1958e: 913

Tychus cordiger Besuchet, 1969e: 408

Tychus distinguendus Besuchet, 1960a: 23

Tychus epiroticus Besuchet, 1964a: 431

Tychus georgicus Besuchet & Sabella, 1999e: 247
Tychus holzschuhi Besuchet & Sabella, 19994: 317
Tychus judaeus Besuchet, 1964a: 428

Tychus laminiger Besuchet, 1969e: 409

Tychus latebrosus Besuchet, 2011: 26

Tychus longicornis Besuchet, 1958e: 914 (= T. balcanicus Reitter)
Tychus lusitanicus Besuchet & Sabella, 1999e: 237
Tychus manicanus Besuchet & Sabella, 1999e: 238
Tychus mundulus Besuchet, 1958e: 915

Tychus paludivagus sicilianus Besuchet & Sabella, 1996: 111
Tychus pelopeius Besuchet & Sabella, 1999d: 312
Tychus persicus Besuchet & Sabella, 1999e: 245
Tychus ponticus Besuchet & Sabella, 1999e: 249
Tychus remaudierei Besuchet, 1969e: 407

Tychus rhodopeus Besuchet & Sabella, 1999d: 313
Tychus sardous Besuchet, 1964a: 432

Tychus sengleti Besuchet & Sabella, 1999d: 315
Tychus striola andalusiacus Besuchet & Sabella, 1996: 107

SLA
360.
361.
302.

CLAUDE BESUCHET’'S WORK

Tychus striola balearicus Besuchet & Sabella, 1996: 105
Tychus viti Besuchet, 2011: 23

Tychus tingitanus Besuchet, 1962b: 364

Tyrodes janetscheki Besuchet, 1970b: 316

IV. List of species and subspecies of Scydmaenidae

30.
12
2.

Cephennium (Cephennium) fraterculum Besuchet, 1971b: 278
Cephennium (Cephennium) machulkai Besuchet, 1971b: 278
Cephennium (Cephennium) paganettii Besuchet, 1971b: 278
Cephennium (Phennecium) galitense Besuchet, 1982e: 238
Cephennium (Phennecium) solarii Besuchet, 1958e: 896
new name for C. romanum Holdhaus, 1924: 21
Cephennodes (s.str.) basilewskyi Besuchet, 1962c: 420
Cephennodes (s.str.) indifferens Besuchet, 1962c: 422
Cephennodes (s.str.) leleupi Besuchet, 1962c: 421
Cephennodes (s.str.) marginatus Besuchet, 1962c: 423
Cephennomicrus fossulatus Besuchet, 1961: 17
Cephennomicrus glaber Besuchet, 1961: 24
Cephennomicrus impressus Besuchet, 1961h: 15
Cephennomicrus jucundus Besuchet, 1961h: 23
Cephennomicrus latipennis Besuchet, 1961h: 21
Cephennomicrus longicornis Besuchet, 1961h: 18
Cephennomicrus pauliani Besuchet, 1961h: 19
Cephennomicrus pusillus Besuchet, 1961h: 24
Cephennomicrus rugosicollis Besuchet, 1961h: 19
Cephennomicrus suturalis Besuchet, 1961h: 22
Cephennomicrus vadoni Besuchet, 1961h: 16
Cephennomicrus validus Besuchet, 1961h: 21

Clidicus loebli Besuchet, 1971a: 254

Clidicus mussardi Besuchet, 1971a: 255

Clidicus quadricollis Besuchet, 1971a: 252

Etelea tingitana Besuchet & Vit, 2004b: 341

Leptocharis algericus Besuchet, 1958e: 916
Nanophthalmus nonveilleri Besuchet & Vit, 2000b: 159
Nanophthalmus serbicus Besuchet & Vit, 2000b: 159
Neuraphes (Pararaphes) toumayeffi Besuchet, 1980d: 192
Scydmaenus aelleni Besuchet, 1981c: 460

Taurablepton asitawandas Besuchet, 19692: 315 (Ablepton)
Taurablepton rutash Besuchet, 1969g: 316 (Ablepton)

V. List of species and subspecies of Ptiliidae

BR © D =

Actidium reticulatum Besuchet, 1971c: 319
Oligella intermedia Besuchet, 1971c: 320
Ptiliolum stockmanni Besuchet, 1971c: 326
Ptilium (Ptilium) cognatum Besuchet, 1971c: 323

503

504 P. HLAVAC

5. Ptilium (Ptilium) scrutandum Besuchet, 1971c: 322
6. Ptilium (Ptilium) timidum Besuchet, 1971c: 322

VI. List of species and subspecies of Leptinidae

1. Leptinus illyricus Besuchet, 1980e: 136
2. Leptinus pyrenaeus Besuchet, 1980e: 139

VII. List of species and subspecies of Cerylonidae:

Axiocerylon baloghi Besuchet & Slipinski, 1988b: 908
Axiocerylon bournei Besuchet & Slipinski, 1988b: 908
Axiocerylon burckhardti Besuchet & Slipinski, 1988b: 908
Axiocerylon decemcostatum Besuchet & Slipinski, 1988b: 922
Axiocerylon ghanense Besuchet & Slipinski, 1988b: 919
Axiocerylon gomyi Besuchet & Slipinski, 1988b: 920
Axiocerylon hammondi Besuchet & Slipinski, 1988b: 916
Axiocerylon humerale Besuchet & Slipinski, 1988b: 914
Axiocerylon loebli Besuchet & Slipinski, 1988b: 923
Axiocerylon luzonicum Besuchet & Slipinski, 1988b: 908
Axiocerylon minimum Besuchet & Slipinski, 1988b: 911
Axiocerylon myops Besuchet & Slipinski, 1988b: 910
Axiocerylon orousseti Besuchet & Slipinski, 1988b: 910
Axiocerylon perkorum Besuchet & Slipinski, 1988b: 910
Axiocerylon roberti Besuchet & Slipinski, 1988b: 916
Axiocerylon solomonense Besuchet & Slipinski, 1988b: 913
Axiocerylon triste Besuchet & Slipinski, 1988b: 911
Axiocerylon variabile Besuchet & Slipinski, 1988b: 916
Axiocerylon venustum Besuchet & Slipinski, 1988b: 919

ni dt ni dt et it dt dt nt pt
CHU ES DES Lou U ED Em

20: Cautomus (s.str.) distinguendus Besuchet, 1972: 119

21. Cautomus (s.str.) elongatus Besuchet, 1972: 118

22. Cautomus (s.str.) latus Besuchet, 1972: 125

23. Cautomus (s.str.) venustus Besuchet, 1972: 126

24. Cautomus (Aculagnathus) pusillus Besuchet, 1972: 138
25. Cautomus (Leptoxycheilus) convexus Besuchet, 1972: 133
26. Cautomus (Leptoxycheilus) longipilis Besuchet, 1972: 134
27. Cautomus (Leptoxycheilus) myops Besuchet, 1972: 130
28. Cautomus (Leptoxycheilus) philippinensis Besuchet, 1972: 128
29. Cautomus (Leptoxycheilus) punctatus Besuchet, 1972: 132
30. Cautomus (Leptoxycheilus) sugerens Besuchet, 1972: 130
Aile. Cautomus (Paracautomus) nitidus Besuchet, 1972: 136
32. Cautomus (Paracautomus) reticulatus Besuchet, 1972: 133
33. Glyptolopus amazonicus Besuchet & Slipiñski, 1987b: 80
34. Glyptolopus convexus Besuchet & Slipiñski, 1987b: 78

35. Glyptolopus peruanus Besuchet & Slipiñski, 1987b: 81

CLAUDE BESUCHET’S WORK 505

VIII. List of species and subspecies of Rhipiporidae:
1. Pirhidius beaumonti Besuchet, 1957: 348

IX. List of species and subspecies of Dryopidae:
Le Geoparnus setifer Besuchet, 1978e: 706

ACKNOWLEDGEMENT

I would like to thank to Jon Cooter for reading and commenting on the manu-
script.

NOTE BY JON COOTER

It has been an honour to read through a draft of this contribution by my good
friend Peter Hlaväë and have the opportunity to add a personal brief note. Although,
like many coleopterists, I had corresponded with Claude Besuchet for a number of
years, it was in 1997 when we actually met. Claude invited me to work in Geneva
Museum for a period of two weeks identifying and checking what Claude regarded as
all the Leiodinae: Leiodini from Swiss institutional and private collections and sundry
extra Swiss specimens from non-Swiss museums. This was part of the revision of the
Swiss list, which as the millennium approached, would have been 100 years old.

As my departure date drew near I received a message from Claude that he
would meet me at Geneva airport and words to the effect that I would have no trouble
in identifying him — indeed this was true, after entry formalities I entered the public
area to see a genial giant of a man, who initially reminded me a little of Jaques Tati,
holding a volume of Freude Harde & Lohse "Die kafer mitteleuropas" rather than a
board with "J.Cooter" written on 1t. I greatly enjoyed the hospitality Claude extended
to me during my stay, which included practicalities of eating out in Geneva, visits to
nature reserves 1n the Geneva area and the Jura as well as the Geneva Insekten Bô6rse
and a very pleasant relaxing afternoon and meal at Claude’s home. Needless to say,
entomologically my every need was catered for — my own office space, Power Mac,
microscope and accommodation in the museum with 24hr access to the collections.

As Peter has stated, Claude regarded us lesser mortals as equals, imparting his
knowledge freely and always ready to help and support colleagues. Congratulations
Claude on your 80th birthday and for your scientific legacy.

PUBLICATIONS OF CLAUDE BESUCHET

I. Contributions on systematics

1. BESUCHET, C. 1952. Larves et nymphes de Plectophloeus (Col. Pselaphidae). Mitteilungen
der Schweizerischen entomologischen Gesellschaft 25 (3): 251-256.

2. BESUCHET, C, 1953. Bibloplectus nouveaux d'Europe centrale (Col. Pselaphidae). Mit -
teilungen der Schweizerischen entomologischen Gesellschaft 26 (3): 225-232.

3. BESUCHET, C. 1954a. Arcopagus (Bythobletus) chevrolati Aubé et carinula Rey (Col.
Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 27 (2):
157-160.

4. BESUCHET, C, 1954b. Arcopagus mimus Dod. Description de la femelle et d'une variété
nouvelle (Col. Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesell-
schaft 27 (4): 435-436.

506 P. HLAVAC

2%.

26:

BESUCHET, C. 1954c. Une nouvelle espèce de Brachygluta du Danemark (Col. Pselaphidae).
Entomologiske Meddelelser 27: 43-45.

BESUCHET, C. 1955a. Monographie des Bibloplectus et Pseudoplectus paléarctiques (Col.
Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 28 (2):
153-209.

BESUCHET, C. 1955b. Arcopagus picteti Tourn. et krügeri Mach. (Col. Pselaphidae).
Mitteilungen der Schweizerischen entomologischen Gesellschaft 28 (3): 274-278.

BESUCHET, C. 1955c. Bibloporus européens décrits par Jeannel (Col. Pselaphidae). Mit-
teilungen der Schweizerischen entomologischen Gesellschaft 28 (3): 278.

BESUCHET, C. 1956a. Révision des genres Zibus, Saulcyella, Aphiliops et description d'un

genre nouveau (Col. Pselaphidae). Mitteilungen der Schweizerischen entomologischen
Gesellschaft 29 (4): 363-372.

. BESUCHET, C. 1956b. Larves et nymphes de Psélaphides (Coléoptères). Revue suisse de

Zoologie 63: 697-705.

. BESUCHET, C. 1956c. Un Euplectus nouveau de Bornéo (Col. Pselaphidae). The Ento -

mologist 89: 88-90.

. BESUCHET, C. 1956d. Biologie, morphologie et systématique des Rhipidius (Col. Rhipi-

phoridae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 29(2):
73-144. Thèse.

. BESUCHET, C. 1956e. Rhipidius abeillei Chob. (Col. Rhipiphoridae). Mitteilungen der

Schweizerischen entomologischen Gesellschaft 29(3): 254.

. BESUCHET, C. 1957. Contribution à l’étude des Rhipidiinae. Bulletin de la Société Vaudoise

des Sciences naturelles 66(293): 341-351.

. BESUCHET, C. 1958a. Bryaxis Kugelann et Bryaxis Leach. Mitteilungen der Schweizerischen

entomologischen Gesellschaft 31 (1): 65-69.

. BESUCHET, C. 1958b. Bryaxis grouvellei Rtt., sculpticornis Guillb. et description d'une

espèce nouvelle (Col. Pselaphidae). Mitteilungen der Schweizerischen entomologischen
Gesellschaft 31 (1): 70-76.

. BESUCHET, C. 1958c. Description de deux Psélaphides nouveaux d'Italie (Coléoptères).

Bollettino del Museo Civico di Storia Naturale di Venezia 11: 7-10.

. BESUCHET, C. 1958d. Descriptions de quelques Psélaphides paléarctiques nouveaux

(Coleoptera). Mitteilungen der Schweizerischen entomologischen Gesellschaft 31 (3-4):
333-338.

. BESUCHET, C. 1958e. Coleoptera Pselaphidae et Scydmaenidae. Revue suisse de Zoologie 65

(4): 891-919.

. BESUCHET, C. 1959a. Quelques remarques concernant le genre Amauronyx Reïtt. et

description d'une espèce nouvelle (Coleoptera Pselaphidae). Archivos del Instituto de
Aclimataciôn 8: 25-27.

. BESUCHET, C. 1959b. Coléoptères Psélaphides et Scydménides de la collection CI. Rey.

Mitteilungen der Schweizerischen entomologischen Gesellschaft 32 (2-3): 328-332.

. BESUCHET, C. 1960a. Coléoptères Psélaphides de la collection J. Sahlberg. Suomen

Hyôünteistieteellinen Aikakauskirja; Annales Entomologici Fennici 26 (1): 11-31.

. BESUCHET, C. 1960b (1959). Description de deux Psélaphides paléarctiques nouveaux

(Coleoptera). Mitteilungen der Schweizerischen entomologischen Gesellschaft 32 (4):
398-400.

. BESUCHET, C. 1960c (1959). Petite revision du genre Glyphobythus Rafïffr. (Col.

Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 32 (4):
401-408.

BESUCHET, C. 1961a. Psélaphides paléarctiques. Espèces nouvelles et notes synonymiques
(Coleoptera). Mitteilungen der Schweizerischen entomologischen Gesellschaft 34 (1):
30-42.

BESUCHET, C. 1961b. Révision des Pselaphostomus et Pselaphogenius ibériques (Col.
Pselaphidae). Eos, Madrid 37: 229-265.

21.

28.

48.

49.

CLAUDE BESUCHET’'S WORK 507

BESUCHET, C. 1961c. Révision des Claviger ibériques (Coleopt. Pselaphidae). Revue suisse
de Zoologie 68: 443-460.

BESUCHET, C. 1961d. Révision du genre Decatocerus SAULCY (Col. Pselaphidae).
Miscellanea Entomologica 1 (4): 91-97.

. BESUCHET, C. 1961e. Deux Psélaphides nouveaux d'Iran (Coleoptera). Stuttgarter Beiträge

zur Naturkunde, 51 (3): 1-3.

. BESUCHET, C. 1961f (1960). Revision du genre Geopselaphus Jeann. (Col. Pselaphidae).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 33 (4): 245-263.

. BESUCHET, C. 1961g. Quelques Bryaxis nouveaux du Caucase (Col. Pselaphidae). Zoolo -

gicheskiy Zhurnal 40 (12): 1827-1831, in Russian.

. BESUCHET, C. 1961h. Recherches sur la faune endogée de Madagascar VI. Description de

quelques Cephennomicrus nouveaux de Madagascar et des Comores. Mémoires de
l'Institut Scientifique de Madagascar (E: Entomologie), sér. E 12: 15-25.

. BESUCHET, C. 1962a. Psélaphides récoltés par M. J. Klapperich en Afghanistan (Coleo-

ptera). Annales Historico-Naturales Musei Nationalis Hungarici (Zool.) 54: 269-270.

. BESUCHET, C. 1962b. Contribution à l'étude des Psélaphides du Maroc (Coleoptera).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 34 (4, 1961): 333-371.

. BESUCHET, C. 1962c. Coleoptera Scydmaenidae: Cephenniini. Mission zoologique de

l'I.R.S.A.C. en Afrique orientale (P. Basilewsky et N. Leleup, 1957). LXII. Annales du
Musée Royal de l'Afrique Centrale, Tervuren (Série 8°: Sciences Zoologiques) 107:
420-423.

. BESUCHET, C. 1963a. Notes sur quelques Brachygluta paléarctiques (Col. Pselaphidae).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 36 (1-2): 27-46.

. BESUCHET, C. 1963b. Troisième contribution à l'étude des Psélaphides du Maroc

(Coleoptera). Archives des Sciences, Genève 16 (2): 217-223.

. BESUCHET, C. 1963c (1962). Psélaphides récoltés au Maroc par M. R. Mussard (Coleoptera).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 35 (3-4): 227-232.

. BESUCHET, C. 1964a. Psélaphides paléarctiques. Espèces nouvelles et notes synonymiques.

IT (Coleoptera). Revue suisse de Zoologie 71 (2): 411-443.

. BESUCHET, C. 1966a. Meliceria RAFFR. et Cyrtoplectus NORM. (Col. Pselaphidae).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 39 (1-2): 49-58.

. BESUCHET, C. 1966b. Revision des Centrophthalmus paléarctiques. Mitteilungen der

Schweïizerischen entomologischen Gesellschaft 39 (1-2): 59-65.

. BESUCHET, C. 1966c. Bryaxis KUGELANN, 1794 and Bythinus Leach, 1817 (Insecta,

Coleoptera): proposed addition to the official list in their original sense. Z. N.(S.) 1642.
Bulletin of Zoological Nomenclature 23 (2-3): 114-116.

. BESUCHET, C. 1968. Psélaphides des Canaries et de Madère (Coleoptera). Mitteilungen der

Schweizerischen entomologischen Gesellschaft 41 (1-4): 275-297.

. BESUCHET, C. 1969a. Rybaxis SAULCY, 1876 (Insecta, Coleoptera): Proposed designation

of a type-species under the plenary powers. Z.N. (S.) 1882. Bulletin of Zoological
Nomenclature 26 (3): 166.

. BESUCHET, C. 1969b. Geopselaphus d'Espagne nouveaux ou peu connus (Col. Pselaphidae).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 42 (1-2): 96-105.

. BESUCHET, C. 1969c. Faronus nouveaux ou méconnus de la péninsule ibérique (Col. Pse -

laphidae). Mitteilungen der Schweïizerischen entomologischen Gesellschaft 42 (1-2):
106-116.

. BESUCHET, C. 1969d. Les Trogaster du groupe de doriae (Coleoptera Pselaphidae).

Memorie della Società Entomologica Italian 48: 211-218.

BESUCHET, C. 1969e. Psélaphides paléarctiques. Espèces nouvelles et notes synonymiques.
IT (Coleoptera). Revue suisse de Zoologie 76 (2): 397-420.

BESUCHET, C. 1969f. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der
Mongolei. 180. Pselaphidae. Reichenbachia 11: 301-304.

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. BESUCHET, C. 1969g. Description de deux Ablepton (Col., Scydmaenidae). Mitteilungen der

Schweizerischen entomologischen Gesellschaft 42(4): 313-316.

. BESUCHET, C. 1969h. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der

Mongolei. 184. Puliidae. Faunistische Abhandlungen Staatliches Museum fur Tierkunde
in Dresden 3(3): 13-14.

. BESUCHET, C. 1970a. Nouveaux Psélaphides des Canaries et de Madère (Coleoptera).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 43 (2): 119-124.

. BESUCHET, C. 1970b. Psélaphides récoltés au Népal par le Prof. H. Janetschek (Coleoptera).

Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 58 (1970):
313-318.

. BESUCHET, C. 197la. Les Clidicus de Ceylan (Col. Scydmaenidae). Mitteilungen der

Schweïzerischen entomologischen Gesellschaft 43(3): 249-257.

. BESUCHET, C. 1971b. 18. Familie: Scydmaenidae [1. Tribus Euthüini, 2. Tribus Cephenniini]

(pp. 273-278). 1n: Freude H., Harde K. W. & Lohse G. A. (eds). Die Käfer Mitteleuropas.
Band 3. Adephaga 2. Palpicornia, Histeroidea, Staphylinoidea 1. Goecke & Evers,
Krefeld, 365 pp.

BESUCHET, C. & SUNDT, E. 1971c. 1n: Freude H., Harde K. W. & Lohse G. A. (eds). Die
Käfer Mitteleuropas. Band 3. Adephaga 2. Palpicornia, Histeroidea, Staphylinoidea 1.
Goecke & Evers, Krefeld, 365 pp.

BESUCHET, C. 1972. Les Coléoptères Aculagnathides. Revue Suisse de Zoologie 79(1):

99-145.

BESUCHET, C. 1974a. Un Clavigerinae nouveau de l'Ethiopie (Col. Pselaphidae). Revue

suisse de Zoologie 81 (4): 887-891.

BESUCHET, C. 1974b. Les Psélaphides cavernicoles de l'Espagne (Coleoptera Pselaphidae).
Miscelanea zoologica 3 (4): 41-69.

BESUCHET, C. 1974c. 24. Familie: Pselaphidae (pp. 305-362). /n: H. Freude, K. W. Harde,
and G. A. Lohse (eds). Die Käfer Mitteleuropas. Vol. 5, Staphylinidae IT (Hypocyphtinae
und Aleocharinae), Pselaphidae. Goecke & Evers, Krefeld, 381 pp.

BESUCHET, C. 1975a. Bibloplectus nouveaux de la région méditerranéenne orientale (Col.
Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 48 (1-2):
31-36.

BESUCHET, C. 1975b. Revision du genre Scotoplectus Reitt. (Col. Pselaphidae). Mitteilungen

der Schweizerischen entomologischen Gesellschaft 48 (3-4): 397-404.

BESUCHET, C. 1975c. Un Halorabyxis nouveau de la Polynésie française (Coleoptera

Pselaphidae). Compte Rendu des Séances, SPHN Genève (N.S.) 10 (2-3): 137-140.

BESUCHET, C. 1976. Contribution à l'étude des Ptiliides paléarctiques (Coleoptera).

Mitteilungen der Schweizerischen entomologischen Gesellschaft 49: 51-71.

BESUCHET, C. 1977a. Les Trogasteropsis de la Catalogne (Col. Pselaphidae). Nouvelle

Revue d’'Entomologie 7 (3): 291-293.

BESUCHET, C. 1977b. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen
Museums in Basel. Coleoptera: Fam. Pselaphidae Subf. Clavigerinae Gen. Fustiger
BREND. mit Einschluss der indischen Arten. Entomologica Basiliensia 2: 261-267.

BESUCHET, C. 1978a. Un Bythinus cavernicole nouveau (B. hauseri) de la Grèce (Coleoptera
Pselaphidae). Annales Musei Goulandris 4: 263-265.

BESUCHET, C. 1978b. Le genre Namunia Reitt. (Coleoptera, Pselaphidae). Revue Suisse de
Zoologie 85 (1): 127-133.

BESUCHET, C. 1978c. Le premier Psélaphide troglobie de la Turquie (Coleoptera). Quaderni
di Speleologia, Circolo Speleologia Romano 3 (1978-79): 69-73.

BESUCHET, C. 1978d. Réintroduction en Suisse du Platypsyllus castoris Rits. (Coleoptera:

Leptinidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 51:
291-292.

. BESUCHET, C. 1978e. Description d’un Dryopide terrestre nouveau de la Malaisie (Coleo -

ptera). Revue suissse de Zoologie 85(4): 705-709.

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CLAUDE BESUCHET’S WORK 509

BESUCHET, C. 1979. Description d'un nouveau Batrisus AUBÉ (Coleoptera, Pselaphidae).
Revue suissse de Zoologie 86 (1): 279-283.

BESUCHET, C. 1980a. Le genre /mirus REITT. (Coleoptera, Pselaphidae). Nouvelle Revue
d'Entomologie 10 (1): 51-58.

BESUCHET, C. 1980b. Contribution à l'étude des Coléoptè res Psélaphides d'Italie et du
Tessin. Revue suissse de Zoologie 87 (2): 611-635.

BESUCHET, C. 1980c. Le genre Panaphantus KIESW. (Col., Pselaphidae). Mitteilungen der
Schweizerischen entomologischen Gesellschaft 53 (2-3): 151-155.

BESUCHET, C. 1980d. Neuraphes et Scydmoraphes de la Suisse, de l’ Ain et de la Haute-
Savoie. Mitteilungen aus der Entomologischen Gesellschaft 30: 189-196.

BESUCHET, C. 1980e. Revision des Leptinus paléarctiques (Coleoptera: Leptinidae). Revue
Suisse de Zoologie 87(1): 131-142.

BESUCHET, C. 1981a. Contribution à l'étude des Batrisodes paléarctiques (Coleoptera:
Pselaphidae). Revue Suisse de Zoologie 88 (1): 275-296.

BESUCHET, C. 1981b. Insects of Saudi Arabia. Coleoptera: Fam. Pselaphidae. Fauna of
Saudi Arabia 3: 243-250.

BESUCHET, C. 1981c. Description d’un Coléoptè re Scydménide cavernicole de la Nouvelle-
Calédonie. Revue suisse de Zoologie 88(2): 459-461.

BESUCHET, C. 1982a. Contribution à l'étude des Bythinini cavernicoles néarctiques (Cole-
optera: Pselaphidae). Revue suisse de Zoologie 89 (1): 49-53.

BESUCHET, C. 1982b. Le genre Neopselaphus JEANN. (Coleoptera: Pselaphidae). Revue
suisse de Zoologie 89 (3): 797-807.

BESUCHET, C. 1982c. Coléoptères rares des laisses de la Dranse (Haute-Savoie). Bulletin
Romand d’Entomologie 1: 97-100.
BESUCHET, C. 1982d (1981). Un nouveau genre de Brachyglutini de la région méditer-

ranéenne orientale (Col. Pselaphidae). Mitteilungen der Schweïzerischen entomolo-
gischen Gesellschaft 54 (4, 1981): 317-320.

BESUCHET, C. 1982e. 238-240: In. Franz, H. Beitrag zur Kenntnis der Coleopterenfauna der
Insel Galita. Sitzungsberichte der Osterreichischen Akademie der Wissenschaften,
Mathematisch-Naturwissenschaftliche Klasse (Abt. I), 191 (5-10): 231-240.

BESUCHET, C. 1983a. Un nouveau genre de Pyxidicerini du sud de l'Inde (Coleoptera:
Pselaphidae). Revue suisse de Zoologie 90 (2): 509-512.

BESUCHET, C. 1983b. Bryaxis nouveaux ou méconnus du Nord de l'Italie (Coleoptera:
Pselaphidae). Revue suisse de Zoologie 90 (3): 769-780.

BESUCHET, C. 1983c. The species of Pselaphidae (Coleoptera) of the Hortobägy National
Park. In: The Fauna of the Hortobägy National Park. Vol. 2. (Ed: Mahunka, S.)
Akadémiai Kiad6, Budapest 197-198.

BESUCHET, C. 1985a. Bythinini cavernicoles nouveaux de France et d'Espagne (Coleoptera:
Pselaphidae). Revue suisse de Zoologie 92 (2): 509-517.

BESUCHETT, C. 1985b. Le genre Acetalius SHARP (Coleoptera: Pselaphidae). Revue suisse
de Zoologie 92 (3): 761-766.

BESUCHET, C. 1985c. 27. Coléoptères Psélaphides. 199: In Expédition Thai-Maros 85 —
Rapport Spéléologique et Scientifique. Edit. Association Pyrénéenne de Spéléologie,
Toulouse: 199.

BESUCHET, C. 1986a. Synonymes et homonyme nouveaux de quelques genres de Pséla-
phides (Coleoptera). Revue suisse de Zoologie 93 (1): 257-264.

BESUCHET, C. 1986b. Coléoptères Psélaphides des îles du Cap Vert (Coleoptera: Psela-
phidae). CFS — Courier, Forschurigsinstitut Senckenberg 81: 41-42.

BESUCHET, C. 1986c. Contribution à l'étude du genre Seracamaurops WINKLER, 1925
(Coleoptera, Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesell-
schaft 59: 459-463.

BESUCHET, C. 1987a. Pselaphini néotropicaux nouveaux (Coleoptera: Pselaphidae).
Archives des Sciences (Genève) 40 (2): 231-240.

510 P. HLAVAC

96. BESUCHET, C. & SLIPINSKI, S. A. 1987b. A review of Glyptolopus Erichson (Coleoptera,
Cerylonidae) with descriptions of new species. Mitteilungen der Schweizerischen Ento-
mologischen Gesellschaft 60: 73-81.

97. BESUCHET, C. 1988a. Description de deux Batrisodes paléarctiques nouveaux (Coleoptera:
Pselaphidae). Revue suisse de Zoologie 85 (2): 433-437.

98. BESUCHET, C. & SLIPINSKI, À. S. 1988b. A review of Axiocerylon Grouvelle (Coleoptera,
Cerylonidae) with descriptions of new species. Revue Suisse de Zoologie 95(3): 901-928.

99. BESUCHET, C. 1989. 24. Familie: Pselaphidae (pp. 240-243). /n: LOHSE, G. A. & W. H.
LUCHT (eds). Die Käfer Mitteleuropas. Vol. 12 (1. Supplementband mit Katalogteil).
Goecke & Evers, Krefeld. 346 pp.

100. BESUCHET, C. 1990. Nouvelle contribution à l'étude des Psélaphides des Canaries (Cole-
optera). Vieraea 18: 161-166.

101. BESUCHET, C. 1991. Révolution chez les Clavigerinae. Revue suisse de Zoologie 98 (3):
499-515.

102. BESUCHET, C. 1993a. Les Coléoptères Psélaphides de l'atoll Fangataufa. Bulletin de la
Société entomologique de France 98 (4): 339-341.

103. BESUCHET, C. 1993b. Psélaphides cavernicoles de Grèce (Coleoptera). Biologia gallo-hel -
lenica 20 (1): 223-229.

104. BESUCHET, C. & SABELLA, G. 1993c. Ricerce sugli Pselaphidae di Sicilia VII. Lo Psela -
phaulax dresdensis (Herbst, 1792) e le sue sottospecie. (Coleoptera: Pselaphidae).
Animalia, Catania 20 (1-3): 87-95.

105. BESUCHET, C. & SABELLA, G. 1996. Ricerche sugli Pselaphidae di Sicilia. X. Revisione
delle sottospecie maghrebine di Tychus striola Guillebeau, 1888, con descrizione di
nuovi taxa (Coleoptera: Pselaphidae). Annales de la Société Entomologique de France
(N.S.) 32 (1): 101-118.

106. BESUCHET, C. 1999a. Psélaphides paléarctiques. Notes taxonomiques et faunistiques (Cole-
optera Staphylinidae Pselaphidae). Revue suisse de Zoologie 106(1): 45-67.

107. BESUCHET, C. 1999b. Psélaphides paléarctiques nouveaux ou méconnus.(Coleoptera Sta-
phylinidae Pselaphinae). Revue suisse de Zoologie 106(4): 789-811.

108. BESUCHET, C. 1990c. Le genre Dicentrius (Coleoptera Staphylinidae Pselaphinae). Mit-
teilungen der Schweizerischen entomologischen Gesellschaft 72: 221-233.

109. BESUCHET, C. & SABELLA, G. 1999d. Descrizione di nuove specie di Tychus della regione
palearctica noc revisione dei Tychus del gruppo armeniacus (Coleoptera: Staphylinidae:
Pselaphinae). Annales de la Société Entomologique de France (N.S.) 35 (3-4): 303-318.

110. BESUCHET, C. & SABELLA, G. 1999. Nouve specie di Tychus della regione palearctica
(Coleoptera: Staphylinidae: Pselaphinae). Mitteilungen der Schweizerischen entomolo-
gischen Gesellschaft 72: 235-258.

111. BESUCHET, C. & SABELLA, G. 2000a. Pselafidi nuovi per la fauna italiana. Naturalista
siciliano IV, XXIV (3-4): 259-269.

112. BESUCHET, C. & VIT, S. 2000b. Les Nanophthalmus Motschulsky d'Europe. Revue Suisse
de Zoologie 107(1): 153-163.

113. BESUCHET, C. 2002 (2001). Bryaxis nouveaux pour la France et l’Italie (Coleoptera, Sta -
phylinidae, Pselaphinae). Mitteilungen der Schweizerischen entomologischen Gesell-
schaft 74: 211-216.

114. PAVICEVIC, D. & BESUCHET, C. 2003a. Bythinini troglobies des Balkans; un genre nouveau
et deux espè ces nouvelles (Coleoptera Staphylinidae Pselaphinae). Mitteilungen der
Schweizerischen Entomologischen Gesellschaft 76: 279-285.

115. NONVEILLER, G., PAVICEVIC, D. & BESUCHET, C. 2003b. Bryaxis tuberculiceps Sp. n.,
Psélaphide cavernicole du Monténégro (Coleoptera Staphylinidae Pselaphinae).
Mitteilungen der Schweizerischen Entomologischen Gesellschaft 76: 287-291.

116. SABELLA, G., BÜCKLE, C., BRACHAT, V. & BESUCHET, C. 2004a. Revision der Palä-
arktischen Arten der Gattung Brachygluta Thomson, 1859 (Coleoptera, Staphylinidae)
(1. Teil). Muséum d'histoire naturelle, Genève pp. 283.

117.

118.

149,

120.

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122.

123;

124.

125:

126

CLAUDE BESUCHET’S WORK 511

BESUCHET, C. & Vir, S. 2004b. Le genre Etelea Csiki, 1909 (Coeoptera: Scydmaenidae:
Cephenniini). Mitteilungen der Schweïizerischen Entomologischen Gesellschaft 77:
337-343.

LÔBL I. & BESUCHET, C. 2004c: Staphylinidae: Pselaphidae, pp. 272-329. /n: I. Lôbl & A.
Smetana (eds). Catalogue of Palearctic Coleoptera. Vol. 2. Stenstrup: Apollo Books,
942 pp.

BESUCHET, C. 2004d: New nomenclatorial and taxonomic acts, and comments. Scyd-
maenidae: Cephennini (pp. 25-26). In: I. Lôbl & A. Smetana (eds). Catalogue of
Palearctic Coleoptera. Vol. 2. Stenstrup: Apollo Books, 942 pp.

BESUCHET, C. 2004e: New nomenclatorial and taxonomic acts, and comments. Staphy-
linidae: Pselaphinae (pp. 28-29). /n: I. Lôbl & A. Smetana (eds). Catalogue of Palearctic
Coleoptera. Vol. 2. Stenstrup: Apollo Books, 942 pp.

BESUCHET, C. 2004f: Scydmaenidae: Cephenniini (pp. 203-206). /n: I. Lôbl & A. Smetana
(eds). Catalogue of Palearctic Coleoptera. Vol. 2. Stenstrup: Apollo Books 942 pp.

BESUCHET, C. & S. KURBATOV. 2007a: Les Bryaxis du Caucase et du secteur oriental des
chaînes Pontiques (Coleoptera: Staphylinidae: Pselaphinae). Russian Entomological
Journal 16(2): 155-206.

BESUCHET, C. & BEKCHIEV, R. 2007b: Description d'un nouveau Batrisodes de la Bulgarie
(Coleoptera: Staphylinidae: Pselaphinae). Historia naturalis bulgarica 18: 75-80.

BESUCHET, C. 2008a. Synonymies et combinaisions nouvelles, revalidations et description
de taxa nouveaux de Pselaphinae (Coleoptera: Staphylinidae). Mitteilungen der
Schweïizerischen Entomologischen Gesellschaft 81: 61-82.

BESUCHET, C. 2008b. Two new cavernicolous Bythinini from Greece (Coleoptera, Staphy-
linidae, Pselapinae) (pp. 161-164). /n: D. Paviéevié & M. Perreau (eds). Advances in the
studies of the subterranean and epigean fauna of the Balkan Peninsula. Volume dedicated
to the memory of Guido Nonveiller. Institute for Nature Conservation of Serbia,
Belgrade, Monographs 22: vin + 564 pp.

. SABELLA, G., BESUCHET, C. & KURBATOV, S. A. 2011. New species of Tychini from Turkey

and Japan (Coleoptera: Staphylinidae: Pselaphinae). Zootaxa 2764: 22-34.

II. Faunistics

L:

10.

BESUCHET, C. & COMELLINI, A. 1964b. Coléoptères endogés des environs de Genève.
Mitteilungen der Schweizerischen entomologischen Gesellschaft 36 ((4) (1963)):
313-320.

BESUCHET, C. & COMELLINI, À. 1965 (1964). Coléoptères endogés des environs de Genève.
Proc. XII Int. Congress of the Entomology, London 459-460.

BESUCHET, C. 1968. Répartition des insectes en Suisse. Influence des glaciations. Mit -
teilungen der Schweizerischen entomologischen Gesellschaft 41 (1-4): 337-340.

BESUCHET, C. 1976. Coléoptères dans les laisses de la Dranse. Arve-Léman Nature 12: 1-6.

BESUCHET, C. 1977. Coléoptères endogés du Tessin. Mitteilungen der Schweizerischen
entomologischen Gesellschaft 49: 305.

BESUCHET, C. 1982. Coléoptères rares des Laisses de la Dranse (Haute-Savoie). Bulletin
Romand d’entomologie 1: 97-100.

BESUCHET, C. 1983. Coléoptères des Alpes suisses atteignant ou dépassant l’altitude de
3000 m. Bulletin Romand d’entomologie 1: 167-176.

BESUCHET, C. 1985. Combien d’espèces de Coléoptères en Suisse. Bulletin Romand d’ento-
mologie 3: 15-25.

BESUCHET, C. & MOESCHLER, P. 1986. Coléoptères cavernicoles dans des sources karstiques
du Jura Neuchatelois (Suisse). Bulletin Romand d’entomologie 4: 155-158.

BESUCHET, C., BURCKHARDT, D. & LÔBL, I. 1987. The "Winkler/Moczarski" eclector as an
efficienr extractor for fungus and litter Coleoptera. The Coleopterists Bulletin 41(4):
392-394.

512 P. HLAVAC

Hi:

BESUCHET, C. 1993a. Ein neuer Käferkatalog für die Schweiz. Mitteilungen der
Schweizerischen entomologischen Gesellschaft 43(3): 90-100.

. BESUCHET, C. & MERKL, O. 1993b. Scydmaenidae, Ptiliidae and Pselaphidae (Coleoptera)

from the Bükk National Park. /n: The Fauna of the Bükk National Park. Hungarian
natural History Museum, Budapest 99-103.

. BESUCHET, C. 1994. Les insectes. Le Malagnou 2: 25-27.
. BESUCHET, C. 1998. Des petits clandestins. Le Malagnou 4: 28-31.

. Popular & Others

127

13;

14.

157

. BESUCHET, C. 1957. Une technique nouvelle pour la préparation de l’édéage des

Microcoléoptères. Mitteilungen der Schweïizerischen entomologischen Gesellschaft 30:
341-342.

. BESUCHET, C. 1958. Un insecte parasite du castor. Bulletin Mensuel des Musées et

Collections de la ville de Genève, 15me année, No. 10.

. BESUCHET, C. 1962. Les myrmécophiles. Bulletin de la Société Vaudoise d’Entomologie, 3:

13-16.

. BESUCHET, C. 1971. Recherches entomologiques à Ceylan. La Revue "Musées de Genève"

No. 116: 3-6.

. BESUCHET, C. 1973. Les Rhinogrades. La Revue "Musées de Genève" No. 137: 10-13.
. BESUCHET, C. 1977. Les Afta, fourmis champignonnistes. La Revue "Musées de Genève"

No-172:2-11;

. BESUCHET, C. 1977. Les Afta, fourmis exotiques champignonnistes. Arve-Léman Nature,

LS ES KE

. BESUCHET, C. 1978. Les Atta, Fourmis champignonnistes. L’Entomologiste, 34(6): 221-231.
. BESUCHET, C. 1979. Que deviennent les Arta? La Revue "Musées de Genève" No. 196:

15-18.

. BESUCHET, C. 1979. Un insecte parasite du Castor. Arve-Léman Nature, 33: 15-17.
. WUEST, J. & BESUCHET, C. 1981. La colonie d’Atrta cephalotes du Muséum de Genève.

Comptes rendus, Assemblée générale, Toulouse, Section française. Union internationale
d’étude des Insectes sociaux, 131-132.

BESUCHET, C. 1983. Rôle du travail sur le terrain et particulièrement des prospections ento-
mologiques outre-mer. Bulletin de la Société entomologique de France, 88: 433-435.
BESUCHET, C. 1986. Les Animaux cavernicoles + Résultats d’une campagne de piègeage
dans les Alpes vaudoises. Le Trou, Groupe Spéléo Lausanne, 44: 16-20.

BESUCHET, C. 1992. Vers un "Catalogue des Coléoptères de Suisse". Bulletin Romand
d’Entomologie, 10: 49-55.

BESUCHET, C. & REZBANYAI-RESER, L. 1999. Angaben zur Verbreitung und Phänologie des
Fanghaftes, Mantispa styriaca (Poda, 1761) (Neuroptera: Mantispidae) in der Schweiz.
Mitteilungen der Entomologischen Gesellschaft Basel, 49(1): 8-18.

REVUE SUISSE DE ZOOLOGIE 118 (3): 513-559; septembre 2011

New species in the Zelotes tenuis-group and new or little known
species in other Zelotes groups (Gnaphosidae, Araneae)

Antoine SENGLET
Route de Begnins, 19, CH-1267 Vich, Switzerland. E-mail: a.senglet@bluewin.ch

New species in the Zelotes tenuis-group and new or little known species
in other Zelotes groups (Gnaphosidae, Araneae). - Notes are given on
mating mechanisms of Z. tenuis and Z. semirufus in the Z. fenuis-group.
Eight species are recognized in the Z. fenuis-group. The following
synonyms and new combinations are proposed: Trachyzelotes manytchensis
Ponomarev & Tsvetkov = Zelotes manytchensis, Z. ruscinensis Simon =
Z. semirufus (L. Koch), Z. fuscotestaceus (Simon) = Z. tenuis (L. Koch),
Z. denisi Marinaro = Z. criniger Denis and Z. sumchi Levy = Z. metellus
Roewer. Z. babunaensis (Drenski) is revalidated. First description are given
of the male of Z. babunaensis, the male of Z. metellus, the female of Z. fla-
gellans (L. Koch), and of the following eight species: Z. alpujarraensis
Sp. n., Z. baeticus Sp. n., Z. chaniaensis Sp. n., Z. cordubensis sp. n., Z. egre-
gioides Sp. n.,Z. hispaliensis Sp. n., Z. laconicus Sp. n. and Z. pediculatoides
Sp. n.

Keywords: Arachnida - taxonomy - mating mechanism.

INTRODUCTION

The study of the & pedipalp in Zelotes has led to the grouping of species. The
Zelotes subterraneus group (Senglet, 2004 106) and the Z. petrensis group (Senglet,
2004: 111) have been discussed earlier. Five additional groups are treated here, the
Z. tenuis, Z. thorelli, Z. atrocaerulaeus, Z. baeticus and Z. metellus-groups.

MATERIAL AND METHODS

Except for the © of Z. prishutovae and the types specimens of Zelotes semi -
rufus, Z. fulvaster and Z. fulvopilosus, all spider material was collected by myself.
Measurements are in millimetres. Vulvae were examined on an excavated microscopic
slide, in lactic acid. Holotypes and paratypes of the species described in here are
deposited in the Natural History Museum of Geneva (MHNG); the other material, 1f
not indicated otherwise, remains in my collection. For details on rearing and cryo-
fixing of mated spiders, see Senglet (2004: 87).

The typical leg spination according to Platnick & Shadab (1983) is: Femora; I,
IT d110, p001; IT, IV d110, p011, r011; patella II r010; tibiae: II p111,v222,r011; IV
222 0 /nEétatarsi LI V20011Pp122/V221, rM2"IN p122,v220;r122. Only
differences to this pattern are given in the text. AME, ALE, PME, PLE and MO refer

Manuscript accepted 03.06.2011

514 A. SENGLET

to anterior median, anterior lateral, posterior median, posterior lateral eyes and to the
median ocular quadrangle (eyes included).

Terminology of genital structures follows Senglet (2004: 88-90). Additional
comments on some structures are:

Embolar radix: This structure (Figs 8-10; Senglet, 2004: 88, fig. la, 1d) seems
to be present in many Zelotinae. In 7rachyzelotes and in many Zelotes groups it is a
solid sclerite at the posterior base of the embolus; in some cases it is articulated or
flexible.

Embolar base: A term for the ventral base of the embolus. Present in different
Zelotinae genera; originating from the prolateral base of the tegulum, it is usually
connected to the embolus with a flexible joint. There 1s no connection to the embolus
in the Zelotes thorelli-group.

Posterior sclerite of terminal apophysis: Where present in Zelotinae, the termi-
nal apophysis 1s a ventral sclerotized extension of the embolar base. The presence of a
posterior sclerite of the terminal apophysis (absent in Drassyllus and Trachyzelotes) is
a synapomorphy of Zelotes s. str. This sclerite may be reaching the root of the radix
(Z. subterraneus, Z. longipes and Z. fhorelli-groups), the middle sector of the radix
(Z. tenuis-group), or even the embolus (Z. baeticus-group).

Intercalary sclerite: Rightly considered as a synapomorphy of the genus by
Platnick & Shadab (1983: 100, fig. 2), it is a widening of the proximal part of the em-
bolar base. Present only in conjunction with the posterior sclerite of the terminal
apophysis, highly variable and often reduced, it seems that its basic function 1s to of-
fer a flexible joint to the terminal apophysis fastened to the posterior segment of the
tegulum (Senglet, 2004: fig. 13).

TAXONOMY AND FAUNISTIC DATA
Genus Zelotes Gistel, 1848
TYPE SPECIES: Zelotes subterraneus (C. L. Koch, 1833)

KEY TO SPECIES GROUPS

Il Embolar radix with sclerotized connection to embolus; no dorsal apo-
physisonembolat base 4 Es LEP ES DRE Zelotes s. str., 2
- Embolar radix with a membranous connection to embolus; a dorsal
apophysis present on embolar base ............ Z. criniger-metellus-group
2 Embolar base a simple transverse sclerite with a retrolateral projection
and a notched link to embolus; posterior sclérite of terminal apophysis
connected:toipostenortesulanbase 2 Z. subterraneus-group
- Different . ::.::2454e80 228 RE 3
3 Embolus not coïlediz. 4.824802 OR RES RER RER À

- Long coiled embolus (turning left on left palp) without direct connection
to embolar base. Embolar base fused to terminal apophysis up to its api-
cal projection. Strong embolar radix and posterior sclerite of terminal
apophysis at a relatively acute angle (Figs 83, 92) ........ Z. thorelli-group
- Elevated arched embolar base with a distad-directed embolus. Posterior
sclerite of terminal apophysis connected to median part of embolar

NEW ZELOTES SPECIES 515

radix. Epigynum with single lateral folds; median sector membranous .

A ES SRE ds a on Se NU Z. tenuis-group
- Embolar base partly fused with terminal apophysis. Terminal apophysis

with ventral and dorsal lobe; its posterior sclerite connected to posterior

tegular base (Figs 139-141, 146-147). Epigynum with single lateral folds

I nn Lu 2 tee ma ue Z. atrocaerulaeus-group
- Embolar base elevated and arched; posterior sclerite of terminal apo-

physis connected directly to embolus (Fig. 133); embolus linked to

embolar base by whitish cuticle along its prolateral margin (Figs 120,

127, 131 & 133); the large embolus carrying a terminal segment in the

shape of a wide tube; its variable ventral protrusion corresponding to

prolateral projection of median apophysis. Lateral folds of epigynum

modified into large lateral pockets; median part membranous (Figs 123,

RO bn are er à Z. baeticus-group

Zelotes tenuis-group

DEFINITION: The male pedipalp has an embolar radix, an intercalary sclerite and
an elevated arched embolar base with a distad-directed embolus. The terminal apo-
physis has a posterior sclerite connected to the median part of the radix in Z. subterra-
neus and Z. longipes-groups (Senglet, 2004: figs la, 1d, 12-14), the connection 1s
situated below the base of the embolar radix. The arched embolar base has a variable
tooth, which is replaced in Z. manitchensis and Z. alpujarraensis sp. n. by a retrolateral
projection. The female has plain lateral epigynal folds enclosing a membranous cuticle.
Except for Z. alpujarraensis, the type of ocular group shown in Figs 54-55 1s shared
by all species of the Z. fenuis-group; the PME are larger or equal to the PLE and
separated by 15 to 35% of their diameter. In Z. alpujarraensis the posterior eyes are of
equal size, separated by 70% of their diameter.

MATING MECHANISM: In Zelotes tenuis (Fig. 1) and Z. fuscorufus (Senglet,
2004: 32-34) the apical part of the embolar base is inserted into the posterior segment
of the epigynal pocket, putting the embolus in contact with the insemination pore. In
Zelotes semirufus (Figs 2-4), however, the proximal apophysis of the embolar base
(Figs 2, 11-12) is inserted into the swollen membranous median cuticle of the epi-
gynum (Fig. 3); the broken prolateral hook of the embolar base of Fig. 4 remains inside
the cuticle in Fig. 3; the transverse cuticle fold filling the gap between embolus and
prolateral embolar base apophysis is visible. The same type of folded median mem-
branous part is found in Z. chaniaensis sp. n. and Z. fulvaster. Figures 5-11 show
details of the male bulbus in copula.

KEY TO THE SPECIES OF THE ZELOTES TENUIS-GROUP

D Mn le... lun. 1
2 ot mt deu laseinmmaiauns de... 8
Il ORAN OC CMD AEIDasSE CORVEx:l 1 Luca failli veih. ... 3

- Apical margin of embolar base concave between distal tooth and
A abandon ADI EE RE. à 2

s16 A. SENGLET

FIGS 1-4

(1) Zelotes tenuis, epigynum in copula, male palp extracted. (2-4) Zelotes semirufus, left male
palp extracted in copula. (2) Left male palp, apical view (compare with Fig. 11). (3) Epigynum.
(4) Broken left male palp (inserted in epigynum shown in Fig. 3).

2 Large apophysis on embolar base, situated prolaterally (Fig. 13) . Z. semirufus
- Small hook-shaped apophysis on embolar base, situated close to

embolus Figs 33-34). be RER NN EN CN Z. fulvaster
ë, A retrolateral projection on embolar base (Figs 61-62, 66-69) ............ 7
- No retrolateral projecuionest ec ER CNP NES RER À
4 Retrolateral tibial apophysis ribbon-like, bifid (Fig. 20) ....... Z. fuscorufus
- Retrolateraliubialapophysis pen 5
5 A small retrolaterally directed apical hook on embolar base (Fig. 53) ..

ED de D Z. chaniaensis sp. n.
- Asmalltrnanpgularapicaltoothonembolanbase "+7 tree ee 6
6 Retrolateral tibial apophysis longer than dorsal length of tibia; width of

pedipalpal tibia more than 3/4 of its dorsal length (Fig. 28); embolus
short(Figs 22-24): tee MORE RE Z. babunaensis

NEW ZELOTES SPECIES 517

post. tegular base

btegular rid i
suvtégular ridge subtegulum subtegular ridge

intercalary sclerite

term. apoph. é Lies median embolar base - 5 K ,._——embolus
post.sclerite IE Rss FPOP ee | . 9
2 RATE intercalary sclerite
embolar
radix ;
terminal apoph. à
embolus À) ERA ; embolar base a cymbium
8 És embolar base tegulum subtegulum
prolateral apoph.
terminal
Forte, apoph. intercalary sclerite
broken
median apoph. F: M” embolar base
3 ne prolateral apoph.

term. apoph.
post. sclerite

terminal apoph. Pr

embolar base glandular duc

subtegular ridge

embolar base / = embolus
prolateral apoph. 10

FIGS 5-11

Zelotes semirufus, left male palp in copula, viewed from the female opisthosoma. (5) Anterior
view. (6) Posterior view. (7) Left view. (8) Id., detail. (9) Right view. (10) Posterior view, detail.
(11) Palp on female with vulva in dorsal view. Bold lines indicate female parts. Scale lines
0.2 mm.

518 A. SENGLET

- Retrolateral tibial apophysis shorter than dorsal length of tibia; width of
pedipalpal tibia 2/3 of its length or less (Figs 40-43) ............. Z. tenuis

g Retrolateral embolar base projection slender, long, curved (Figs 66-69) .....
TN UE du Me us Doha Z. alpujarraensis sp. n.

- Retrolateral embolar base projection short, membranous (Figs 60-62) ..
RE Lin Z. manitchensis

8 Epigynal plate much wider than long (Figs 21, 63,73 & 77)............14
- Epigynal plate longer than wide or slichtiywider...,. 28... 9
9 Lateral epigynal fold more or less curved, with a posterior rounded bend . . 10

- Lateral epigynal fold almost straight, with an acute posterior bend;
lateral vulval pouch posteriorly widened; median ducts connected on

médian side of spermathecac (Fies 37-59 V5) 0 Z. fulvaster
10 Width of anterior epigynal margin equal to or less than epigynal width ..... 11
- Anterior epigynal margin wider than epigynal width (Figs 29, 57) ........ 12
11 Short median ducts leaving the median epigynal cuticle undivided (Figs

LOST SERRE ce ... DORE Z. semirufus
- Posterior part of median cuticle divided by median ducts and folds (Figs

AS AO). a 2 80 MO de ie AMOR NP RE EEE Z. tenuis
12 Short median ducts leaving the median epigynal cuticle undivided (Figs

TOP DEEE 2 CRE TT LL TR CR EE Z. alpujarraensis sp. n.

Posterior area of median cuticle divided by median ducts and folds . ...... 13
13 ShOrt CUrVvEd'meduan ducts (Fies 57-SS) RE Z. chaniaensis sp. n.
- Straight or slightly curved long median ducts with an acute bend, close

to thé inseminatonpore (Fiss 30-35 ee Re Z. babunaensis
14 Short median ducts (Fig -210)47 52 SR RE Z. fuscorufus
- Bons médian ducs iss 63-64)... RE Z. manytchensis
Zelotes semirufus (L. Koch, 1882) Figs 2-17, 74

Prosthesima semirufa L. Koch, 1882: 636, pl. 20, fig. 15 (description of ©).
Zelotes ruscinensis Simon, 1914: 157, 169, 219, figs 295, 346 (description of d) syn. n. —
Senglet, 2004: 104, figs 47-50.
For previous synomymy, see Senglet (2004: 104) and Platnick (2011).
TYPE MATERIAL: BM1915.3.5.6100, Natural History Museum (London); 1 ® from
Menorca; received on loan through the courtesy of Mrs Janet Beccaloni.

OTHER MATERIAL EXAMINED: SPAIN, Levant / Murcia, Alicante, Elche (in palm grove),
38°17'N 00°42'W; 1 & (last moult 08.07.2002); 15.05.2002. — Estremadura, Caceres, west of
Jarandilla (litter in vegetation), 40°08'N 05°40'W; 1 © (last moult 17.08.2009); 16.06.2009. —
Andalusia, Cordoba, Almodôvar del Rio (Breña dam), 37°50'N 05°04'W; 1 © (last moult
18.08.2009); 01.06.2009. — Huelva, Puerto Gil /Aracena (cork-oak leaf litter), 37°53'N 06°29'W;
1 ® (last moult 21.08.2009); 07.06.2009. — Huelva, Alajar /Aracena, 37°53'N 06°40'W; 6 G,
5 © (last moults of & 02 to 28.07.2009, of $ 12.07 to 23.08.2009); 09.06.2009.

DIAGNOSIS: The male of Z. semirufus is easily distinguishable from those of
other species of the tenuis-group by its large prolateral hook on the embolar base, and
the female by its wide lateral pouches with folded cuticle, its narrow anterior epigynal
margin and its widely separated median ducts (Figs 12-17, 74).

DESCRIPTION: See Senglet, 2004: 104.

NEW ZELOTES SPECIES 519

FIGsS 12-21

(12-17) Zelotes semirufus. (12-15) Left male palp. (12) Ventral view. (13) Distal part, cleared,
retrolateral view. (14) Id., dorsal view. (15) Tibia, retrolateral view. (16) Koch's type, epigynum.
(17) Id., vulva, dorsal view. (18-21) Zelotes fuscorufus. (18-20) Left male palp. (18) Ventral
view. (19) Prolateral view. (20) Tibia, retrolateral view. (21) Epigynum (on left) + vulva, dorsal
view. Bold lines indicate epigynal folds. Scale lines 0.2 mm.

520 A. SENGLET

Zelotes fuscorufus (Simon, 1878) Figs 18-21, 73

Prosthesima fusco-rufa Simon, 1878: 95, pl. 14, fig. 28 (description of 4 & ®).

Zelotes fuscorufus.-— Senglet, 2004: 102, figs 1b-c, 32-34, 51-54.

For redescription and previous literature, see Senglet (2004: 102) and Platnick (2010).
DIAGNOSIS: The male of Z. fuscorufus is easily distinguishable from males of

other species of the tenuis-group by the bifid retrolateral tibial apophysis and the more

distad-directed embolar base, and the female by its epigynum wider than long (Figs

18-21, 73).

Zelotes babunaensis (Drensky, 1929), Figs 22-31,76
here removed from the synonymy of Z. fenuis

Echemus babunaensis Drensky, 1929: 5, 59, pl. 1, figs 3-4 (description of 9).

Zelotes babunaensis.— Deltshev & Blagoev, 2001: 110 (transfer of $ from Echemus).

Zelotes tenuis.-— Deltshev, 2003: 137, fig. 8, not figs 9-10 = Z. renuis (synonymy of © , misiden-
tification).— Chatzaki ef al., 2003: 80, figs 108-110, 113-114 (misidentification), not figs
105-107, 111-112 = Z. chaniaensis sp. n.

MATERIAL EXAMINED: GREECE, Macedonia, Thessaloniki, Loutra Apollonias (in litter),
40°39'N 23°24'E; 1 9 (last moult 17.09.2004); 09.09.2004. — Epiros, Thesprotia, near Neraï da,
39°31'N 20°24'E; 1 G ; 21.06.1998. — Thessalia, Larissa, Omélion-Tempé, 39°53'N 22°37'E;
1 & (1 palpus lost), 1 8 ; 17.06.1970. — Sterea Hellas, Attiki, Marathon lake, 38° 10'N 23°54'E;
1 ® (with vulva in microvial); 11.07.1968. — Phthiotidas, Trägana, 38°37'N 23°07'E,; 1 d;
13.07.1968. — Phthiotidas, Theologos, 38°39'N 23°12'E; 1 d,2 © (with vulva in microvial, last
moults of $ 22.5, of $ 27.05 and 25.06.1998); 20.05.1998. — Eubea, Loutra Aidipsou, 38°53'N
22°59'E; 1 d; 29.05.1983. — Peloponnesus, Laconia, east of Lira (in Pinus litter), 36°39'N
22°5S8'E; 1 &, 1 S(vulva in microvial, last moults of 4 06.06.2005, of 9 30.06.2005);
03.10.2004. — Argolida, east of Ligourion (in Lentiscus litter), 37°38'N 23°04'E; 1 d (last moult
05.06.2005); 01.10.2004. — Argolida, Palaia Epidauros, 37°39'N 23°09'E; 1 ® (vulva lost);
05.06.1981. — Cyclades, Naxos, north-west of Sangri, 37°03'N 25°27'E; 1 d (last moult
06.06.1998); 05.06.1998. — Cyclades, Koronido (650-750m), 37°09'N 25°37'E; 1 4,2 (last
moult of 1 8 28.06.1998); 08.06.1998. — Cyclades, Kato Potamia, 37°06'N 25°26'E; 2 à ,1 %
(last moult of 8 12.06.1998); 10.06.1998. — Cyclades, Dimitra /Ano Sangri, 37°02'N 25°26'E;
2 4,1 9 (last moults of 1 4 12.06, 8 22.07.1998); 11.06.1998.

DESCRIPTION: Prosoma tawny-brown, with faint blackish markings. Opistho-
soma blackish, with medium-long dark hairs. Legs blackish to middle of femora, tarsi
tawny. Leg spination: Metatarsi I VO00; IT v220. — 4 from Naxos: Total length 6.33.
Prosoma 2.50 long, 1.90 wide, 1.00 wide at level of posterior eyes. Eye sizes and
interdistances: AME 0.07, ALE 0.13, PME 0.11, PLE 0.09; AME-AME 0.06, AME-
ALE 0.01, PME-PME 0.03, PME-PLE 0.05, ALE-PLE 0.06. MOQ length 0.29, front
width 0.44, back width 0.50. Clypeus: 0.11 from AME, 0.07 from ALE. Pedipalp (Figs
22-28): Patella dorsally longer than tibia. Strong and short pedipalpal tibia. Retrolateral
tibial apophysis equal or longer than dorsal length of tibia (shorter in Z. fenuis) and
width of pedipalpal tibia less than dorsal length of tibia (Fig. 28). Embolus short (F1gs
22-24). Scutum 1/5 of opisthosoma length. — ® from Naxos: Total length 6.00. Pro -
soma 2.20 long, 1.60 wide, 0.92 wide at level of posterior eyes. Eye sizes and inter-
distances: AME 0.07, ALE 0.10, PME 0.10, PLE 0.08; AME-AME 0.03, AME-ALE
0.01, PME-PME 0.03, PME-PLE 0.03, ALE-PLE 0.06. MORQ length 0.23, front width
0.38, back width 0.45. Clypeus: 0.08 from AME, 0.06 from ALE. Epigynum (Fig. 29):
Anterior anchoring pockets wider than lateral folds. Vulva (Figs 30-31): Long median
ducts, straight or slightly curved, with a sharp bend, close to the insemination pore.

NEW ZELOTES SPECIES s21

22 23

embolar

embolar apophysis haematodocha

embolar
radix

embolar base

terminal apophysis posterior sclerite 26

EC
PT Dee ER LITE
-
LL
1
me”

en
mm

FIGS 22-31

Zelotes babunaensis. (22-28) Left male palp. (22) Ventral view (median apophysis expanded),
from Crete. (23) Id., from Eubea. (24) Id., from Naxos. (25) Id., retrolateral view. (26) Distal part
cleared, dorsal view. (27) Apical view. (28) Diagram of tibia, retrolateral view. (29) Epigynum
from Eubea. (30) Id., vulva, dorsal view. (31) Id., from Naxos. Bold lines indicate epigynal folds.
Scale lines 0.2 mm.

322 A. SENGLET

REMARKS: Drensky's (1929: pl. 1, fig. 3) and Deltshev’s (2003: fig. 8) illus-
trations leave no doubt about the validity of this species. It is more abundant in Greece
than the rare Z. tenuis, both are sympatric with Z. fulvaster.

Zelotes fulvaster (Simon, 1878) Figs 32-39, 75

Prosthesima fulvastra Simon, 1878: 96, pl. 14, fig. 30 (description of ®).

Zelotes fulvaster.— Simon, 1914: 168, 219, fig. 345 (description of 9). — Jézéquel, 1962: 603,
fig. 26 (description of 9).

Zelotes tenuis.— Deltshev, 2004: 72, figs 9-10 (misidentification).

TYPE MATERIAL: Muséum National Histoire Naturelle, Paris, collection Simon, jar 568,
no. 1748, 1 © deprived of epigynum and 1 slide PM52, presumably from Porto Vechio, Corse
(Simon, 1878: 96 and Jézéquel, 1962: 603).

OTHER MATERIAL EXAMINED: GREECE, Macedonia, Thessaloniki, Aghios Vassilios,
40°40'N 23°07'E; 1 9% (last moult 25.07.1998); 14.06.1998. — Thessaloniki, West of Aghios
Vassilios (vegetation), 40°41'N 23°05'E; 2 d, 1 ® (last moults of 4 25 and 27.06, of 9%
27.06.2008); 31.05.2008. — Thessaloniki, East of Apollonia (under stones), 40°37'N 23°32'E;
1 G; 01.06.2008. — Sterea Hellas, Phthiotidas, Theologos (Glyphada), 38°39'N 23°14'E; 2 9
(with vulva in microvial); 20.06.1970. — Phthiotidas, Near Malesina, 38°37'N 23°13'E; 1 à (last
moult 09.06.1998); 21.05.1998. — Same; 1 & ,1 9 (last moult of 9 12.06.2008); 08.06.2008. —
Phthiotidas, north of Malesina, 38°38'N 23°14'E; 1 9 ; 19.06.2008. — Eubea, Theologos, 38°29'N
23°4T'E; 1 4,1 © (last moult of 9 28.06.2008); 11.06.2008. — Peloponnesos, Argolida, Palaia
Epidauros, 37°39'N 23°09'E; 1 © (last moult 29.07.1998); 14.05.1998. — Cyclades, Naxos, north
of Aghios Prokopios (under vegetation), 37°05'N 25°21'E; 2 4 (with palpus in microvial),
1 $ (last moults of 4 14. And 18.06, of 9 24.07.1998); 06.06.1998. — Cyclades, Naxos, Kato
Potamia, 37°06'N 25°26'E; 1 9% (last moult 25.06.1998); 10.06.1998. IRAN, Esfahan,
Falävariän, 32°34'N 51°31'E,; 1 d ; 14.06.1974. — Fars, Khohkiluyeh, Arow, 30°35'N 50°43'E;
1 4,2 © (with palpus and vulva in microvial); 24.05.1974. — Khohkiluyeh, Basht, 30°20'N
51°15'E; 2 9; 25.05.1974. — Khohkiluyeh, Bishäpour, 29°47'N 51°35'E; 1 d ; 28.05.1974. —
Khohkiluyeh, Aliäbäd, 30°01'N 53°00'E; 1 % ; 09.06.1974.

DESCRIPTION: Prosoma tawny, With faint blackish marking. Opisthosoma dor-
sally blackish, ventrally grey, covered with medium-long dark hairs. Legs tawny.
Iranian specimens entirely tawny. — 4 from Naxos (in parentheses from Arow, Iran):
Total length 4.75. Prosoma 1.86 (2.00) long, 1.40 (1.46) wide, 0.70 (0.80) wide at level
of posterior eyes. Eye sizes and interdistances: AME 0.06 (0.07), ALE 0.10 (0.11),
PME 0.11,PLE 0.07 (0.86); AME-AME 0.06, AME-ALE 0 .01,PME-PME 0.01, PME-
PLE 0.02, ALE-PLE 0.04. MOQ length 0.24, front width 0.34, back width 0.41.
Pedipalp (Figs 32-36): Tibia and patella dorsally of same length. Width of pedipalpal
tibia and length of retrolateral apophysis about equal to dorsal length of tibia (cf. Fig.
28). Median apophysis with a large rounded concave retrolateral flap. Small hook-
shaped, retrolaterally directed apical projection present on apical margin of embolar
base (Figs 33-34). Hook separated from the short embolus by a concave ventral margin
of embolar base. Retrolateral tibial apophysis dorsally arched. Scutum occupying 30%
of opisthosoma length. Leg spination: Metatarsus I v000; some large Iranian specimens
with additional retrolateral spines on metatarsi III & IV. — ® from Naxos: Total length
5.20. Prosoma 2.25 long, 1.75 wide, 0.91 wide at level of posterior eyes. Eye sizes and
interdistances: AME 0.08, ALE 0.12, PME 0.12, PLE 0.09; AME-AME 0.04, AME-
ALE 0.01, PME-PME 0.03, PME-PLE 0.04, ALE-PLE 0.04. MOQ length 0.30, front
width 0.42, back width 0.47. Epigynum (Figs 37, 75): Side of epigynal fold almost
straight, making an acute posterior bend. Posterior part of epigynal pouch strongly

NEW ZELOTES SPECIES 323

ee
| à 7

FIGs 32-39

Zelotes fulvaster. (32-36) Left male palp. (32) Prolateral view. (33) Ventral view. (34) Id. Iranian
specimen. (35) Retrolateral view. (36) Apical view. (37) Epigynum. (38) Vulva, dorsal view. (39)
Id., Type "PMS2" in MNEN Paris. Bold lines indicate epigynal folds. Scale lines 0.2 mm.

widened. Vulva (Figs 38-39): Lateral pouch widened, dorsally with a cuticular fold
along its posterior margin, this fold sometimes visible on epigynum as an oblique
darker line. Leg spination: Metatarsus I v000.

Zelotes tenuis (L. Koch, 1866) Figs 40-49, 72
Drassus tenuis L. Koch, 1866: 101, pl. 4, figs 65-66 (description of à).

Zelotes tenuis.— Platnick, 1989: 489 (new combination). — Levy, 1998: 131, figs 78-81.
Prosthesima fusco-testacea Simon, 1878: 97, pl. 14, fig. 31 (description of ©) syn. n.

524 A. SENGLET

Zelotes fuscotestaceus.-— Simon, 1914: 168,218, fig. 344 (description of 9 ).— Denis, 1952: 123,
fig. 19. — Jézéquel, 1962: 604, fig. 30.

Prosthesima circumspecta Simon, 1878: 94, pl. 14, fig. 26 (description of 4 and 9). Chyzer
& Kulczynski, 1897: 205, pl. 8, figs 2, 20.

Zelotes circumspectus.-— Simon, 1914: 157,168, 219, figs 295-296, 342-343. - Denis,1952: 123,
fig. 20. — Jézéquel, 1962: 604, fig. 27.

Prosthesima pallida O. P. -Cambridge, 1874: 383, pl. 51, fig. 11 (description of 4 and ©).

Zelotes pallidus.-— Platnick & Shadab, 1983: 185, figs 259-262 (description of 4 and 9).

MATERIAL EXAMINED: GREECE, Macedonia, Thessaloniki, Aghios Vassilios, 40°40'N
23°07'E; 1 4,3 9 (last moults of 9 18-27.06.1998); 14.06.1998. — Thessalia, Larissa, Omélion-
Tempé, 39°53'N 22°37'E; 1 9 ; 17.06.1970. — Sterea Hellas, Phthiotidas, Theologos (Glyphada),
38°39'N 23°14'E; 1 9 ; 20.06.1970. — Same; 1 & (last moult 18.06.2005); 04.06.1978. — West of
Malesina, 38°37'N 23°13'E; 1 d ; 25.09.2004. — Same; 1 9 (last moult 12.06.2008); 08.06.2008.
— SPAIN, Catalonia, Barcelona, Gelida, 41°27'N 01°51'E; 1 4 (with darker opisthosoma, 1 juv.);
14.06.1971. — Levant / Murcia, Valencia. La Albufera, 39°21'N 00°19'W; 2 & ; 16.06.1971. —
Montroy, 39°20'N 00°35'W; 1 %; 22.06.1971. — Alicante, Elche (under palm grove), 38°17'N
00°42'W; 4 4,3 % (last moults of 1 & 05.06.2002, of 8 11-14.06.2002); 15.05.2002. — Elda,
38°30'N 00°47'W; 2 & ; 19.06.1971. —- Murcia, Archena, 38°07'N 01°17'W; 1 9; 30.06.1971. —
Same; 2 4,2 % (last moults of 4 26.05.2002, of £ 10 and 21.07.2002); 17.05.2002. — Castilla
/ Leon, Valladolid, Peñafiel, 41°35'N 04°08'W; 1 G ; 23.06.2002. - Nueva Castilla / La Mancha,
Madrid, Navalcarnero, 40°18'N 03°56'W; 2 G; 12.06.1969. — Madrid, Aldea del Fresno (rio
Alberche), 40°19'N 04°13'W; 1 9; 13.06.1969. — Toledo, Escalona del Alberche, 40°10'N
04°24'W; 1 G: 14.06.1969. — Toledo, Cardiel de los Montes (rio Alberche), 40°02'N 04°39'W:
1 4,1 ©; 15.06.1969. — Toledo, Urda /Consuegra, 39°25'N 03°42'W; 2 ©; 12.08.1969. -
Albacete, Hellin, 38°29'N 01°37'W; 1 G ; 29.06.1971. — Albacete, La Gineta (rio Jucar, fine leaf
litter), 39°10'N 01°58'W; 6 G ; 16.05.2002. — Ciudad Real, Caracuel (laguna), 38°50'N 04°04'W;
3 ©: 04.08.1969. — Estremadura, Caceres, Jarandilla (rio Tietar, in litter on sand), 40°01'N
05°37'W; 1 4,4 9 (last moult of 1 9 03.08.2009); 15.06.2009. - Caceres, west of Jarandilla (in
litter in vegetation), 40°08'N 05°40'W; 2 4,1 9 (last moult of £ 22.06.2009); 16.06.2009. —
Badajoz, south of Venta del Culebrin /Monesterio, 37°58'N 06°14'W; 1 d; 19.06.1969. —
Badajoz, Venta del Culebrin /Monesterio, 38°01'N 06°13'W; 11 &,1 %; 19.06.1969. — Same;
3 G (last moults of 2 &, 07 and 25.06.2002); 04.06.2002. — Badajoz, Rio Guadalemar
/Garbayuela, 39°03'N 04°59'W; 2 9; 17.08.1969. — Badajoz, Rio Sillo (Higuera la Real),
38°06'N 06°41'W; 2 G (last moult of 1 4 11.06.2002); 09.06.2002. — Badajoz, south of
Monesterio (under stones in vegetation), 38°03'N 06°14'W; 10 4,5 % (last moults of 3 d
6-12.06.2010, of 4 £ 07.5-20.06.2010); 05.06.2009. — Andalucia, Almeria, Adra (La Albufera),
36°45'N 02°57'W; 2 9 ; 08.07.1971. — Same (cultivated dry pond), 36°46'N 02°58'W; 4 4,2 9;
25.05.2002. — Granada, La Räbita /Albuñol, 36°45'N 03°10'W; 3 9; 09.07.1971. — Ugijar,
36°59'N 03°04'W; 1 9: 15.07.1971. — Jaen, Cañada de las Hazadillas (under Pinus), 37°39'N
03°43'W; 3 4,1 @ (last moults of & 03.06.2002, of 8 11.07.2002); 30.05.2002. — Cordoba,
Palma del Rio; 1 G; 03.06.1967. — Cordoba, Palma del Rio, 37°43'N 05°18'W; 4 4,3 9;
26.06.1969. — Cordoba. Almodévar del Rio (Breña dam). 37°50'N 05°04'W; 1 9 ; 28.06.1969. —
Cordoba, Pantano de la Breña (evergreen oak litter), 37°51'N 05°04'W; 5 4,8 9 (last moults of
4 G 2-12.06.2002, of ? 02.6-16.07.2002); 01.06.2002. — Cordoba, Almodôévar del Rio (Breña
dam), 37°50'N 05°04'W; 1 & (last moult & 14.06.2009); 01.06.2009. — Cordoba, Peñarroya,
38°17'N 05°16'W; 1 4,3 9 (with vulva in microvial); 30.06.1969. — Cordoba, Los Villares
/Cordoba (Eucalyptus litter), 37°58'N 04°49'W; 1 & (last moult 02.06.2009); 31.05.2009. —
Cordoba, Palma del Rio (Retortillo dam, leaf litter), 37°51'N 05°22'W; 1 &: 03.06.2009. —
Malaga, Ronda, 36°46'N 05°13'W; 1 %; 21.07.1969. — Malaga, Estepona, 36°25'N 05°11'"W;
1 ®; 25.07.1969. — Malaga, Torre de Mar, 36°44'N 04°07'W; 2 9 ; 27.07.1969. — Malaga, Valle
de Abdalagis /Antequera, 36°56'N 04°41'W; 1 9; 28.07.1969. — Sevilla, Lebrija (rio. del
Salado); 2 & ; 06.06.1967. — Sevilla, Alcala del Rio, 37°31'N 05°59'W; 3 9; 22.06.1969. —
Sevilla, rio Viar /Castilblanco, 37°42'N 05°53'W; 1 © ; 24.06.1969. — Same, 37°43'N 05°53'W;
8 4,10 ? (with palpus and vulva in microvial, last moults of 3 4 09.6-9.07.2002, of 6 © 07.6-
27.07.2002); 02.06.2002. — Sevilla, Cantillana, 37°37'N 05°50'W; 1 d ; 26.06.1969. — Sevilla,
Alanis, 38°02'N 05°11'W; 1 & ; 01.07.1969. — Sevilla, Sanlücar la Mayor, 37°22'N 06° 14'W;

NEW ZELOTES SPECIES 525

terminal apoph.
posterior sclerite

RLCTTCLLS

ne
pen e

LE

FIGs 40-49

Zelotes tenuis. (40-44) Left male palp. (40) Greek specimen, ventral view. (41) Id., retrolateral
view. (42) Spanish specimen, ventral view. (43) Id., retrolateral view. (44) Id., cleared, dorsal
view. (45) Greek specimen, epigynum. (46) Id., vulva, dorsal view. (47) Spanish specimen, epi -
gynum. (48) Id., vulva, dorsal view. (49) Id., variant. Bold lines indicate epigynal folds. Scale
lines 0.2 mm.

526 A. SENGLET

4 S; 17.07.1969. — Sevilla, Coripe (rio Guadalete), 36°58'N 05°26'W; 1 %; 18.07.1969. —
Sevilla, east of Cazalla de la Sierra (deep leaf litter), 37°57'N 05°45'W; 2 4,3 $ (last moults of
1 G 16.06.2009, of 2 9 29.06.2009); 04.06.2009. — Sevilla, Rivera de Benalija (Pintado dam),
38°02'N 05°55'W, 4 4,3 % (last moults of 2 & 5 and 12.06.2009, of $ 21.7-02.08.2009):
05.06.2009. — Cadiz, Algodonales, 36°53'N 05°27'W, 1 ©, 19.07.1969. — Huelva, Santa Olalla
(rio Cala), 37°55'N 06°11'W,2 % ,04.07.1969. — Same, 2 6 ,05.06.2002. — Huelva, Zufre (rivera
Huelva), 37°51'N 06°19'W, 1 ® ,04.07.1969. - Huelva, Zalamea la Real, 37°41'N 06°39'W, 1 &,
2 © ,10.07.1969. — Huelva, Moguer, 37°16'N 06°50'W, 1 % , 11.07.1969. — Huelva, Torre de Oro
/]Mazagén, 37°05'N 06°43'W, 1 ©, 13.07.1969. — Huelva, Alajar /Aracena, 37°52'N 06°40'W;
2 4,1 %; 07.07.1969. — Same (cork-oak leaf litter), 37°52'N 06°41'W; 1 &,1 % (last moult of
® 08.08.2009); 09.06.2009. — Huelva, Puerto Gil /Aracena (cork-oak leaf litter), 37°53'N
06°29'W; 10 G (3 large males, others small), 3 $ (last moults of 2 8 17.06 and 20.07.2009);
07.06.2009. — Huelva, Fuenteheridos (deep leaf litter), 37°54'N 06°40'W; 3 ,3 © (last moults
of 1 G 14.06.2009, of £ 14.06-02.07.2009); 12.06.2009. PORTUGAL, Extremadura, Setubal,
Azinheira dos Bairros, 38°04'N 08°25'W; 1 9 ;: 02.08.1971. FRANCE, Corse, Nord Corse, Ponte
Leccia; 2 G ; 02.06.1971. — Nord Corse, Above Ponte Leccia (under gravel, river); 4 @ , 10 9;
20.06.1999. — Nord Corse, Above Barchetta (Golo river, under dry vegetation); 2 4,2 9;
21.06.1999. — Nord Corse, Biguglia pond /Bastia; 1 ®; 23.06.1999. — Sud Corse, Ajaccio
(Gravone river); 1 4; 01.06.1971. — Sud Corse, Portigliolo /Propriano (under vegetation &
stones); 1 ® ; 17.06.1999. - Sud Corse, Sartène, Acorane bridge; 3 4,2 ® (last moults of $ 20
and 22.06.1999); 19.06.1999. — Provence / Côte d'azur, Var, Porquerolle (in litter), 43°00'N
06°12'E; 2 G ,4 9 (last moults of 4 13.06.2001, of $ 19.07.2001); 23.06.2001. — Languedoc,
Herault, Les Ouglou /Agde, (Etang de Thau), 43°20'N 03°33'E; 3 ® (last moult of 1 9%
08.07.2001); 26.06.2001. — Pyrenees Orientales, Arles sur Tech, 42°27'N 02936'E; 1 9;
01.07.2001. — ITALY, Sardinia, Sassari, Platamona /Porto Torres (damp leaf litter); 6 & ,4 9;
25.05.1999. — Sassari, Castelsardo:; 8 & ,5 © (last moults of 2 & 5 and 20.06.1999, of 4 © 05.06-
03.07.1999); 26.05.1999. — Sassari, Stagno di Calich /Alghero; 3 4 ,3 9 ; 29.05.1999. — Sassari,
Porto di Vignola; 1 © (last moult 27.07.1999); 12.06.1999. — Nuoro, lago alto de Flumendosa;
8 4,8 % (last moults of & 11-27.06.1999, of 7 $ 10.6-31.07.1999); 08.06.1999. — Nuoro, Cala
Ginepro /Orosei (under Juncus); 1 4,3 ® (last moults of 13-16.06.1999); 10.06.1999. —
Cagliari, Quartu (laguna); 1 ®; 15.09.1968. — Same; 6 d, 14 © (spiders with strong size
variation); 03.06.1999. — Calabre, Cosenza, Tarsia /Crati valley; 1 9 ; 04.08.1968.

DESCRIPTION: Prosoma tawny-brown, with faint marking. Opisthosoma dor-
sally blackish, ventrally grey, covered with medium-long dark hairs. Legs with
blackish marking up to mid-length of femora, tarsi tawny. Leg spination: Metatarsi I
v000; IT v220. — 4 from Greece (from Spain in parentheses): Total length 4.8 (6.0).
Prosoma: 2.25 (2.8) long, 1.58 (1.94) wide, 0.86 (1.07) wide at level of posterior eyes.
Eye sizes and interdistances: AME 0.08 (0.10), ALE 0.11 (0.14), PME 0.10 (0.14), PLE
0.10 (0.10); AME-AME 0.06 (0.07), AME-ALE 0.01 (0.01), PME-PME 0.02 (0.03),
PME-PLE 0.03 (0.04), ALE-PLE 0.06 (0.07). MOQ length 0.24 (0.31), front width
0.40 (0.48), back width 0.44 (0.53). Clypeus: 0.08 (0.12) from AME, 0.04 (0.07) from
ALE. Width of pedipalpal tibia 2/3 of its length or less (cf. Fig. 28). Retrolateral tibial
apophysis shorter than dorsal length of tibia. Small triangular apical projection on em -
bolar base. Scutum occupying 20% of opisthosoma length. ?: From Greece (from
Spain in parentheses): Total length 5.0 (7.50). Prosoma: 1.96 (2.83) long, 1.43 (2.12)
wide, 0.80 (1.21) wide at level of posterior eyes. Eye sizes and interdistances: AME
0.08 (0.10), ALE 0.11 (0.14), PME 0.11 (0.16), PLE 0.11 (0.13); AME-AME 0.04
(0.05), AME-ALE 0.01 (0.01), PME-PME 001 (0.03), PME-PLE 0.02 (0.03), ALE-
PLE 0.05 (0.07). MOQ length 0.28 (0.35), front width 0.38 (0.50), back width 0.41
(0.57). Clypeus: 0.08 (0.13) from AME, 0.07 (0.07) from ALE. Epigynum and vulva
(Figs 45-49, 72). Anterior epigynal margin narrower (rarely equal to) than width of

NEW ZELOTES SPECIES ss

epigynal folds (Figs 45-49). Posterior vulval pouches small. Epigynal folds reaching
middle length of epigynum in its centre, forming a large triangle separating the oblique
lateral membranous areas.

REMARKS!: Jézéquel's (1962: fig. 30) drawing of the vulva of Z. fuscotestaceus
(presumably the holotype: "tube 1897, Plouharnel, Morbihan") shows the vulva of a Z.
tenuis. Simon (1914, footnote on page 168) himself expressed doubts about the status
of Z. fuscotestaceus in relation to Z. tenuis ( under circumpectus). I have not been able
to see the type specimen of Z. fuscotestaceus.

Zelotes chaniaensis Sp. n. Figs 50-58, 79

Z. tenuis. — Chatzaki et al., 2003: 80, figs 105-107, 111-112 (rare form, misidentification), not
figs 108-110, 113-114 = Z. babunaensis.

HOLOTYPE: GREECE, Crete, Chania, Episkopi, 35°30'N 23°46'E; 4 (with palpus in
microvial, last moult 04.01.2000); 10.10.99.

PARATYPE: Same locality and collecting date as for holotype; 1 ® (with vulva in
microvial, last moult 19.01.2000).

ETYMOLOGY: The species name, an adjective, refers to the city of Chania on the
island of Crete.

DIAGNOSIS: The male differs from that of Z. tenuis by a hook-like distal embo-
lar base tooth (Figs 50-53); the female with shorter median ducts; anterior anchoring
pockets wider than lateral folds and folded cuticle in median sector (Figs 57-58, 79).

DESCRIPTION: Prosoma tawny-brown, with faint marking. Opisthosoma
dorsally blackish, ventrally grey, covered with medium-long dark hairs. Legs with faint
marking,; tarsi tawny. Leg spination: Metatarsi I v210; II v220. — d : Total length 5.50.
Prosoma: 2.20 long, 1.66 wide, 0.90 wide at level of posterior eyes. Eye sizes and
interdistances: AME 0.08, ALE 0.10, PME 0.10, PLE 0.08; AME-AME 0.04, AME-
ALE 001, PME-PME 0.04, PME-PLE 0.03, ALE-PLE 0.04. MOQ length 0.27, front
width 0.40, back width 0.44. Clypeus: 0.11 from AME, 0.06 from ALE. Pedipalp (Figs
50-53): Patella dorsally slightly longer than tibia. Width of pedipalpal tibia 3/4 of its
length or less (cf. Fig. 28). Retrolateral tibial apophysis shorter than dorsal length of
tibia. Scutum occupying 1/4 of opisthosoma length. — © : Total length 5.40. Prosoma:
2.33 long, 1.55 wide, 0.80 wide at level of posterior eyes. Eye sizes and interdistances:
AME 0.07, ALE 0.11, PME 0.08, PLE 0.08; AME-AME 0.06, AME-ALE 0.01, PME-
PME 0.04, PME-PLE 0.028, ALE-PLE 0.05. MORQ length 0.26, front width 0.37, back
width 0.41. Clypeus: 0.07 from AME, 0.06 from ALE. Epigynum (Figs 57-58).

REMARK: Z. chaniaensis seems to be a Cretan endemic replacing Z. fenuis on
that island.

Zelotes manytchensis (Ponomarev & Tsvetkov, 2006) comb. n. Figs 59-64, 77
Trachyzelotes manytchensis Ponomarev & Tsvetkov, 2006: 11, figs 18-19 (description of 4
and ®).

MATERIAL EXAMINED: IRAN, Khohkiluyeh, Dogonbadän, 30°22'N 50°47'E; 1 4,2 9
(with palpus and vulva in microvials); 21.05.1974.

DIAGNOSIS: 4 pedipalp (Figs 59-62): The retrolateral loop of the embolar base,
which reaches far below the level of terminal apophysis, and its membranous retro-

528 A. SENGLET

embolar base hook

FIGS 50-58

Zelotes chaniaensis sp. n. (50-53) Left male palp. (50) Prolateral view. (51) Ventral view. (52)
Retrolateral view. (53) Embolar base, ventral view. (54) Male, ocular group. (55-58) Female.
(55) Ocular group. (56) Left chelicera, ventral view. (57) Epigynum. (58) Vulva, dorsal view.
Bold lines indicate epigynal folds. Scale lines 0.2 mm.

NEW ZELOTES SPECIES 529

(/
2)
CAC
2

é sperm duct

4 Embolar base

ÿ FAITS

Embolar haematodocha
Embolar base
secundary duct
emb. base :
projection Embolar radix
Embolar base ——>\(" se 62

Term. apoph. post. sclerite

PAZ

FIGs 59-64

Zelotes manytchensis. (59-62) Left male palp. (59) Prolateral view. (60) Ventral view. (61)
Retrolateral view. (62) Apex of palpal organ, cleared, dorsal-retrolateral view. (63) Epigynum.
(64) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale lines 0.2 mm.

530 A. SENGLET

lateral projection distinguish Z. manytchensis from the other species of the renuis-
group. 9: Epigynum (Figs 63, 77): lateral folds wide apart, with large convex lateral
pouches.

DESCRIPTION: Prosoma and legs tawny-brown, lighter tarsi and metatarsi.
Opisthosoma greyish brown, covered with short to medium-long hairs. Tarsi I, II
entirely scopulate, metatarsi I, II in apical half. — & : Total length 4.50. Prosoma: 1.80
long, 1.45 wide. Eye sizes and interdistances: AME 0.07, ALE 0.10, PME 0.10, PLE
0.07; AME-AME 0.05, AME-ALE 0.01, PME-PME 001, PME-PLE 0.02, ALE-PLE
0.02. MOQ length 0.20, front width 0.35, back width 0.38. Clypeus: 0.07 from AME;
0.04 from ALE. Pedipalp tawny (Figs 59-62). Tibia dorsally slightly shorter than
patella. Retrolateral apophysis longer than tibia (cf. Fig. 28). Embolar base forming a
large and low retrolateral loop wearing a membranous projection; a secondary duct
(Fig. 62) uniting projection to tip of embolus, as observed in Drassyllus (Miller, 1967:
table 3, figs 3, 5). Scutum occupying 1/3 of opisthosoma length. Leg spination:
Metatarsus I v000. — 9: Total length 6.20. Prosoma: 1.84 long, 1.36 wide, 0.74 wide
at level of posterior eyes. Eye sizes and interdistances: AME 0.07, ALE 0.10, PME
0.08, PLE 0.08; AME-AME 0.05, AME-ALE 001, PME-PME 0.02, PME-PLE 0.04,
ALE-PLE 0.04. MOQ length 0.21, front width 0.35, back width 0.38. Epigynum (Figs
63-64, 77) with large convex lateral pouches. Vulva (Fig. 64) with almost straight
median ducts.

REMARK: Ponomarev & Tsvetkov (2006) described long cheliceral setae for this
species. The female resembles a Trachyzelotes but lacks a cheliceral brush. The geni-
talia show a close relationship with the Zelotes tenuis species group.

Zelotes alpujarraensis sp. n. Figs 65-71, 78

HOLOTYPE: SPAIN, Andalucia, Granada, Puerto del Lino, 1200m, 36°48'N 03°18'W; d:
26.05.2002.

PARATYPES: Same locality as for holotype; 1 & ,3 ? (with palpus and vulva in microvial)
(last moults of 2 © 02.06 and 27.07.2002).
ETYMOLOGY: The species name refers to the Alpujarras Mountains.

DESCRIPTION: Large species, prosoma and legs dark brown. Opisthosoma
blackish, covered with black bristles and prostrate copper-coloured hairs giving a shiny
appearance. Leg spination: Tibiae II r111, IV r211; metatarsus II v220. — 4 paratype:
Total length 7.50. Prosoma: 3.33 long, 2.55 wide, 1.50 wide at level of posterior eye.
Eye sizes and interdistances: AME 0.08, ALE 0.13, PME 0.10, PLE 0.10; AME-AME
0.07, AME-ALE 0.01, PME-PME 0.07, PME-PLE 0.08, ALE-PLE 0.08. MOQ length
0.30, front width 0.51, back width 0.59. Clypeus: 0.10 from AME, 0.07 from ALE.
Pedipalp (Figs 65-69): Tibia 1.5x longer than wide. Retrolateral tibial apophysis
shorter than tibia. Patella dorsally longer than tibia. Embolar base developed as a large
prolateral bow, with a curled retrolateral projection and a distad-directed embolus.
Large, more or less circular terminal apophysis bearing a retrolateral ridge. Scutum
occupying 1/4 of opisthosoma length. 9 : Total length 6.60. Prosoma: 3.00 long, 2.36
wide, 1.28 wide at level of posterior eye. Eye sizes and interdistances: AME 0.08, ALE
0.11, PME 0.08, PLE 0.08; AME-AME 0.07, AME-ALE 0.01, PME-PME 0.07, PME-

NEW ZELOTES SPECIES

531

66

embolar radix

+ x
terminal apophysis embolar radix 68
posterior sclerite

etre

+”

FIGS 65-71

Zelotes alpujarraensis sp. n. (65-69) Left male palp. (65) Prolateral view. (66) Ventral view. (67)
Retrolateral view. (68) Apex of palpal organ, cleared, dorsal view. (69) Id., apical view. (70)
Epigynum. (71) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale lines 0.2 mm.

532 A. SENGLET

PLE 0.07, ALE-PLE 0.07. MOQ length 0.26, front width 0.44, back width 0.51.
Clypeus: 0.10 from AME, 0.07 from ALE. Epigynum (Fig. 70). Epigynal plate almost
square, With short posterior pouches. Vulva (Fig. 71) showing sclerotized median ducts
and large spermathecae.

REMARKS!: The genitalia of this large and dark species correspond to those of
the Z. tenuis-group, especially in the male pedipalpal structure. The male posterior
ocular row differs from that of the Z. fenuis-group (Figs 54-55) in having smaller eyes
of equal size separated by 0.7 of their diameter.

Zelotes thorelli-group

DEFINITION: Dark coloured spiders. Long coiled embolus without direct
connection to embolar base. Coil turning left on left palp. Embolar base more or less
fused to terminal apophysis. Strong embolar radix and posterior sclerite of terminal
apophysis at a relatively acute angle (Figs 83, 92, 101).

KEY TO THE SPECIES OF THE Z. THORELLI-GROUP

l Large embolar base projection (Figs 81, 89, 100). Epigynum with
median. DOSteTIOT NOLCR SE thorelli-subgroup, 2
- Short embolar base projection (Figs 108, 115). Epigynum with median
POSITIOr P'OUSION EE EN EN CP CT fulvopilosus-subgroup, 5
2 Short posterior epigynal notch; short convolution of median vulval duct
(Figs 85-86). Slim transverse embolar base projection (Fig. 81)........... 4
- Median posterior epigynal notch occupying 1/3 or more of epigynal
plate; convolution of median vulval duct reaching anterior margin of
epigynum; embolar base projection wide or depressed (Figs 89, 98, 100) ....3
3 Embolar base projection wide, distad-directed. Posterior epigynal notch
hafrow (Figs 89,95)..-1225 SC OR E Z. thorelli
- Embolar base projection narrow, ribbon-like; embolus with two narrow
apophyses (Figs 98, 100-101). Posterior epigynal notch triangular

(Fig 102) 2. 4, RS PI EE Z. laconicus sp. n.
+ Pedipalp widened, with narrow transverse retrolaterad-directed embolar

projection (Figs 81-83). Copulatory duct coiled in numerous narrow

turnsi(Fios 86-87) Mau Nr NE MR Re Z. pediculatoides sp. n.

- Pedipalp elongated, with narrow embolar projection retrolateral-distad
directed (Fig. 96; Di Franco, 1994: figs 5-8)..Copulatory duct coiled in

FN TÜNS 20025 Aie 4e HO NS AS CE TS Z. lagrecai
- Copulatory duct coiled in numerous wide turns (Di Franco, 1994: fig. 9;
Levy,2009:fies 65-66) Male unknown en Z. pediculatus
5 One-loop retrolateral embolar coil almost reaching base of bulbus.
Lateral epigynal pouches elongated, in a posterior position (Figs 106-
107.109) 50e RE Z. flagellans

- One-loop retrolateral embolar coil reaching mid-length of bulbus. Short
lateral pouches extended laterally (Figs 113-114, 116) ....... Z. fulvopilosus

NEW ZELOTES SPECIES 393

FIGS 72-79

Photos of epigyna. (72) Zelotes tenuis. (73) Z. fuscorufus. (74) Z. semirufus. (75) Z. fulvaster.
(76) Z. babunaensis. (77) Z. manytchensis. (78) Z. alpujarraensis. (79) Z. chaniaensis. Scale
lines 0.2 mm.

534 A. SENGLET

Zelotes thorelli-subgroup

SPECIES INCLUDED: Z. thorelli, Z. laconicus Sp. n., Z. pediculatoides sp. n.,
Z. lagrecai and probably Z. pediculatus (male unknown).

Zelotes pediculatoides sp. n. Figs 80-87

HOLOTYPE: SPAIN, Levant / Murcia, Archena, 38°07'N 01°17'W; G (last moult
14.09.2002); 17.05.2002.

PARATYPES: Same locality as for holotype; 1 & (with palpus in microvial), 1 ® (last
moult 20.09.2002). — Spain, Albacete, Hoya Gonzalo (under stones, evergreen oak), 38°55'N
01°34'W; 3 9; 16.05.2002. — Spain, Estremadura, Caceres, Jarandilla (rio Tietar in litter on
sand), 400 L'N 05°37'W; 1 9; 15.06.2009 — Spain, Andalucia, Sevilla, Rio Viar /Castilblanco,
37°43'N 05°53'W; 2 9 ; 02.06.2002.

OTHER MATERIAL EXAMINED: SPAIN, Valencia, La Albufera, 39°21'N 00°19'W: 1 ©;
16.06.1971. — Castellon, Villanueva de Alcolea, 40°14'N 00°03'E; 1 @ ; 07.09.1971. — Alicante,
Elda, 38°30'N 00°47'W; 3 9 (with vulva in microvial); 19.06.1971. — Alicante, Bullas, 38°02'N
01°39'W; 2 9; 04.07.1971. — Same locality (under vegetation); 1 $; 18.05.2002. — Nueva
Castilla /La Mancha, Madrid, Navalcarnero, 40°18'N 03°56'W; 1 9: 12.06.1969. — Madrid,
Aldea del Fresno (rio Alberche), 40°19'N 04°13'W; 1 9; 13.06.1969.

ETYMOLOGY: The species name reflects the close relationship with Z. pedicu-

latus Marinaro.

DESCRIPTION: Large dark species. Prosoma dark brown, with black bristles.
Opisthosoma black, covered with greyish adpressed hairs and black bristles. Legs dark
brown; tarsi feebly lightened. Tarsi and metatarsi I, IT entirely scopulate. Posterior eye
row Straight to slightly recurved. — 4 paratype: Total length 6.46. Prosoma 3.00 long,
2.30 wide, 1.08 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.07,
ALE 0.14, PME 0.10, PLE 0.10; AME-AME 0.07, AME-ALE 001, PME-PME 005,
PME-PLE 0.07, ALE-PLE 0.09. MOQ length 0.28, front width 0.43, back width 0.52.
Clypeus: 0.13 from AME, 0.08 from ALE. Pedipalp (Figs 80-84): Patella dorsally
longer than tibia. Tibia wider than long. Retrolateral tibial apophysis twice as long as
tibia. Terminal apophysis partly fused to embolar base at level of apical projection (Fig.
83). Basal segment of embolar base narrow (? intercalary sclerite); its apical segment
with inflated membranous wall. Embolar radix embedded in a large posterior embolar
haematodocha, articulated on the posterior tegular base over the terminal apophysis
posterior sclerite. Embolar base with long narrow dorsally furrowed apical projection
guiding the slim embolus. Coiled embolus bearing a membranous retrolateral flange
for about 1/3 of its length; hair-thin tip of embolus resting in a cymbial furrow. Scutum
occupying 40% of opisthosoma length. — 9 : Total length 8.00. Prosoma 3.10 long, 2.35
wide, 1.26 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.09,
ALE 0.12, PME 0.11, PLE 0.11; AME-AME 0.07, AME-ALE 0.01, PME-PME 0.05,
PME-PLE 0.07, ALE-PLE 0.09. MOQ length 0.30, front width 0.46, back width 0.57.
Clypeus: 0.13 from AME, 0.07 from ALE. Epigynum and vulva (Figs 85-87): Vulval
coils often visible through the cuticle in a transverse to longitudinal position.

Zelotes thorelli Simon, 1914 Figs 88-95

Zelotes thorelli Simon, 1914: 163, 172, 214, figs 320-321, 357 (description of 4 and 9). —
Jézéquel, 1962: 525, fig. 5 (description of ®).
MATERIAL EXAMINED: SPAIN, Catalonia, Gerona, Port de la Selva-Qadaquès (under
Cistus), 42°18'N 03°13'E; 1 4,3 © (last moult of 4 26.09.2004); 23.05.2004. — Gerona, north

NEW ZELOTES SPECIES 535

embo

83

post. tegular base

——.
PCR F
+
LE, Fr À
AL
« pe
*
*“
LE]
-
-
Ten mmnn 6

FiGs 80-87

Zelotes pediculatoides sp. n.(80-84) Left male palp. (80) Prolateral view. (81) Ventral view. (82)
Retrolateral view. (83) Apex of palpal organ, cleared, retrolateral view. (84) Id., dorsal view. (85)
Epigynum. (86) Vulva, ventral view. (87) Id., dorsal view. Bold lines indicate epigynal folds.
Scale lines 0.2 mm.

326 A. SENGLET

of Portligat (under rocs & Cistus), 42°18'N 03°17'E; 2 9 ; 24.05.2004. — Gerona, Cap de Creus
(under rosemary, Cistus, Erica), 42°19'N 03°17'E; 3 ; 24.05.2004. — Castilla / Leon, Valladolid,
Peñafiel, 41°35'N 04°08'W; 1 & (last moult 23.08.2002); 23.06.2002. — Avila, Arenas San
Pedro; 1 & (with palpus in microvial); 19.09.1967. - Nueva Castilla / La Mancha, Albacete, La
Gineta (rio Jucar, fine leaf litter), 39°10'N 01°58'W; 1 4,1 $ (last moult of 4 11.09.2002);
16.05.2002. — Estremadura, Caceres, Montänchez, 39°14'N 06°08'W; 1 ®%; 10.06.2002. —
Badajoz, Venta del Culebrin /Monesterio, 38°02'N 06°13'W; 1 9; 04.06.2002. — Andalucia,
Almeria, Puerto Maria, 37°42'N 02°10'W; 1 4,2 @ (last moults of $ 09.09.2002, of 1 ©
04.10.2002); 19.05.2002. — Granada, Puebla de Don Fadrique (Pinus + rosemary litter), 38°00'N
02°27'W; 1 ® (last moult 09.09.2002); 19.05.2002. - Granada, La Vidriera / Pinar pass, 38°03'N
02°34'W; 1 4,3 © (last moult of & 18.07.2002); 20.05.2002. — Granada, La Calahora (in pine
forest), 37° 10'N 03°03'W; 3 9; 23.05.2002. - Granada, Road Puerto de la Ragua (in pine forest),
37°09'N 03°03'W; 1 G,1 % (last moult of ® 03.09.2002); 24.05.2002. — Granada, above
Capileira (1850m), 36°58'N 03°20'W; 1 9 ; 27.05.2002. — Granada, Collado del Muerto (Sierra
Nevada, 1450m), 37°08'N 03°28'W; 1 9%; 29.05.2002. — Jaen, Sierra de Cazorla (Fuente del
Oso), 37°55'N 02°56'W; 2 4,2 @ (last moult of 1 & 17.09.2002); 21.05.2002. — Jaen, Sierra de
Cazorla (Linarejas), 37°55'N 02°55'W; 2 9 ; 22.05.2002. — Jaen, Tiscar pass, 37°48'N 03°03'W;
3 9; 23.05.2002. — Jaen, Cañada de las Hazadillas (under Pinus), 37°39'N 03°43'W: 1 4,3 9%
(last moults of & 25.09.2002, of 1 $ 02.10.2002); 30.05.2002. — Jaen, Puente de la Sierra,
37°41L'N 03°46'W; 1 9; 31.05.2002. — Cordoba, Breña dam (in evergreen oak litter), 37°51'N
05°04'W; 1 à (last moult 21.09.2002); 01.06.2002. — Cordoba, Villaviciosa road to Espiel); 1 9;
29.06.1969. — Sevilla, Alanis; 1 9 : 01.07.1969. — Huelva, Linares de la Sierra /Aracena, 37° 54'N
06°37'W; 1 9; 05.07.1969. — Same; 2 9; 05.06.2002. — Huelva, Alajar /Aracena, 37°52'N
06°41'W; 2 $ (with vulva in microvial); 07.07.1969. — Same; 2 4,2 $ (last moults of & 17-
27.09.2002, of 1 ® 23.10.2002); 07.06.2002. — Huelva, north of La Nava (schist litter), 38°00'N
06°45'W::1 ©: :09:06.2002:="*Huelva Santa Olalla (0 Cala), 37255 'N DIN RREE
04.07.1969. — FRANCE, Provence /Cote D'Azur, Basses-Alpes, La Pourcine /Limans, 43° 59'N
OS°44'E; 2 9; 21.06.2001. — Basses-Alpes, Aubenas-des-Alpes, 43°57'N OS°48'E; 1 9;
21.06.2001. — Languedoc, Gard, Vénéjan; 1 ® (with vulva in microvial); 01.10.1998. — Gard,
Aude, Bedos pass /Mouthoumet, 42°58'N 02°34'E; 1 9; 30.05.2004. — Eastern Pyrenees,
Banyuls sur Mer, 42°28'N O03°O07'E; 1 d (last moult 28.08.2001); 27.06.2001. — Eastern
Pyrenees, Les Abeilles /Banyuls, 42°28'N 03°04'E; 1 4,2 9 (last moult of & 01.10.2001);
28.06.2001. — Eastern Pyrenees, Cerbère (under stones, in Cistus), 42°27'N 03°09'E; 2 4,3 ©
(last moults of & 1-24.09.2004, of 1 8 29.09.2004); 22.05.2004. — Eastern Pyrenees, above
Banyuls sur Mer (350m), 42°28'N 03°08'E; 2 9; 23.05.2004. ANDORRA, St Julià de Loria
(1200m), 42°27'N 01°29'E; 1 d,1 $ (last moult of & 17.09.2001); 08.07.2001. — Same (oak
litter); 1 &,1 © (last moults of & 19.08.2004, of £ 14.09.2004); 25.05.2004.

DESCRIPTION: Prosoma and legs dark brown. Opisthosoma black, with black
adpressed hairs and black bristles. Posterior eye row straight or slightly recurved. — à:
Total length 5.90. Prosoma 2.56 long, 2.00 wide, 0.90 wide at level of posterior eyes.
Eye sizes and interdistances: AME 0.07, ALE 0.11, PME 0.09, PLE 0.09; AME-AME
0.05, AME-ALE 001, PME-PME 0.04, PME-PLE 0.06, ALE-PLE 0.08. MOQ length
0.24, front width 0.37, back width 0.46. Clypeus: 0.11 from AME, 0.08 from ALE.
Pedipalp (Figs 88-93): Patella dorsally slightly longer than tibia. Tibia wider than long.
Retrolateral tibial apophysis longer than tibia. Retrolateral tip of terminal apophysis
ventrad-directed. Median apophysis with a wide hook and a tapering, strongly devel-
oped, retrolaterad-directed apical lobe. Ventrally sclerotized embolar base with a wide
distad-directed projection. Large embolus coiled into one revolution, with a wide in-
flatable membranous flange. Scutum occupying 35% of opisthosoma length.-— 9: Total
length 6.20. Prosoma 2.73 long, 2.00 wide, 0.10 wide at level of posterior eyes. Eye
sizes and interdistances: AME 0.09, ALE 0.10, PME 0.90, PLE 0.09; AME-AME 0.07,
AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.05, ALE-PLE 0.08. MOQ length 0.28,

NEW ZELOTES SPECIES S31

3 “terminal apoph.
LT, post. sclerite
terminal Le res" 92
apophysis

e*

-
CTP ET LLLS

FIGs 88-96

(88-95) Zelotes thorelli. (88-94) Left male palp. (88) Prolateral view. (89) Ventral view. (90)
Retrolateral view. (91) Apex of palpal organ, cleared, apical view. (92) Id., retrolateral view. (93)
Cymbium discarded, retrolateral view. (94) Vulva, dorsal view. (95) Epigyna, two forms. (96)
Zelotes lagrecai, vulva, dorsal view. Bold lines indicate epigynal folds. Scale lines 0.2 mm.

538 A. SENGLET

front width 0.37, back width 0.46. Clypeus: 0.13 from AME, 0.10 from ALE.
Epigynum (Fig. 95). Epigynal notch almost reaching anterior part of median sector.
Vulva (Fig. 94) characterized by coiled anterior part of median ducts reaching past an-
terior epigynal margin.

Zelotes lagrecai Di Franco, 1994 Fig. 96
Zelotes lagrecai Di Franco, 1994: 217, figs 5-8 (description of 4 and 9).

MATERIAL EXAMINED: SPAIN, Andalusia, Almeria, Adra (La Albufera), 36°45'N
02°57'W; 1 $ (with vulva in microvial); 08.07.1971.

REMARK: Di Franco’s (1994: fig. 6) drawing of the male clearly shows that this
species belongs to the Z. fhorelli group. The epigynum of the single female, which I
collected in the proximity of a large pond, corresponds entirely to the original drawing
(Di Franco, 1994: fig. 7), but has the coiled copulatory duct somewhat reduced (Fig.
96). The biotope has since been completely destroyed for intensive cultivation.

Zelotes laconicus sp. n. Figs 97-104

HOLOTYPE: GREECE, Peloponnesus, Laconia, Viglafia /Neapolis (in Juncus litter on
sand), 36°31'N 22°59'E; G ; 04.10.2010.

PARATYPES: Same locality as for holotype ; 1 4, 2 ® (with vulva and palpus in
microvial). — 1 ® , same, 04.10.2004.

ETYMOLOGY: The species name, an adjective, refers to the Greek province of
Laconia.

DESCRIPTION: Large dark species. Prosoma dark brown, with black bristles.
Opisthosoma black, covered with numerous black bristles. Legs dark brown, tarsi
feebly lightened. Tarsi and metatarsi I, II entirely scopulate. Posterior eye row straight
to slightly recurved. One tooth on retromargin of chelicera. — & holotype: Total length
5.00. Prosoma: 2.25 long, 1.70 wide, 0.86 wide at level of posterior eyes. Eye sizes and
interdistances: AME 0.06, ALE 0.10, PME 0.08, PLE 0.08, AME-AME 0.06, AME-
ALE 001, PME-PME 0.04, PME-PLE 0.04, ALE-PLE 0.06. MOQ length 0.21, front
width 0.31,back width 0.37. Clypeus: 0.08 from AME, 0.07 from ALE. Pedipalp (Figs
97-101): Patella dorsally longer than tibia. Tibia slightly wider than long. Retrolateral
übial apophysis about as long as dorsal length of tibia. Intercalary sclerite narrow.
Grooved embolar base with a ribbon-like depressed retrolateral projection. Embolus
coiled; its external part sclerotized, with a wide internal membranous flange reaching
end of terminal projection of embolar base (Figs 100-101); at that point emitting an
inner twisted apophysis linked with a membrane to the threadlike embolus; an elon-
gated and twisted basal embolar apophysis also present. Scutum occupying 40% of
opisthosoma length. — ® paratype: Total length 6.33. Prosoma: 2.56 long, 1.92 wide,
1.06 wide at level of posterior eyes. Eye sizes and interdistances: AME 0.06, ALE 0.11,
PME 0.10, PLE 0.10; AME-AME 0.04, AME-ALE 001, PME-PME 0.04, PME-PLE
0.06, ALE-PLE 0.07. MORQ length 0.27, front width 0.38, back width 0.46. Clypeus:
0.90 from AME, 0.70 from ALE. Epigynum (Fig. 102) wider than long. Sclerotized
epigynal plate with a triangular posterior median notch reaching almost mid-length of
plate. Vulva (Figs 103-104) characterized by uncoiled bag-like anterior part of median
ducts reaching anterior epigynal margin.

NEW ZELOTES SPECIES 539

embol &- apical and
LP ee RE basal embolar

apophyses

embolar base
projection

grooved Fa
embolar base /::

embolar radix

100

terminal apoph! 7°

spermophor
post. sclerite ; F or Y

ff

FIGs 97-104

Zelotes laconicus sp. n. (97-101) Left male palp. (97) Prolateral view. (98) Ventral view. (99)
Retrolateral view. (100) Apex of palpal organ, apical view, cymbium discarded. (101) Id.,
cleared, retrolateral view. (102) Epigynum. (103) Vulva, ventral view. (104) Id., dorsal view.
Bold lines indicate epigynal folds. Scale lines 0.2 mm.

540 A. SENGLET

Z. fulvopilosus-subgroup
SPECIES INCLUDED: Z. flagellans, Z. fulvopilosus.

Zelotes flagellans (L. Koch, 1882) Figs 105-111

Prosthesima flagellans L. Koch, 1882: 635, pl. 20, fig. 14 (description of & ).
Zelotes fulvopilosus.-— Machado, 1949: 15, fig. 10 (misidentification).

MATERIAL EXAMINED: SPAIN, Andalusia, Granada, La Calahora (in pine forest), 37° 10'N
03°03'W;2 4,2 9 (with palpus and vulva in microvial); 23.05.2002.

DESCRIPTION: Prosoma dark brown. Legs tawny to brown, with darker patella.
Opisthosoma blackish, covered with black bristles and prostrate copper-coloured hairs.
— d: Total length 4.00. Prosoma 1.84 (1.95) long, 1.43 (1.54) wide, 0.73 wide at level
of posterior eyes. Eye sizes and interdistances: AME 0.05, ALE 0.08, PME 0.07, PLE
0.07; AME-AME 0.04, AME-ALE 0.01, PME-PME 0.06, PME-PLE 0.04, ALE-PLE
0.05. MOQ length 0.21, front width 0.28, back width 0.34. Clypeus: 0.08 from AME,
0.06 from ALE. Pedipalp (Figs 105-108): Retrolateral tibial apophysis 1.5 times dorsal
length of tibia. Tegulum with a ventral sclerotization reaching level of median apo-
physis. Intercalary sclerite present. Terminal apophysis (Fig. 108) carrying a cone-
shaped dorsal projection concealed by the cymbium, fused with embolar base to a
single grooved sclerite (cf. Fig. 115 showing Z. fulvopilosus). Embolar base carrying a
conical apical projection attached to embolar haematodocha. Median apophysis situ-
ated in apical position between apical projection and retrolateral tip of terminal apo-
physis. Very large, one-turn-coiled embolus reaching below proximal quarter of
tegulum length. Leg spination: Tibiae III r221, IV r211; metatarsi IT, IV v221. Tarsi
and metatarsi I, Il entirely scopulate. Scutum occupying almost 50% of opisthosoma
length. — ?: Total length 5.60. Prosoma 2.60 long, 2.00 wide, 1.00 wide at level of
posterior eyes. Eye sizes and interdistances: AME 0.06, ALE 0.10, PME 0.10, PLE
0.08; AME-AME 0.06, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.06, ALE-PLE
0.07. MOQ length 0.23, front width 0.36, back width 0.47. Clypeus: 0.12 from AME,
0.10 from ALE. Epigynum (Fig. 109). Epigynal plate sclerotized between lateral and
posterior folds. Deep lateral epigynal pouches situated in posterior sector of folds;
anterior epigynal margin as wide as epigynal plate. Vulva (Figs 110-111): Posterior
intromission. Copulatory duct with two revolutions. Median ducts reaching anterior
part of epigynal plate. Leg spination: Femur IV p001, tibiae LIT, IV p211,r211; meta-
tarsi III, IV v221, v221. Tarsi and metatarsi I, II entirely scopulate.

REMARK: I have not seen the type material, but Koch'’s original drawing leaves
no doubt about the identity of the specimens examinated.

Zelotes fulvopilosus (Simon, 1878) Figs 112-118

Prosthesima fulvopilosa Simon, 1878: 61, pl. 14, fig. 14 (description of & and 9).
Zelotes fulvopilosus. — Simon, 1914: 178, 214, figs 317-318, 375 (description of 4 and 9).
Jézéquel, 1962: 598, fig. 3.

TYPE MATERIAL: Museum National Histoire Naturelle Paris, collection Simon; 1 à , jar
577, AR1947; 1 9 , jar 577; AR1935 (together with 2 9 of Z. flagelans), no locality given.

OTHER MATERIAL EXAMINED: SPAIN, Catalonia, Gerona, Port de la Selva-Qadaquès
(under stones, Cistus), 42°18'N 03°13'E; 1 9 (last moult 30.06.2004); 23.05.2004. — Gerona, Val

NEW ZELOTES SPECIES 541

embolar base
projection

embolar base
projection

term. apoph.
post. sclerite

._ embolar base
emb. radix projection

115

FiGs 105-115

(105-111) Zelotes flagellans. (105-108) Left male palp. (105) Prolateral view. (106) Ventral
view. (107) Retrolateral view. (108) Apex of palpal organ, cymbium discarded, prolateral view.
(109) Epigynum. (110) Vulva, ventral view. (111) Id., dorsal view. (112-115) Zelotes fulvo -
pilosus, left male palp (specimen in coll. Simon). (112) Prolateral view. (113) Ventral view. (114)
Retrolateral view. (115) Dorsal view, cymbium discarded. Bold lines indicate epigynal folds.
Scale lines 0.2 mm.

542 A. SENGLET

de Blanya (4 km south of Capsacosta pass), 42°14'N 02°23'E; 1 d, 1 $; 25.05.2004. —
Castilla/Leon, Avila, road to parking Gredos/Hoyo del Espino (1470m), 40°18'N 05°12'W; 1 9;
18.06.2009. — Nueva Castilla/La Mancha, Albacete, Hoya Gonzalo (under stones, evergreen
oak), 38°55'N 01°34'W; 2 9%; 16.05.2002. — Andalusia, Almeria, Puerto Maria, 37°42'N
02°10'W; 1 © (last moult 24.05.2002); 19.05.2002. — Granada, La Vidriera / Pinar pass, 38°03'N
02°34'W:;: 2 4,3 © (last moult of 1 © 31.05.2002); 20.05.2002. — Granada, Prado de Zangarrilla
(Sierra Nevada, under stones, pasture, 2000m) 37°07'N 03°26'W; 1 4 ; 29.05.2002. — Jaen, Sierra
de Cazorla (Fuente del Oso), 37°55'N 02°56'W; 4 4,4 $ (with vulva and palpus in microvial,
last moult of 1 © 27.05.2002); 21.05.2002. — Jaen, Sierra de Cazorla (Linarejas), 37°55'N
02°55'W; 1 9 ; 22.05.2002. — Cordoba, Almodévar del Rio (Breña dam), 37° 50'N 05°04'W; 1 9:
01.06.2009. — Sevilla, Alanis, 38°02'N 05°11'W; 1 d ; 01.07.1969. — Huelva, Linares de la Sierra
/Aracena, 37°54'N 06°37'W; 1 9; 05.06.2002. — Huelva, Alajar /Aracena, 37°52'N 06°40'W/;
1 : 07.07.1969. — Same; 2 9 ; 07.06.2002. — Same (cork-oak leaf litter); 3 $ ; 09.06.2009. —
Huelva, Fuenteheridos (deep leaf litter), 37°54'N 06°40'W; 1 9%; 12.06.2009. - FRANCE,
Languedoc, Eastern Pyrenees, above Banyuls sur Mer (350m),42°28'N 03°08'E; 1 © (last moult
07.06.2004); 23.05.2004.

DESCRIPTION: Prosoma dark brown. Legs brown, with lightened tarsi. Opistho-
soma black, covered with black bristles. — 4 from Spain, Fuente del Oso (in paren-
theses 4 from same locality): Total length 5.33. Prosoma 2.35 (2.18) long, 1.86 (1.74)
wide, 0.88 (0.84) wide at level of posterior eyes. Eye sizes and interdistances: AME
0.07, ALE 0.10, PME 0.09, PLE 0.09; AME-AME 0.05, AME-ALE 0.01, PME-PME
0.04, PME-PLE 0.04, ALE-PLE 0.05. MORQ length 0.21, front width 0.34 (0.21), back
width 0.40 (0.31). Clypeus: 0.06 from AME, 0.05 from ALE. Pedipalp (Figs 112-115):
Patella dorsally longer than tibia. Tibia wider than long. Retrolateral tibial apophysis
short, dorsally arched. Tegulum with ventral sclerotization reaching level of median
apophysis. Intercalary sclerite present. In ventral view embolar base projection visible
behind terminal apophysis. Terminal apophysis (Fig. 115) fused with embolar base into
a single grooved sclerite. One-turn-coiled embolus reaching mid-length of tegulum.
Embolus arising from embolar radix not connected to embolar base. Tarsi I, II entirely
scopulate, metatarsi I, IT in apical half. Leg spination: Tibiae IIT, IV p211; metatarsi I,
VI v221. Scutum occupying 35% of opisthosoma length. ® from Spain, Fuente del
Oso (in parentheses $ from same locality): Total length 5.50 (5.25). Prosoma 2.25
(1.94) long, 1.65 (1.48) wide, 0.94 (0.08) wide at level of posterior eyes. Eye sizes and
interdistances: AME 0.07, ALE 0.10 (0.09), PME 0.09 (0.08), PLE 0.09 (0.08); AME-
AME 0.04, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.05 (0.04), ALE-PLE 0.05.
MOAQ length 0.21 (0.18), front width 0.34 (0.30), back width 0.38 (0.37). Clypeus: 0.10
from AME, 0.05 from ALE. Epigynum (Fig. 116): Anterior anchoring pockets small,
close to each other. Lateral epigynal pouches shallow, extended along the folds. Vulva
(Figs 117-118): Median ducts situated in posterior half of epigynal plate. Tarsi and
metatarsi I, II entirely scopulate. Leg spination: Tibiae IT p211,r111, IV p211,r211;
metatarsi III, VI v221.

Zelotes baeticus-group

DEFINITION: Light coloured spiders. Male palp: Retrolateral tibial apophysis
with flattened and widened tip. A short intercalary sclerite present. Elevated and arched
embolar base connected to embolus. Posterior sclerite of terminal apophysis connected
directly to embolus (Fig. 133); embolus linked to embolar base by whitish cuticle along
its prolateral margin (Figs 120, 127, 131, 133). Embolus with a large posterior basal

NEW ZELOTES SPECIES 543

sector (Fig. 122), and a large terminal segment in the shape of a wide tube, the later
open (Fig. 129) or flattened (Figs 128, 132); its variable ventral protrusion corres-
ponding to prolateral projection of median apophysis. Epigynum: Lateral folds modi-
fied into large lateral pockets; median part membranous (Figs 123, 134). Cheliceral
groove: Retromargin with large prolateral conical tooth and two smaller teeth behind;
promarginal teeth (from ental to ectal): Medium, large, medium and five decreasing in
size.
SPECIES INCLUDED: Z. baeticus Sp. n., Z. hispaliensis Sp. n. and

Z. cordubensis Sp. n.

KEY TO THE SPECIES OF THE Z. BAETICUS-GROUP

Il Prolateral apical protrusion of median apophysis of bulbus large and
conspicuous, bent retrolaterally (Figs 131, 133); embolus wide open;
lateral epigynal pockets covering lateral folds ............ Z. baeticus sp. n.

- Prolateral apical protrusion of median apophysis reduced and cone-
pd ii I20m2 7); embolus #lätiened :/. : /. 1. 4... Arte... 2

2 Embolar base with terminal projection; embolus with ventral inden-
honscmale Gnknown 2.4. .1.0-0L......0 Z. cordubensis sp. n.

- Embolar base without terminal projection; embolus without ventral
indentation; lateral epigynal pockets separated from lateral folds . .....
he 1, er. Z. hispaliensis sp. n.

Zelotes hispaliensis sp. n. Figs 119-125

HOLOTYPE: SPAIN, Andalusia, Huelva, Alajar /Aracena, 37°52'N 06°41'W; d;
07.06.2002.

PARATYPES: Same locality as for holotype; 3 4,5 © (last moults of 1 4 09.06.2002, of
2 ® 20.06 and 07.07.2002). — Same; 2 & (with palpus in microvial); 07.07.1969. — Same (cork-
oak leaf litter); 2 $ (last moults 12 and 13.06.2009); 09.06.2009. — Spain, Huelva, Puerto de
Alajar (820m, evergreen oak leaf litter), 37°53'N 06°40'W; 2 4 ; 10.06.2000.

ETYMOLOGY: The species name is the adjective of Hispalis, the roman name of
Sevilla.

DESCRIPTION: Prosoma tawny, adorned with very fine adpressed hairs and
medium-sized black bristles. Opisthosoma covered with black adpressed hairs and
black bristles. Posterior eye row slightly recurved. — 4 paratypes (smaller 4 in paren-
theses): Total length 4.75 (4.50). Prosoma 2.25 (1.90) long, 1.72 (1.47) wide, 0.91
(0.78) wide at level of posterior eyes. Eye sizes and interdistances: AME 0.06, ALE
0.10, PME 0.08 (0.07), PLE 0.08 (0.07); AME-AME 0.06 (0.05), AME-ALE 001,
PME-PME 0.04, PME-PLE 0.06, ALE-PLE 0.06. MOQ length 0.23 (0.19), front width
0.37 (0.33), back width 0.44 (0.43). Clypeus: 0.08 from AME, 0.07 from ALE.
Pedipalp (Figs 119-122): Patella dorsally longer than tibia. Tibia wider than long.
Retrolateral tibial apophysis with widened and truncated tip, provided with a small
tooth. Intercalary sclerite small. Terminal apophysis oblique, in prolateral position; its
posterior sclerite directly connected to embolus, linked to embolar base by transparent
cuticle. Hook of wide triangular median apophysis forming a gutter. Apical protrusion
on median apophysis small, conical (Fig. 120). Embolar base projection absent.

544 A. SENGLET

embolus ventral embolar radix
protrusion

À post. sector
of embolus

122

FIGS 116-125

(116-118) Zelotes fulvopilosus. (116) Epigynum. (117) Vulva, ventral view. (118) Id., dorsal
view. 119-125 Zelotes hispaliensis sp. n. (119-122) Left male palp. (119) Prolateral view. (120)
Ventral view. (121) Retrolateral view. (122) Apex of palpal organ, cymbium discarded, retro-
lateral-apical view. (123) Epigynum. (124) Vulva, ventral view. (125) Id., dorsal view. Bold lines
indicate epigynal folds. Scale lines 0.2 mm.

NEW ZELOTES SPECIES 545

Embolus with a prolateral protrusion (F1g. 122).Leg spination: Metatarsi I, II v220; IT,
IV v221, some large specimens with an additional spine on femur IT pO11. Tarsi and
metatarsi I, II entirely scopulate. Scutum occupying 1/4 of opisthosoma length. — ®
(smaller specimen in parentheses): Total length 5.66. Prosoma 2.90 (1.96) long, 1.90
(1.43) wide, 1.11 (0.78) wide at level of posterior eyes. Eye sizes and interdistances:
AME 007, ALE 0.11, PME 0.07, PLE 0.08; AME-AME 0.07, AME-ALE 001, PME-
PME 0.05, PME-PLE 0.07, ALE-PLE 0.06. MOQ length 0.26, front width 0.46, back
width 0.54. Clypeus: 0.11 from AME, 0.08 from ALE. Epigynum (Fig. 123): Lateral
anchoring pockets not reaching posterior level of lateral folds; darker posterior medi-
an marking due to median vulval ducts. Vulva (Figs 124-125).Leg spination: Femur II
p011, metatarsi III v221,r122; IV r122, some large specimens with an additional spine
on tibia II r111.

REMARKS: Z. hispaliensis Was found in evergreen oak leaf litter and seems to
occur only in a narrow range of biotops.

Zelotes cordubensis sp. n. Figs 126-128

HOLOTYPE: SPAIN, Andalusia, Cordoba, Los Villares /Cordoba (Pinus litter), 37°59'N
04°48'W; G ; 31.05.2009.

PARATYPE: Spain, Cordoba, Breña dam (evergreen oak litter), 37°51'N 05°04'W; 1 G;
01.06.2002.

ETYMOLOGY: The species name, an adjective, 1s derived from Corduba, the
Roman name of Cordoba.

DESCRIPTION: Prosoma light coloured, adorned with very fine adpressed hairs
and medium-sized black bristles. Opisthosoma covered with black adpressed hairs and
black bristles. — 4 holotype (paratype in parentheses): Total length 5.00 (5.50).
Prosoma 2.25 (2.50) long, 1.72 (1.90) wide, 0.87 (1.00) wide at level of posterior eyes.
Eye sizes and interdistances: AME 0.07 (0.07), ALE 0.08 (0.10), PME 0.07 (0.08),
PLE 0.07 (0.08); AME-AME 0.05, AME-ALE 0.01, PME-PME 0.04 (0.06), PME-PLE
0.07, ALE-PLE 0.06 (0.07). MOQ length 0.21, front width 0.40, back width 0.50.
Clypeus: 0.10 from AME, 0.07 from ALE. Posterior eye row slightly recurved.
Pedipalp (Figs 126-128): Patella dorsally longer than tibia, the latter wider than long.
Retrolateral tibial apophysis less long than 1.3 of cymbium and slightly longer than
tibia, flexed backward, flattened and widened at tip. Terminal apophysis oblique, in a
prolateral position, depressed, its margin tapering to the tip; its posterior sclerite
directly connected to embolus, linked to embolar base by whitish cuticle. In ventral
view apical protrusion of median apophysis partially concealed by terminal apophysis
(Fig. 127). Embolar base with rounded apical projection protruding beyond junction
with embolus (Figs 126-128). Embolus flattened and enlarged, with a ventral indenta-
tion. Tarsi I, Il entirely scopulate; metatarsi in apical half. Leg spination: Femur II
p011; metatarsi I, II v220; III, IV v221. Scutum occupying 1/4 of opisthosoma length
or less.

REMARKS: Z. cordubensis sp. n. is closest to Z. hispaliensis Sp. n. due to the
reduced apical protrusion of the median apophysis, and to Z. baeticus Sp. n. due to the
presence of a ventral indentation on the embolus and due to a terminal projection on

546 A. SENGLET

the embolar base. The differences from the two sister species justify the description of
a new species. The specimens examined were collected from evergreen oak and Pinus
leaf litter.

Zelotes baeticus sp. n. Figs 129-136

HOLOTYPE: SPAIN, Estremadura, Badajoz, rio Sillo (Higuera la Real), 38°06'N
06°41'W; d (last moult 12.06.2002); 09.06.2002.

PARATYPES: Same locality as for holotype; 1 4 ,1 © (last moults of 4 27.06.2002, of %
09.06.2002). — 2 d'; same; 14.06.2009. — Spain, Andalusia, Cordoba, road Posada-Villaviciosa
(under stones in Pinus litter), 37°00'N 05°05'W; 1 & (last moult 19.06.2009); 02.06.2009. —
Spain, Huelva, north of La Nava (schist litter), 38°00'N 06°45'W; 3 4,2 % (last moults of 2 4
16.06.2002, of 1 © 11.08.2002); 09.06.2002. — Spain, Huelva, Alajar /Aracena (under stones and
in vegetation litter), 37°53'N 06°40'W; 1 4,2 9 (last moults of 9 20.07 and 03.08.2009);
09.06.2009.

OTHER MATERIAL EXAMINED: SPAIN, Sevilla, Rivera de Benalija (Pintado dam), 38°02'N
05°55'W; 1 © (last moult 24.06.2009); 05.06.2009. — Sevilla, Rio Viar /Castilblanco, 37°42'N
05°53'W,; 1 6 ; 24.06.1969.

ETYMOLOGY: The species name 1s the adjective of Baetis, the Roman name of
the river Guadalquivir.

DESCRIPTION: Prosoma tawny, adorned with very fine adpressed haïirs and
medium-sized black bristles. Opisthosoma covered with black adpressed hairs and
black bristles. Posterior eye row straight or slightly recurved. — & paratype from Rio
S1llo,: Total length 5.70. Prosoma 2.50 long, 1.90 wide, 0.93 wide at level of posterior
eyes. Eye sizes and interdistances: AME 0.07, ALE 0.10, PME 0.07, PLE 0.07; AME-
AME 0.05, AME-ALE 001, PME-PME 0.06, PME-PLE 0.07, ALE-PLE 0.06. MOQ
length 0.24, front width 0.40, back width 0.49. Clypeus: 0.11 from AME, 0.86 from
ALE. Pedipalp (Figs 129-133): Patella dorsally longer than tibia. Tibia wider than long.
Retrolateral tibial apophysis with parallel margins, truncated, provided with a small
tooth. Intercalary sclerite small. Terminal apophysis oblique, in a prolateral position,
its apical part wide, depressed into a shallow groove (Fig. 130); its posterior sclerite
directly connected to embolus, linked to embolar base by whitish cuticle. Wide hook
of median apophysis triangular, forming a gutter. Median apophysis with a large blunt
apical protrusion (Figs 131, 133) corresponding to a rounded ventral indentation in
basal half of terminal sector of embolus (Fig. 132). Embolar base with an apical pro-
jection with a triangular dorsal flap. Tarsi I, Il entirely scopulate, metatarsi I, II in
apical half. Leg spination: Metatarsi I, II v220; IIE, IV v221; some large specimens with
an additional spine on femur IT p011. Scutum occupying 1/5 of opisthosoma length. —
® paratype from La Nava: Total length 6.20. Prosoma 2.00 long, 1.43 wide, 0.83 wide
at level of posterior eyes. Eye sizes and interdistances: AME 0.07, ALE 0.10, PME
0.06, PLE 0.07; AME-AME 0.04, AME-ALE 0.01, PME-PME 0.05, PME-PLE 0.06,
ALE-PLE 0.06. MOQ length 0.21, front width 0.35, back width 0.41. Clypeus: 0.07
from AME, 0.57 from ALE. Epigynum (Fig. 134): Lateral epigynal pockets partially
covering lateral folds, extended over posterior level of folds (Fig. 134). Vulva (Figs
135-136). Tarsi and metatarsi I, II entirely scopulate. Leg spination: Tibia II r111;
metatarsi IT, IV v221; some large specimens with an additional spine on femur II p011.

REMARKS: Z. baeticus seems to occur in a wide range of biotops: Below stones
in Pinus, grass and schist litter.

NEW ZELOTES SPECIES sa7

emb. base
projection

embolus

— mn
ns
SE

emb. radix post. segment
term. apoph. of embolus
post. sclerite 4 RE

PS

emb. base
projection

embolus

{1

133

median apoph.
apical protrusion

FIGS 126-136

(126-128) Zelotes cordubensis sp. n., left male palp. (126) Holotype, prolateral view. (127 Id.,
ventral view. (128) Paratype, cymbium discarded, retrolateral view. (129-136) Zelotes baeticus
sp. n. (129-133) Left male palp. (129) Embolus complex, dorsal view. (130) Prolateral view.
(131) Ventral view. (132) Retrolateral view. (133) Apical view, cymbium removed. (134)
Epigynum. (135) Vulva, dorsal view. (136) Id., ventral view. Bold lines indicate epigynal folds.
Scale lines 0.2 mm.

548 A. SENGLET

Zelotes atrocaeruleus-group

DEFINITION: Terminal apophysis with ventral and dorsal lobe, partly or entirely
fused with the embolar base.
SPECIES INCLUDED: Z. atrocaeruleus (Simon) and Z. /atreillei (Simon).

Zelotes atrocaeruleus (Simon, 1878) Figs 137-145

Prosthesima atrocaerulea Simon, 1878: 73, pl. 14, fig. 16 (description of & and ©).
Zelotes atrocoruleus.-— Müller, 1967: 260, pl. 1, figs 15-18, pl. 5, fig. 7 (description of 4 and ©).
— Grimm, 1985: 238, figs 279, 292-293 (description of 4 and 9).

MATERIAL EXAMINED: SWITZERLAND, Vaud, near Genolier, 46°26'N O06°14'E;:
1 (with vulva in microvial, last moult 09.06.2001); 30.05.2001. GREECE, Macedonia,
Thessaloniki, east of Chortiatis (under stones in pasture), 40°37'N 23°07'E; 2 4 (with palpus in
microvial), 1 ® (last moults 4 1 and 11.07.2005, ® 08.07.2005); 07.09.2004. — Thessaloniki,
Aghios Vassilios, 40°40'N 23°07'E; 2 9%; 14.06.1998. — Kastoria, north of Kastoria (pass),
40°35'N 21°18'E,; 2 9 ; 20.06.1998. — Sterea Hellas, Phthiotidas, Aghios Konstantinos-Agnadi,
38°44'N 22°52'E; 1 d (last moult 08.06.2005); 18.09.2004. — Phthiotidas, West of Malesina,
38°37N 23°13'E: 1 4, 1 9; 2105.1968. — Same: 1 d’, 2° 9° (last moults ° 01061998)
21.05.1998. — Same; 1 d (with palpus in microvial, last moult 08.08.2005); 25.09.2004. —
Phthiotidas, Near Malesina, 38°37'N 23°13'E; 1 9 ; 08.06.2008. — Phthiotidas, east of Theologos
(under stones, Cistus), 38°39'N 23°12'E; 1 9 ; 20.06.2008. — Eubea, Loutra Aidipsou, 38°53'N
22°59'E; 1 d; 29.05.1983. — Peloponnesus, Elis, Andritsaina, 37°30'N 21°53'E; 2 G, 1 9;
29.05.1981. — Laconia, north-west of Monemvasie (under stones), 36°43'N 22°59'E; 1 9 (last
moult 07.06.2005); 03.10.2004). — Argolida, Palaia Epidauros (in Pinus litter), 37°39'N 23°09'E;
1 & (with palpus in microvial, last moult 02.07.2005); 01.10.2004.

REMARKS: The Greek 4 & (Figs 137-143) correspond to illustrations by Grimm
(1985: fig. 279) and Müller (1967: figs 15-18). The ventral lobe of the terminal apo-
physis is relatively narrow in Fig. 138, it is variable in width and sometimes serrated.
The Swiss ® (Figs 142-143) corresponds to illustrations by Grimm (1985: figs
292-293) and Miller (1967: fig. 7). The Greek $ $ differ (Figs 144-145) by longer
epigynum and glandular duct, and by a shorter median duct.

DESCRIPTION: See Grimm (1985: 238, figs 279, 292-293).

Zelotes latreillei (Simon, 1878) Figs 146-147

Prosthesima latreillei Simon, 1878: 62 (description of 4 and ©).

Zelotes latreillei. —- Simon, 1914: 165, 177, 214, figs 330-331, 371. — Miller, 1967: 270, pl. 3,
figs 14-15, pl. 4, fig. 9 (description of & and 9). — Grimm, 1985: 201, figs 2, 7,9, 220-
221,241, 258-259 (description of 4 and ®). — Senglet, 2004: 117, figs 102-103 (9).

MATERIAL EXAMINED: GREECE, Macedonia, Thessaloniki, west of Aghios Vassilios,
40°40'N 23°05'E; 2 G; 08.09.2004. — Same; 1 & (last moult 03.08.2008); 31.05.2008. —
FRANCE, Languedoc, Aude, Carcanet forest (in pasture), 42°41'N 02°08'E; 1 4,6 ® (with
vulva in microvial, last moults 4 24.08.2001, 2£ 30.08.2001); 04.07.2001. — Aude, Carcanet
forest (in pasture edge), 42°40'N 02°09'E; 1 & ,1 $ (last moults 4 15.08.2001, £ 15.09.2001);
04.07.2001. — Eastern Pyrenees, above Les Fourquets (1800m), 42°26'N 02°25'E; 1 9;
30.06.2001. — Eastern Pyrenees, Les Fourquets, 42°26'N 02°23'E; 1 d (last moult 0.08.2001);
30.06.2001 (1600m). — Eastern Pyrenees, Les Angles, /Matemale forest (1700m), 42°33'N
O2°04'E; 2 9; 03.07.2001. - PORTUGAL, Tras-Os-Montes, Vila Real, Cortico /Montalegre
(900m), 41°46'N 07°47'W; 1 9 ; 30.08.1969.

DESCRIPTION: See Grimm (1985: 201, figs 2, 7,9, 220-221, 241, 258-259 ).

NEW ZELOTES SPECIES 549

embolar base

term. apophysis

retrolateral-dorsal lobe
term. apophysis
ventral lobe

term. apophysis
posterior sclerite

term. apophysis
prolateral-dorsal lobe

embolar radix

term. apophysis
prolateral-dorsal
lobe /

term. apoph.
ventral lobe

141

fused emb. base -

median apoph.
terminal apoph.

term. apoph.
dorsal lobe

embolar radix interc. sclerite

147

FIGS 137-147

(137-145) Zelotes atrocaeruleus. (137-141) Left male palp. (137) Prolateral view. (138) Ventral
view. (139) Cleared, retrolateral view. (140) Id., dorsal view. (141) Id., cymbium discarded,
apical view. (142-143) Female from Switzerland. (142) Epigynum. (143) Vulva, dorsal view.
(144-145) Female from Greece. (144) Epigynum. (145) Vulva, dorsal view. (146-147) Zelotes
latreillei, left male palp, cymbium discarded. (146) Prolateral view. (147) Dorsal view. Bold
lines indicate epigynal folds. Scale lines 0.2 mm.

550 A. SENGLET

Zelotes subterraneus-group

DEFINITION: Embolar base a simple transverse sclerite with a retrolateral
projection and a notched link to embolus; posterior sclerite of terminal apophysis
connected to posterior tegular base.

SPECIES INCLUDED: Z. aeneus (Simon), Z. apricorum (L. Koch), Z. clivicola
(L. Koch), Z. cyanescens Simon, Z. egregius Simon, Z. gallicus Simon, Z. pseudo-
apricorum Schenkel, Z. subterraneus (C. L. Koch) and Z. egregioides n. sp. (the only
species of this group treated here).

Zelotes egregioides sp. n. Figs 148-156
Zelotes fuscipes.-—Machado, 1941: 15, figs 10-14 (misidentification). — Jézéquel, 1962: 600, fig.
11 (misidentification). — Miller, 1967: 263, pl. 2, fig. 4, pl. 5, fig. 8 (misidentification).
HOLOTYPE: SPAIN, Castilla / Leon, Avila, Peñanegra pass (under stones), 40°25'N
05°18'W; & (last moult 01.07.2009); 17.06.2009.

PARATYPES: Same locality as for holotype; 1 4,1 © (with palpus and vulva in microvial,
prosoma d partly destroyed, last moults 4 20.07.2009, © 04.08.2009). — Same locality as for
holotype; 3 4,1 © (last moults 4 20.07-11.08.2009, 9 06.08.2009). — Spain, Avila, Parking
Gredos /Hoyo del Espino (1780m, in pasture), 40°16'N 05°14'W; 1 & (last moult 08.07.2009);
18.06.2009. — Spain, Avila, Tornavaca pass (under stones), 40°16'N 05°40'W; 2 &,1 % (last
moults & 02.7 and 07.08.2009, © 24.07.2009); 17.06.2009. — Spain, Badajoz, Rio Sillo (Higuera
la Real), 38°06'N 06°41'W; 1 G,1 ® (with palpus and vulva in microvial, last moults 4
27.09.2009, % 20.10.2009); 14.06.2009.

OTHER MATERIAL EXAMINED: SPAIN, Aragon /Navarra, Saragosse, Ariza, 48°18'N
02°05'W; 3 8 (with vulva in microvial); 24.09.1967. -Salamanca, El Cabaco, 40°32'N 06°08'W:
1 G; 12.08.1971. — Estremadura, Caceres, west of Jarandilla (litter in vegetation), 40°08'N
05°40'W; 1 © (last moult 01.10.2009); 16.06.2009. — Andalusia, Granada, La Vidriera /Pinar
pass, 38°03'N 02°34'W; 1 d (last moult 02.08.2002); 20.05.2002. — Granada, Puerto de la Ragua
(2000m), 37°07'N 03°02'W; 1 & (last moult 01.07.2002); 24.05.2002. —- PORTUGAL, Tras-Os-
Montes, Vila Real, Vilarandelo - S. Lourenço /Chaves, 41°40'N 07°21'W; 1 & 1 & juv.,
29.08.1969. — Vila Real, Cortico /Montalegre, 41°46'N 07°47'W; 1 d; 30.08.1969 (900m). —
Beira Alta, Guarda, Maceira /Fornos de Algodres (Casal do Monte), 40°44'N 07°24'W,1 4,19
(with palpus and vulva in microvial); 09.08.1971.

ETYMOLOGY: The species name indicates a close relationship with Z. egregius
Simon.

DIAGNOSIS: d : Close to Z. aeneus and Z. egregius (see Senglet, 2004: figs
55-57) with respect to terminal apophysis and embolar base; differs from these two
species by the absence of an acute prolateral ventral tooth on the embolar base and by
the absence of a median membranous flap on the embolus. — ©: Differs from that of
Z. egregius (see Senglet, 2004: figs 59-61) by the strongly retrolaterad-directed and
longer glandular duct. Different from Z. murcidus (see Simon, 1914: fig. 374; Jézéquel,
1962: fig. 14, slide PM 53, col. Simon examined), by the wider epigynal plate and the
glandular ducts reaching 40% of the total epigynal length (in Z. murcidus only 30%).

DESCRIPTION: Large dark species. Prosoma dark brown, with black bristles.
Opisthosoma black, covered with greyish adpressed hairs and black bristles. Legs dark
brown, with lighter tarsi. Posterior eye row straight to slightly recurved. — & paratype
from Peñanegra, (holotype in parenthesis): Total length 6.00 (5.53). Prosoma: 2.70
(2.38) long, 2.10 (1.95) wide, 1.08 (0.93) wide at level of posterior eyes. Eye sizes and

NEW ZELOTES SPECIES 551

_ emb.base embolus
R projection

——
—

FIGs 148-156

Zelotes egregioides sp. n. (148-152) Left male palp. (148) Retrolateral view. (149) Ventral view.
(150) Retrolateral view. (151) Embolus complex cleared, dorsal view. (152) Tip of embolus.
(153) Vulva, short median duct type, dorsal-lateral view. (154) Id., dorsal view. (155) Vulva,
long median duct type, dorsal view. (156) Epigynum. Bold lines indicate epigynal folds. Scale
lines 0.2 mm.

592 A. SENGLET

interdistances: AME 0.06, ALE 0.11, PME 0.10, PLE 0.10; AME-AME 0.06, AME-
ALE 001, PME-PME 0.05 (0.04), PME-PLE 0.06 (0.05), ALE-PLE 0.08 (0.07). MOQ
length 0.27 (0.23), front width 0.38 (0.36), back width 0.44 (0.43). Clypeus: 0.14 from
AME, 0.11 from ALE. Pedipalp (Figs 148-152): Patella dorsally longer than tibia.
Tibia dorsally wider than long. Retrolateral tibial apophysis longer than tibia. Scutum
occupying 1/4 of opisthosoma length. Leg spination: Metatarsi I VOOO; IT, IV v221.
Tarsi I, II entirely scopulate, metatarsi in apical 3/4. — ® paratypes from Peñanegra:
Total length 5.53 (5.50). Prosoma: 2.55 (2.25) long, 1.90 (1.73) wide, 1.03 (0.90) wide
at level of posterior eyes. Eye sizes and interdistances: AME 0.05, ALE 0.10, PME
0.08, PLE 0.08; AME-AME 0.07, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.05,
ALE-PLE 0.70. MOQ length 0.25, front width 0.26, back width 0.31. Clypeus: 0.11
from AME, 0.08 from ALE. Epigynum and vulva (Figs 153-156). Anterior extension
of median ducts variable (in Figs 153-154 short, in 155-156 long).Leg spination:
Metatarsi IT, IV v221. Tarsi and metatarsi I, II entirely scopulate.

REMARKS: Adult 4 & occur from July to September, © from July to October.
The 4 examined by Machado (1941: fig. 10) could belong to Zelotes egregius because
of the presence of an acute prolateral tooth on the embolar base.

Zelotes criniger-metellus-group

DEFINITION: This heterogeneous group differs notably from Zelotes s. str. by:
Absence of intercalary sclerite; radix with a dorsal apophysis, radix connected to em-
bolus by a membranous link; embolar base with a prolateral-dorsal apophysis. Due to
the presence of a posterior sclerite on the terminal apophysis, the shape of the ocular
group and the general appearance, I leave this group in the genus Zelotes.

KEY TO THE SPECIES OF THE Z. CRINIGER-METELLUS-GROUP

I Embolus elongated and:coiled\ is 160) Z. criniger
- Embolus short and massive (Figs 165,175).......... Z. metellus-subgroup
Zelotes criniger Denis, 1937 Figs 157-164

Zelotes criniger Denis, 1937: 1036, plate 2, fig. 4 (description of 4). — Di Franco, 1987: 149,
figs 8, 11 (4).

Zelotes denisi Marinaro, 1967: 693, fig. 9 (description of ©), syn. n. — Di Franco, 1987: 152,
figs 12-13 (). — Deltshev er al. 2006: 712, figs 12-13 (9).

MATERIAL EXAMINED: SPAIN, Andalusia, Cordoba, Breña dam, Almodôvar del Rio (evergreen
oak litter), 37°51'N 05°04'W; 4 G, 6 ® (with palpus, and vulva in microvial);
01.06.2002. — Cordoba, Almodôévar del Rio (Breña dam), 37°50'N 05°04'W: 2 4,5 9;
01.06.2009. — Sevilla, east of Cazalla de la Sierra (deep leaf litter), 37°57'N 05°45'W;
1 ® ; 04.06.2009.

DESCRIPTION: Prosoma brown, adorned with fine adpressed hairs. Opisthosoma
black, covered with medium adpressed haïirs. Legs brown, with darker femora.
Posterior median eyes larger than posterior laterals in slightly recurved line. — & from
Breña dam: Total length 4.20. Prosoma: 1.76 long, 1.33 wide, 0.64 wide at level of
posterior eyes. Eye sizes and interdistances: AME 0.71, ALE 0.08, PME 0.10, PLE
0.07; AME-AME 0.04, AME-ALE 0.01, PME-PME 0.02, PME-PLE 0.04, ALE-PLE

NEW ZELOTES SPECIES 533

terminal apoph.
& post. sclerite

161

Embolar base ire :
dorsal apophysis ZZS

PLLLETT LS

-
+
-
LEP EL Le
__
,
“

:
CR
‘

’ L

: 1:

= ;

FIGs 157-164

Zelotes criniger. (157-161) Left male palp. (157) Prolateral view. (158) Ventral view. (159)
Retrolateral view. (160) Cymbium and subtegulum discarded, apical view. (161) Id., cleared,
prolateral view. (162) Epigynum. (163) Vulva, ventral view. (164) Id., dorsal view. Bold lines
indicate epigynal folds. Scale lines 0.2 mm.

554 A. SENGLET

0.04. MOQ length 0.24, front width 0.31, back width 0.37. Clypeus: 0.10 from AME,
0.07 from ALE. Pedipalp (Figs 157-161). Patella dorsally longer than tibia. Tibia dor-
sally wider than long. Retrolateral tibial apophysis curved dorsally, tip rounded and
provided with a small tooth. Ventrad-directed terminal apophysis cone-shaped, its
strong posterior sclerite connected to mid part of radix. Large folded embolar base with
a narrow connection to its dorsal apophysis (remains of an intercalary sclerite ?).
Embolar base with an acute mesal apophysis and expanding into a large retrolateral
lobe (Figs 160-161). Embolar radix fused to dorsal side of lobe of embolar base; fused
sclerite, shaped into a large bowl with ventral convexity, reaching embolus through a
membranous fold. Leg spination: Metatarsi I, IT v220; I r122. Tarsi I, II entirely sco-
pulate. Scutum occupying 40% of opisthosoma length. — ® from Breña dam: Total
length 3.20. Prosoma: 1.80 long, 1.28 wide, 0.67 wide at level of posterior eyes. Eye
sizes and interdistances: AME 0.07, ALE 0.08, PME 0.10, PLE 0.08; AME-AME 0.04,
AME-ALE 001, PME-PME 0.02, PME-PLE 0.03, ALE-PLE 0.03. MORQ length 0.24,
front width 0.36, back width 0.39. Clypeus: 0.08 from AME, 0.04 from ALE.
Epigynum (Fig. 162) with large and sinuous lateral folds extended close to anterior
anchoring pockets. Median epigynal surface membranous. Additional lateral pockets
present. Vulva in dorsal view with large spiralled pouches almost reaching anterior
anchoring pockets (Figs 163-164); sclerotized elements present in the posterior dorsal
part. Leg spination: Metatarsi I V210; II v210; I r122.

Zelotes metellus-subgroup

DEFINITION: Male palp different from that of most Zelotes by a terminal apo-
physis and an embolar base on a common fused base, intercalary sclerite absent and a
dorsal acute apophysis on embolar base and on embolar radix present. Embolar radix
linked to embolus by membranous cuticle. Embolus short and wide. Retrolateral
posterior lamina present on tegulum. Epigynum: Median part membranous; a charac-
teristic horseshoe-like band bordering posterior part of epigynum.

SPECIES INCLUDED: Z. prishutovae Ponomarev & Tsvetkov and Zelotes metel-
lus Roewer.

KEY TO THE SPECIES OF THE Z. METELLUS- SUBGROUP

l d : Posterior apophyses of embolar base and embolar radix sticking out
of bulbus in its dorsal-prolateral part; tip of embolus blunt (Figs
167-169). 9: Anterior sacs of vulva elongatéd, not fused to epigynum
Frs E70) Lire eee MCE EE er EEE Z. prishutovae
- d : Posterior apophyses of embolar base and embolar radix shorter, not
visible on resting pedipalp; tip of embolus tapering (Figs 172, 175). ©:
Anterior sacs of vulva short, fused to cuticle of epigynum (Fig. 177) ..
LR PR, OUR INR ME LR RE Z. metellus

Zelotes prishutovae Ponomarev & Tsvetkov, 2006 Figs 165-170

Zelotes prishutovae Ponomarev & Tsvetkov, 2006: 13, figs 25-26 (description & and ©).
Camillina metellus.-— Chatzaki, Thaler & Mylonas, 2003: 48, figs 3-7 (not figs 8-9 = Z. metellus).

NEW ZELOTES SPECIES 533

embolus emb. base
post. apophysis

emb. radix

emb. radix
post. apophysis

embolar
haematodocha

embolar
radix

embolar radix

Ÿ RE Î
term. apoph. post. sclerite he TA RÈE
terminal apop ysis\ a.

emb. base ë emb. base
post. apophysis £ post. apophysis
165 NES 166
X se

FIGS 165-170

Zelotes prishutovae. (165-169) Left male palp. (165) Cymbium and subtegulum discarded,
prolateral view. (166) Id., cleared dorsal view. (167) Prolateral view. (168) Ventral view. (169)
Retrolateral view. (170) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale lines
0.2 mm.

MATERIAL EXAMINED: GREECE, Peloponnesus, Laconia, south-west of Monemvasia,
36°40'N 23°01'E; 2 G; 11.05.1998. — Crete Lassithi 2 km west of Istro; 1 ©; 02.06.1997 (leg.
Chatzaki, MHNG).

DESCRIPTION: Prosoma dark brown. Opisthosoma black, covered with medium-
long adpressed hairs. Legs brown, with lighter tarsi and metatarsi. Posterior eyes small,
in a Straight row. — d : Total length 3.33. Prosoma: 1.43 long, 1.17 wide, 0.57 wide at
level of posterior eyes. Eye sizes and interdistances: AME 0.04, ALE 0.07, PME 0.06,
PLE 0.06; AME-AME 0.28, AME-ALE 0.01, PME-PME 0.03, PME-PLE 0.03, ALE-
PLE 0.04. MOQ length 0.14, front width 0.23, back width 0.29. Clypeus: 0.06 from
AME, 0.04 from ALE. Pedipalp (Figs 165-169): Patella dorsally longer than tibia.

556 A. SENGLET

Tarsus width same as dorsal length. Tip of embolus wide. Apophysis of embolar radix
elongated. Tarsi I, Il entirely scopulate, metatarsi I, Il in apical third. Leg spination:
Metatarsi Ï VO00; III, IV v221. Scutum occupying 1/5 of opisthosoma length. — $:
Vulva (Fig. 170): Inflated and elongated glandular ducts with scattered pores.

Zelotes metellus Roewer, 1928 Figs 171-178

Zelotes metellus Roewer 1928: 110, pl. 1, figs 16-17 (description of ®).

Camillina metellus. — Chatzaki, Thaler & Mylonas, 2003: 48, figs 8-9 (not d in figs 3-7 =
Z. prishutovae).

Zelotes sumchi Levy, 1998: 151, figs 128-130 (description of 4) syn. n. — Levy, 2009: 41, figs
87-88 (description of ©).

MATERIAL EXAMINED: IRAN, Fars, Firouzäbäd, 28°52'N 52°32'E,; 1 d ,1 %, 1 immature
G ; 06.06.1974. — Kavär, 29°12'N 52°37'E; 1 d ; 05.06.1974.

DESCRIPTION: Prosoma brown. Opisthosoma black, covered with medium-long
adpressed hairs. Legs brown, with lighter tarsi and metatarsi. Posterior eyes small, in a
straight row. — & from Firouzäbäd (in parentheses from Kavar): Total length 4.75
(3.80). Prosoma: 1.85 (1.60) long, 1.45 (1.23) wide, 0.77 (0.59) wide at level of
posterior eyes. Eye sizes and interdistances: AME 0.06 (0.05), ALE 0.10 (0.09), PME
0.08 (0.07), PLE 0.08 (0.07); AME-AME 0.05 (0.04), AME-ALE 001, PME-PME
0.04 (0.03), PME-PLE 0.03, ALE-PLE 0.04 (0.03). MOQ length 0.20, front width
0.34, back width 0.40. Clypeus: 0.17 from AME, 0.08 from ALE. Pedipalp (Figs 171-
176): Patella dorsally longer than tibia. Tarsus width same as dorsal length. Embolus
wide, with tapering tip. Tarsi I, IT entirely scopulate, metatarsi I, IT in apical two thirds.
Leg spination: Tibia HI r111; metatarsus I v000. Scutum occupying 1/5 of opisthosoma
length. — ©: Total length 5.70. Prosoma 2.20 long, 1.66 wide; 0.87 wide at level of
posterior eyes. Eye sizes and interdistances: AME 0.06, ALE 0.11, PME 0.08, PLE
0.10; AME-AME 0.06, AME-ALE 0.01, PME-PME 0.04, PME-PLE 0.06, ALE-PLE
0.04. MOQ length 0.25, front width 0.38, back width 0.45. Clypeus: 0.11 from AME,
0.08 from ALE. Vulva (Figs 177-178): Lateral anterior vulval sacs fused to epigynal
cuticle, visible on epigynum. Posterior eye row slightly recurved. Tarsi and metatarsi
I, IT scopulate.

REMARKS: Chatzaki ef al. (2003: 48) transferred this species to Camillina on the
basis of a bifid terminal apophysis on the male palp and did not take into account the
very distinct ocular pattern of Camillina. The ocular pattern of Z. metellus is clearly of
the Zelotes type, its terminal apophysis 1s not bifid.

NEW ZELOTES SPECIES

embolar base

radix apoph.

term. apoph.
post. sclerite

FiGs 171-178

557

radix apoph.
ch <embolus

rt
post.

Zelotes metellus. (171-176) Left male palp. (171) Prolateral view. (172) Ventral view. (173)
Retrolateral view. (174) Cymbium discarded, dorsal view. (175) Id., apical view. (176) Id.,
prolateral view. (177) Vulva, ventral view. (178) Id., dorsal view. Bold lines indicate epigynal

folds. Scale lines 0.2 mm.

558 A. SENGLET

ACKNOWLEDGMENTS

Many thanks to Dr Christine Rollard of the National Museum of Natural
History in Paris, and to Mrs Janet Beccaloni of the Natural History Museum in London
for making type material available; to Jürg Wunderlich (Hirschberg) for commenting
the manuscript; to Dr Peter. J. Schwendinger for his constant support and his precious
help in editing this paper, and to Dr André Piuz (both of the Geneva Museum of
Natural History) for producing the Scanning Electron Microscope pictures; to the
Geneva Museum of Natural History for covering the charges of these pictures.

REFERENCES

CHATZAKI, M. THALER, K. & MYLONAS, M. 2003. Ground spiders (Gnaphosidae; Araneae) from
Crete and adjacent areas of Greece. Taxonomy and distribution. III. Zelotes and allied
genera. Revue suisse de Zoologie 110(1) 45-89.

CHYZER, C. & KULCZYNSKI, W. 1897. Araneae hungariae, 2. Editio Academiae Scientiarum
Hungaricae, Budapest, pp. 151-366.

DELTSHEV, C. 2003. A critical review of the spider species (Araneae) described by P. Drensky in
the period 1915-1942 from the Balkans. Berichte des Naturwissenschaftlich-
Medizinischen Vereins in Innsbruck 90: 135-150.

DELTSHEV, C. 2004. Spiders (Araneae) from Sandanski-Petrich Valley (SW Bulgaria).
Mitteilungen aus dem Museum für Naturkunde in Berlin, Zool. Reihe 80: 71-76.
DELTSHEV, C. & BLAGOEV, G. 2001. A critical check list of Bulgarian spiders (Araneae). Bulletin

of the British Arachnological Society 12: 110-138.

DELTSHEV, C. BOSMANS, R. DE SPIEGELAERE, W. & PROVOOST, L. 2006. Zelotes balcanicus
sp. n. a new and widespread species from the Balkan Peninsula (Araneae, Gnaphosidae).
Revue suisse de Zoologie 113: 711-716.

DENIS, J. 1937. On a collection of spiders from Algeria. Proceedings of the Zoological Society
of London 1936: 1027-1060.

DENIS, J. 1952. Notes d'aranéologie marocaine. I. Les Zelotes du Maroc. Revue francaise
d’Entomologie 19: 113-126.

Di FRANCO, F. 1987. Gnaphosidae (Arachnida, Araneae) dell'Isola di Salina (Isole Eolie).
Animalia 13: 137-157.

Di FRANCO, F. 1994. Nuovi dati sulle specie di Zelotes del Maghreb (Araneae, Gnaphosidae):
Considerazioni morfologiche e descrizione di Zelotes lagrecai n. Sp. Animalia 19:
213-225.

DRENSKY, P. 1929. Paiatzi (Aranea) ot tzentralna 1 iougo-zapadna Makedonia. Spisanie na
Bälgarska akademija na naukitë, Sofia. 39: 1-76.

GRIMM, U. 1985. Die Gnaphosidae Mitteleuropas (Arachnida, Araneae). Abhandlungen des
Naturwissenschaftlichen Vereins Hamburg 26: 1-318.

JÉZÉQUEL, J. F. 1962. Contribution à l'étude des Zelotes femelles (Araneidea [sic], Labidognatha,
Gnaphosidae) de la fauna française (2e note). Bulletin du Muséum national d'Histoire
naturelle, Paris 33: 594-610.

KOCH, L. 1866. Die Arachniden-Familie der Drassiden, Hefte 1-6. J. L. Lotzbeck, Nürnberg,
pp. 1-304, pls 1-12.

KOCH, L. 1882. Zoologische Ergebnisse von Exkursionen auf den Balearen. Il: Arachniden und
Myriapoden. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 31:
625-678.

LEVY, G. 1998. The ground-spider genera Setaphis, Trachyzelotes, Zelotes, and Drassyllus
(Araneae: Gnaphosidae) in Israel. /srael Journal of Zoology 44: 93-158.

LEVY, G. 2009. New ground-spider genera and species with annexed checklist of the Gnapho-
sidae (Araneae) of Israel. Zootaxa 2066: 1-49.

NEW ZELOTES SPECIES 559

MACHADO, A. de B. 1941. Araignées nouvelles pour la faune portugaise (Il). Memorias e
Estudos do Museu Zoologico da Universidade de Coimbra 117: 1-xvi, 1-60.

MACHADO, À. de B. 1949. Araignées nouvelles pour la faune portugaise (IT). Memorias e
Estudos do Museu Zoologico da Universidade de Coimbra 191: 1-69.

MARINARO, J. -Y. 1967.Les araignées d'Afrique du Nord. I. Sur une collection de Drassidae à
peigne metatarsal d'Algérie. Bulletin de la Société Zoologique de France 92: 687-704.

MILLER, F. 1967. Studien über die Kopulationsorgane der Spinnengattung Zelotes, Micaria,
Robertus und Dipoena nebst Beschreibung einiger neuen oder unvollkommen bekannten
Spinnenarten. Prirodovedné Prace ustavu Ceaskoslovenske Akademie Ved v Brno (N.S.)
1: 251-298.

PLATNICK, N. I. & SHADAB, M. U. 1983. A revision of the American spiders of the genus Zelotes
(Araneae, Gnaphosidae). Bulletin of the American Museum of Natural History 174:
97-192.

PLATNICK, N. I. 2011. The World Spider Catalog, Version 11.5. American Museum of Natural
History, online at: http://research. amnh. org/entomology/spiders/catalog. html.
PONOMAREV, À. V. & TSVETKOV, A. S. 2006. New and rare spiders of family Gnaphosidae
(Aranei) from a southeast of Europe. Caucasian entomological Bulletin 2: 5-13.
ROEWER, C. F. 1928. Zoologische Streifzüge in Attika, Morea, und besonders auf der Insel Kreta.
XI. Araneae. Abhandlungen herausgegeben vom Naturwissenschaftlichen Verein zu

Bremen 27: 92-123, pls 1-2.

SENGLET, À. 2004. Copulatory mechanisms in Zelotes, Drassyllus and Trachyzelotes (Araneae,
Gnaphosidae), with additional faunistic and taxonomic data on species from southwest
Europe. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 77: 87-119.

SIMON, E. 1878. Les arachnides de France, 4. Roret, Paris, 334 pp.

SIMON, E. 1914. Les arachnides de France, 6, 1re partie: Synopsis générale et catalogue des
espèces françaises de l'ordre des Araneae. Roret, Paris, 308 pp.

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REVUE SUISSE DE ZOOLOGIE 118 (3): 561-598; septembre 2011

Linyphiid spiders (Araneae, Linyphiidae) from Pakistan and India

Andrei V. TANASEVITCH
Centre for Forest Ecology and Production, Russian Academy of Sciences,
Profsoyuznaya Str. 84/32, Moscow 117997, Russia. E-mail: and-tan@mail.ru

Linyphiid spiders (Araneae, Linyphiidae) from Pakistan and India. - 37
linyphiid species are recorded from Pakistan and India, 15 of which are
described as new: Acartauchenius himalayensis Sp. n., Agyneta pakistanica
sp. n., Anguliphantes nepalensoides Sp. n., Gongylidioides keralaensis
sp. n., G. pectinatus Sp. n., Halorates concavus Sp. n., Indophantes tonglu
Sp. n., Pelecopsis indus Sp. n., Tapinocyboides bengalensis sp. n., Tchat-
kalophantes baltistan Sp. n., Tiso incisus Sp. n., T. (?) indianus Sp. n.,
Walckenaeria saetigera Sp. n. À new genus, Paracymboides gen. n., 1s
erected for Paracymboides tibialis sp. n. (the type species) and P. aduncus
sp. n. One new synonym is established: Walckenaeria nepalensis
Wunderlich, 1972 syn. n. = W. martensi Wunderlich, 1972. A distribution
pattern is indicated for many species. Seven species, 1.e., Agyneta nigripes
(Simon, 1884), Archaraeoncus prospiciens (Thorell, 1875), Ceratinella
wideri (Thorell, 1871), Maso sundevalli (Westring, 1851), Microbathy -
phantes palmarius (Marples, 1955), Porrhomma pygmaeum (Blackwall,
1834), and Tenuiphantes tenuis (Blackwall, 1852) are recorded from the
Himalayas for the first time.

Keywords: Arachnida - new genus - new species - new record — Himalayas.

INTRODUCTION

The fauna of Pakistan and India is not only of interest in its own right but also
because both lie in two zoogeographical regions: the northern, mountain territories, be-
long to the Palaearctic region, but the main territories lie in the Oriental region. AÏI 29
linyphiid species known from Pakistan were described or recorded from the Palaearctic
part, the Karakorum mountain system (Caporiacco, 1934, 1935). The linyphid fauna
of India amounts to 38 species; 20 of them known from the Palaearctic part, mainly
from Kashmir (Thaler, 1987, Tanasevitch, 1987), another 20 species are known from
the Oriental part, mainly from Sri Lanka (see Simon, 1894, Helsdingen 1985, etc.);
among them only two species, Indophantes bengalensis Saaristo & Tanasevitch, 2003
and Neriene birmanica (Thorell, 1887), are common to the Palaearctic and Oriental
parts of the county.

The material on which this paper is based was collected mainly in the
Himalayas of Pakistan and India; a small material comes from northern localities in
West Bengal (at the foot of the Himalayas), and from the southern part of India, Kerala
and Tamil Nadu.

Manuscript accepted 24.06.2011

562 A. V. TANASEVITCH

MATERIAL AND METHODS

This paper deals with linyphiids collected in Pakistan and India and kept at the
Muséum d'histoire naturelle, Geneva, Switzerland (MHNG). À small additional
material was also available from the Museo Civico di Storia Naturale di Verona, Italy.
Unless otherwise stated, all material is deposited in the MHNG. Some paratypes and
non-type specimens are in the collection of the Zoological Museum of the Moscow
State University, Moscow, Russia. Sample numbers are given in square brackets.

The chaetotaxy of Erigoninae is given in a formula (e.g., 2.2.1.1) which refers
to the number of dorsal spines on tibiae I-IV. In Micronetinae, the chaetotaxy is given
in a different formula, e.g., Ti I: 2-1-1-2(1), which means that tibia I has two dorsal
spines, one pro-, one retrolateral spine, and two or one ventral spine (the apical spines
are disregarded). The sequence of leg segment measurements 1s as follows: femur +
patella + tibia + metatarsus + tarsus. AIl measurements are given in mm. All scale lines
in the figures correspond to 0.1 mm.

The terminology of genitalic structures in Micronetinae follows that of Saaristo
& Tanasevitch (1996); in Erigoninae it mainly follows that of Tanasevitch (1998) and
Hormiga (2000). The systematic nomenclature follows Platnick (2011), except for the
generic concepts of Agyneta Hull, 1911 and Halorates Hull, 1911.

ABBREVIATIONS

E Convector (according to Tanasevitch, 1998: 423)
CAT Personal collection of Andreï Tanasevitch

D Duct

DE Duct entrance

DSA Distal suprategular apophysis

E Embolus

ED Embolic division

EP Embolus proper

LÉ Lamella characteristica

MED Membrane of embolic division

MHNG Muséum d'histoire naturelle, Geneva, Switzerland
MM Median membrane

MSNV Museo Civico di Storia Naturale di Verona, Italy
MT Membranous tissue

PS Proscape

PMP Posterior median plate

R Radix

S Suprategulum

SMF Senckenberg Museum, Frankfurt a. M., Germany
TA Terminal apophysis

Th Thumb of embolus

ZMMU Zoological Museum of the Moscow State University, Moscow, Russia

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 563

RESULTS

Acartauchenius himalayensis sp. n. Figs 1-8
HOLOTYPE: & (MSNV), PAKISTAN, Northern Areas, Gilgit District, Bagrot Valley,
2500 m a.s.1.; 20.VI.2008; leg. L. Latella.
ETYMOLOGY: The new species is named after the mountain system where it
occurs; Latin adjective.

DIAGNOSIS: À. himalayensis Sp. n. is characterized by the peculiar shape of the
male carapace and palpal tibia.

DESCRIPTION: Male. Total length 1.58. Carapace 0.70 long, 0.55 wide, pale or-
ange-brown, modified as shown in Figs 1-2. Head elevation carrying several strong
curved spines directed backward. Chelicerae 0.25 long, stridulatory fields clear. Legs
pale brown. Leg I 2.08 long (0.60+0.18+0.50+0.42+0.38), IV 2.10 long (0.60+
0.15+0.60+0.45+0.30). Chaetotaxy 2.2.1.1, spines weak, their length about same as
diameter of segment. TmI 0.35. Metatarsus IV without trichobothrium. Palp (Figs
3-8): Tibia with an oblong prolateral outgrowth and a small tooth apically. Para-
cymbium small, hook-shaped. Distal suprategular apophysis developed as a long flat
stripe curved distally. Embolus fusiform, widened distally, with a sharp distinct tooth
before widening. Abdomen 0.95 long, 0.60 wide, dark grey.

Female unknown.

TAXONOMIC REMARKS: The new species is similar to À. asiaticus (Tanasevitch,
1989) and A. monoceros (Tanasevitch, 1989) known from Turkmenistan and
Uzbekistan, respectively (Tanasevitch, 1989), but it differs clearly by the the peculiar
shape of the male carapace and palpal tibia.

DISTRIBUTION: Known from the type locality only.

Agyneta nigripes (Simon, 1884) Figs 9-15
MATERIAL: 2 6 ,4 $ (MSNV), PAKISTAN, Northern Areas, Gilgit District, Passu, Passu
Glacier (in the glacier), 2635 m a.s.1.; 30.X.2008; leg. L. Latella.

COMPARATIVE MATERIAL EXAMINED: Agyneta nigripes, more than 100 4 & 9, AUS-
TRIA, Mt Glockner, 1900-2580 m a.s1.; 1978-1980; leg. K. Thaler. — 2 4,3 © (CAT), RUS-
SIA, Arkhangelsk Area, 30 km W of Tobseda Village, 4 km S of Lake Peschanka-To (68°42°N,
53°10'E); VIL.1984; leg. A. Tanasevitch. — 1 4 (CAT), Murmansk Area, Kola Peninsula,
Dal’niye Zelentsy (69°07°’N, 36°04°E); 4.-26.VII.2009; leg. A. Babenko. — 8 G ,10 ? (ZMMU),
Taymyrskiy Autonomous Region, Efremova River, meadow; 7.VIL.-1.VIIL.2004; leg. D. Osipov.
—2 4,2 9 (ZMMU), Magadan Area, upper reaches of Kolyma River, near Sibit-Tyellakh,
Aborigen Field Station; 4.-14.VI.1982; leg. S. Bukkhalo.

TAXONOMIC REMARKS!: The genitalia of A. nigripes from northern Pakistan are
not significantly different from those of conspecific specimens from other parts of the
distribution range (see Figs 9-15). Small variation in the number, shape and arran-
gement of teeth on the lamella characteristica exist even between the left and right palp
of the same specimen.

REMARKS: This is the southernmost known locality of À. nigripes. The species
is here recorded from the Himalayas and Pakistan for the first time.

RANGE: Holarctic arcto-alpine.

564 A. V. TANASEVITCH

7,8 [

FIGs 1-8
Acartauchenius himalayensis Sp. n., & male holotype. (1-2) Carapace, lateral and dorsal views,
respectively (not to scale). (3-5) Right palp, retrolateral, prolateral and proventral views,

respectively. (6) Palpal tibia, dorsal view. (7) Distal suprategular apophysis and embolus. (8)
Embolus.

Agyneta pakistanica Sp. n. Figs 16-25

HOLOTYPE: à , PAKISTAN, Punjab, environs of Islamabad, ca 550 m a.s.1., Lake Rawal,
on bank near water; 26.1V.1984; leg. S. Vit [PAK-84/25].

PARATYPE: 1 d , from same locality, collected together with the holotype.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 565

ui

er
a ae se v É
anse la dde a) 4

9, 11-15

10

FiGs 9-15

Agyneta nigripes (Simon, 1884), & specimen from Passu Glacier, Pakistan. (9) Right palp, retro-
lateral view. (10) Cymbium , prolateral view. (11) Palpal tibia, dorsal view. (12) Palpal tibia and
paracymbium, retrolateral view. (13) Embolic division. (14-15) Lamella characteristica, different
aspects.

ETYMOLOGY: The specific name is an adjective taken from the type locality.

DIAGNOSIS: The new species is characterized by the peculiar shape of the palpal
tibia and of the lamella characteristica.

DESCRIPTION: Male (holotype). Total length 1.63. Carapace 0.73 long, 0.50
wide, pale brown, with a darker margin. Chelicerae 0.28 long. Legs pale brown. Leg I
2.46 long (0.65+0.18+0.58+0.60+0.45), IV 2.46 long (0.68+0.18+0.59+0.63+0.38).
Chaetotaxy: all tibiae with two dorsal spines. TmI 0.25. Metatarsus IV without

566 A. V. TANASEVITCH

FIiGs 16-25

Agyneta pakistanica sp. n., à paratype. (16) Right palp', retrolateral view. (17) Cymbium,
prolateral view. (18) Palpal tibia, dorsal view. (19-21) Palpal tibia and paracymbium, lateral
view, different aspects. (22) Embolic division. (23) Embolus. (24-25) Lamella characteristica,
different aspects.

trichobothrium. Palp (Figs 16-25): Tibia apically with two pointed teeth: one very
small, the other elongated to a process. Cymbium without any posterior outgrowths.
Lamella characteristica small, with a short L-shaped projection basally. Proximal part
of lamella characteristica relatively wide; distal part very thin, awl-shaped. Embolus
sigmoid, carrying a few very small teeth basally. Abdomen 0.85 long, 0.50 wide, pale

grey.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 567

Female unknown.

DISTRIBUTION: Known from the type locality only.

Anguliphantes nepalensis (Tanasevitch, 1987)

MATERIAL: 1 © , PAKISTAN, Hazara District, Käghän Valley, 1450 m a.s.1., Malkandi
Forest, near foot of rock; 29.VI.1985; leg. S. Vit [PAK-85/2]. — 1 G , INDIA, Uttar Pradesh,
Garhwal, above Pauri, 1900 m a.s.1., dry Quercus forest on N slope, sifting leaf litter and moss;
28.X.1979; leg. I. Lôbl [28]. — 1 ©, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m
a.s.]., near top, sifting litter in forest; 18.X.1978; leg. C. Besuchet & I. Lôbl [19].

COMPARATIVE MATERIAL EXAMINED: 1 d, NEPAL, Goropani Forest between Kali
Gandaki Valley and Pokhara Valley, Punhill near Goropani, 3050-3100 m a.sl., edge of
Rhododendron and Abies forest, sifting litter and mosses; 8.X.1983; leg. I. Lôbl & A. Smetana
[31] (new locality).

VARIABILITY: This species shows some variability in certain genital structures,
such as size of the rounded swelling on the paracymbium, and shape and density of the
fringed margin of the embolus (see Tanasevitch & Saaristo, 2006). Nevertheless the
shape of the distal part of the lamella characteristica 1s consistent enough and is a good,
well-visible character to distinguish this species.

REMARKS: À. nepalensis Was previously known from numerous localities in
Nepal (Tanasevitch, 1987; Tanasevitch & Saaristo, 2006). The species is here recorded
from Pakistan and India for the first time.

RANGE: Himalayan.

Anguliphantes nepalensoides sp. n. Figs 26-31
HOLOTYPE: à , INDIA, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m a.s.1.,
near top, sifting in forest; 18.X.1978; leg. C. Besuchet & I. Lôbl [19].
ETYMOLOGY: The species name, an adjective, points out the similarity of the
new species and À. nepalensis (Tanasevitch, 1987).

DIAGNOSIS: The new species is characterized by the peculiar shape of the
lamella characteristica.

DESCRIPTION: Male. Total length 2.20. Carapace 0.98 long, 0.83 wide, pale
brown, almost yellow, with a greyish margin. Chelicerae 0.43 long. Legs yellow. Legs
or 1ts distal segments mostly lost, Fel & II 1.23 long. Chaetotaxy unclear, but probably
equal to that of À. nepalensis, 1.e. Fe I: O-1-0-0: Ti LE: 2-1-1-0, IT: 2-0-1-0, III-IV:
2-0-0-0; Mt I-IIT: 1-0-0-0. TmIl 0.11. Metatarsus IV without trichobothrium. Palp (Figs
26-31): Tibia flattened distally. Cymbium with a small posterodorsal conical tubercle.
Paracymbium with a rounded swelling in mesal part. Distal part of lamella charac -
teristica fork-shaped, upper branch twice longer than lower one, with a small tooth
between both branches. Upper edge of lamella characteristica carrying a spear-shaped
outgrowth directed backward. Embolus fringed at margin. Abdomen 1.18 long, 0.88
wide, pale grey.

Female unknown.

TAXONOMIC REMARKS: The new species 1s very similar to À. nepalensis, but
can be easily distinguished by the shape of the distal part of the lamella characteristica.
See also above, under Remarks to À. nepalensis.

DISTRIBUTION: Known from the type locality only.

568 A. V. TANASEVITCH

FIiGs 26-31

Anguliphantes nepalensoides sp. n., 8 holotype. (26) Right palp, retrolateral view. (27) Palpal
tibia and paracymbium, dorsal view. (28) Embolic division. (29) Embolus. (30-31) Lamella
characteristica, different aspects.

Archaraeoncus prospiciens (Thorell, 1875)

MATERIAL: 1 ? (MSNV), PAKISTAN, Northern Areas, Gilgit District, Bagrot Valley
(36°01°36.7"N, 74°33°57.6"E), 2700 m a.s.; 18.VI.2008; leg. L. Latella & R. Ahmed. — 1 9
(MSNV ), Gilgit District, Naltar Valley (36°12°34.4°N, 74°08°20.6"E), 3000 m a.s.1.; 1.X1.2008;
leg. L. Latella.

REMARKS: The species is here recorded from the Himalayas and Pakistan for
the first time.

RANGE: Ancient Mediterranean.

Caviphantes pseudosaxetorum Wunderlich, 1979

MATERIAL: 1 ®, PAKISTAN, Hazara District, Käghän Valley, 2000 m a.s.., Ghnwool
Valley, Makhair Forest, litter under Viburnum; 30.VI.1985; leg. S. Vit [PAK-85/4]. — 1 9,
Punjab, environs of Islamabad, Lake Rawal, ca 550 m a.s.l., forest, dry litter; 24.1V.1984; les. S.
Vit [PAK-84/24]. —- 1 ©, N of Lake Rawal, 3.1V.1986, leg. S. Vit. — 1 ? (ZMMU), Ghnwool
Valley, Malkandi Forest, 1600 m a.s.., near water, among stones; 30.VI.1985; leg. S. Vit
[PAK85/7].-— 1 4 ,Swat District, Malam Jabba, 2500-2600 m a.s.1., Abies forest, sifting leaf litter

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 569

and moss; 18.V.1983; leg. C. Besuchet & I. Lôbl [17b]. — 1 © ,S of Saidu Sharif, Murghazar,
sifting leaf litter under Platanus, 1300 m a.s.1.; 8.V.1983; leg. C. Besuchet & I. Lôbl [2b].-— 1 ?,
above Miandam, 2400-2500 m a.s.1., Abies forest, sifting leaf litter and moss; 17.V.1983; leg. C.
Besuchet & I. Lôbl [15b].-1 4 , INDIA, Himachal Pradesh, Kulu Valley, Chijoga (S of Manali),
1900 m a.s.1., from rodent burrow; 14.X.1988; leg. S. Vit [22]. —3 © , Uttar Pradesh, Garhwal,
4 km S of Bhatwari, 1400 m a.sl.; 23.X.1979; leg. I. Lôbl [1979/22]. — 4 4,2 ®, Madras
(= Chennai), Palni Hills, 10 km W of Kodaïikanal, 2300 m a.s.., timberline, degraded forest,
sifting moss under fern; 13.X1.1972; leg. C. Besuchet & I. Lôbl [1972/25a]. — 1 4,3 ©, Palni
Hills, 10 km NW of Kodaïkanal, 2150 m a.s.., edge of Rhododendron forest with fern, sifting
litter near river; 15.X1.1972; leg. C. Besuchet & I. Lôbl [1972/27]. —- 2 4,7 $ (ZMMU),
Kodaikanal, 2100 m a.s.l., forest below town, sifting litter; 11.X1.1972; leg. C. Besuchet & I.
Lôbl [1972/22]. - 2 4,5 ®, 10 km W of Kodaikanal, 2350 m a.s.., degraded forest, under
stones; 13.X1.1972; leg. C. Besuchet & I. Lôbl [1972/25b].- 1 & ,1 © ,23 km W of Kodaïikanal,
Lake Berijam, 2150 m a.s.l., Rhododendron forest, sifting litter; 14.X1.1972; leg. C. Besuchet &
I. Lôbl [1972/26].-— 1 9 , Nilgiri, Ootacamund, 2150 m as... sifting litter under bushes in ravine;
21.X1.1972; leg. C. Besuchet & I. Lôbl [1972/41a].-—1 % , Meghalaya, Khasi Hills, Mawphlang,
1800 m a.s.., forest, sifting litter; 28.X.1978; leg. C. Besuchet & I. Lôbl [1978/32b]. — 5 9,
Kerala, Cardamom Hills, near Munnar, Muttapatti, 1700 m a.s.1, forest with fern, sifting litter;
24.XT1.1972; leg. C. Besuchet & I. Lbl [48].

COMPARATIVE MATERIAL EXAMINED: Caviphantes pseudosaxetorum, SMF 29677, 1 %
paratype, NEPAL, Thakkhola, from Kali Gandaki Valley to main Himalayan Mt. Ridge,
Chadziou Khola Valley near Ghasa, 2330 m a.s.1., 31.X.1969, leg. J. Martens.

DISTRIBUTION: The species was originally described from the Nepal Himalayas
(Wunderlich, 1979). Later, it was found in Japan (Ono ef al., 1991), China (Gao ef al.,
1992), and just recently in Lebanon (Tanasevitch, 2011). The species 1s here recorded
from Pakistan and India for the first time.

RANGE: South Palaearctic-Oriental.

Ceratinella wideri (Thorell, 1871)

MATERIAL: 1 4 , PAKISTAN, Hazara District, Käghän Valley, Naran, Lake Saiful Muluk,
3100 m a.s.l., litter under Saxifraga; 4.-5.VII.1985; leg. S. Vit [PAK-85/20].- 1 d , Swat District,
above Utrot, 2500 m a.s.., Abies & Cedrus forest, moss and leaf litter among fallen trunks;
13.V.1983; leg. C. Besuchet & I. Lôbl [11e]. — 1 9, Malam Jabba, 2500-2600 m a.s.., Abies
forest, sifting leaf litter and moss; 18.V.1983; leg. C. Besuchet & I. LGbl [17b].

REMARKS: The discovery of both sexes of this species in the Pakistan
Himalayas makes its determination reliable. The species 1s here recorded from the
Himalayas and Pakistan for the first time.

RANGE: Palaearctic.

Erigone dentipalpis (Wider, 1834)

MATERIAL: 1 & (MSNV), PAKISTAN, Karakorum, Baltistan, Shalabot, 1700 m as;
15.VIIL.1976; leg. G. Osella. — 2 $ (MSNV), Katzaran, 2200 m asl.; 23.VIL.1976; leg. G.
Osella. — 2 9 (MSNV), Northern Areas, Gilgit District, Ghangche, Kaplu Ghwari; 27.VI.2008;
leg. L. Latella. - 2 $ (MSNV), Skardu District, 2300 m a.s.1.; 27.VI.2008; leg. G. Osella. — 1 4
(MSNV), Skardu, Pakova, 2300 m a.sl.; 1.VIL.1976; leg. G. Osella. — 1 4 (MSNV), Naltar
Valley (36°12°34.4”N, 74°08’20.6"E), 3000 m a.sl.; 1.XI1.2008; leg. L. Latella. — 1 9, Swat
District, Saidu Sharif, 1000 m a.s.]., river bank, under stones and cow-dungs; 11.V.1983; leg. C.
Besuchet & I. Lôbl [1983/8].

REMARKS: In the Himalayas this species was hitherto known only from
Kashmir (Cambridge, 1885) and Karakorum (Caporiacco, 1935).

RANGE: Holarctic.

570 A. V. TANASEVITCH

Erigone prominens Bôüsenberg & Strand, 1906

MATERIAL: 2 G , PAKISTAN, Punjab, environs of Islamabad, Lake Rawal, ca 550 m
a.s.l., on bank near water; 26.1V.1984; les. S. Vit [PAK-84/25].

REMARKS: In the Himalayas the species was hitherto known only from Nepal
(Wunderlich, 1983) and it is here recorded from Pakistan for the first time.

Gongylidiellum confusum Thaler, 1987

TYPE MATERIAL EXAMINED: Gongylidiellum confusum, SMF 33728, 4 © paratypes,
INDIA, Kashmir, Pahalgam, coniferous forest, 2400 m a.s.1.; 14.V.1976; leg. J. Martens. —- SMF
33730,2 G paratypes, Sonamarg, Nichinai-Tal, 3100-3200 m a.s.1.; 9.V1.1976; leg. J. Martens.

OTHER MATERIAL: 4 ® (ZMMU), PAKISTAN, Hazara District, Ghnwool Valley,
Malkandi Forest, 1600 m a.s.1., near water, among stones; 30.VI.1985; leg. S. Vit [PAK-85/7]. —
14,1 ®,Swat District, above Utrot, 2500 m a.s.]., Abies & Cedrus forest, sifting moss and leaf
litter among fallen trunks; 13.V.1983; leg. C. Besuchet & I. Lôbl [11e].— 1 % , same, 2500-2600
m a.s.l., sifting rotten Abies wood; 14.VI.1983; leg. C. Besuchet & I. Lôbl [12c].— 1 © , same,
2600 m a.s.1., wet meadow, sifting mosses & rotten wood; 13.V.1983; leg. C. Besuchet & I. Lôbl
[11d].—1 ©, Chitral, Madaglasht, 2700 m a.s.1., sifting under Salix near river; 26.V.1983; leg.
C. Besuchet & I. Lôbl [27b].-—2 % , Chitral, above Bumburet, valley to Pass Ustui, 2700 m a.s..,
sifting leaf litter under Viburnum; 25.V.1983; leg. C. Besuchet & I. Lôbl [25c]. — 1 ©, INDIA,
Uttar Pradesh, Garhwal, 2 km E of Dhanolti, northern slope, 2250 m a.s.1., near source, sifting
mosses and leaf litter under Rhododendron and Abies; 21.X.1979; les. I. Lôbl [19].-1 4,29
(ZMMU), Garhwal, 10 km E of Dhanolti, 2450 m a.s.I., northern slope, Quercus forest, sifting
litter;, 21.X.1979; leg. I. Lôbl [21b]. — 1 4 (ZMMU), Himachal Pradesh, 10 km W of Simla,
Jutogh, 2000 m a.s.1., leaf litter near foot of rock; 29.X.1988; leg. S. Vit. [88/37].

REMARKS: The species was hitherto known only from Kashmir (Thaler, 1987)
and is here recorded from Pakistan for the first time.

RANGE: Himalayan.

Gongylidiellum nepalense Wunderlich, 1983

TYPE MATERIAL EXAMINED: Gongylidiellum nepalense, SMF 31701,1 & ,2 © paratypes,
NEPAL, Thaksang above Tukche, coniferous forest, 3150-3400 m a.s.1.; 5.-10.VIL.1970; leg. J.
Martens.

OTHER MATERIAL: 8 ? , INDIA, West Bengal, Darjeeling District, Tonglu, 3100 m a.s.l.,
near top, sifting in dwarf forest under brushes on pasture; 16.X.1978; leg. C. Besuchet & I. Lôbl
[16b].-—1 G,6 ©, Tonglu, 3100 m a.s.l., near top, under stones; 16.X.1978; leg. C. Besuchet &
I. Lôbl [16a].- 1 4,2 £ (ZMMU), same, 3100 m a.s.l., near top, sifting in dwarf forest under
brushes on pasture; 16.X.1978; leg. C. Besuchet & I. Lôbl [16b]. — 1 4,2 ? (ZMMU), same,
2700 m a.s.1., forest, sifting near litter: 16.X.1978: leg. C. Besuchet & I. LôbI [17].

REMARKS: The species was originally described from Nepal (Wunderlich,

1983) and is here recorded from India for the first time.

RANGE: Himalayan.

Gongylidioides keralaensis sp. n. Figs 32-38, 114

HOLOTYPE: 4 , INDIA, Kerala, Cardamom Hills, near Munnar, Muttapatti, 1700 m a.s.l,
forest, sifting litter under tree ferns; 24.X1.1972; leg. C. Besuchet & I. Lôbl [48].

PARATYPES: 2 © , from same locality, collected together with the holotype.

ETYMOLOGY: The species name, an adjective, refers to the Indian State where
the new species was found.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 571

FiGs 32-37

Gongylidioides keralaensis sp. n., 4 holotype. (32-33) Right palp, retrolateral and prolateral
views, respectively. (34) Palpal tibia, dorsal view. (35) Distal suprategular apophysis. (36-37)
Embolic division, different aspects.

DIAGNOSIS: The new species 1s characterized by the peculiar shape of the male
palpal tibia, as well as by the shape of the ventral epigynal plate.

DESCRIPTION: Male. Total length 2.35. Carapace 1.13 long, 0.85 wide, pale
brown, unmodified, with blurred grey median stripe and dark margin. Chelicerae 0.50
long, unmodified, stridulatory area distinct. Legs pale brown. Leg I 4.18 long
(1.13+0.30+1.10+1.00+0.65), IV 4.71 long (1.00+1.28+0.93+0.95+0.55). Chaetotaxy
2.2.1.1, spines long and stout. TmI 0.74. AI metatarsi with a trichobothrium. Palp (Figs
32-37): Tibia with a large prolateral process basally bent at a right angle. Cymbium
without posterodorsal outgrowth. Tegulum small, ending with rounded protegulum.
Distal suprategular apophysis long and wide well protruding forward, membranous

5172 A. V. TANASEVITCH

FiGs 38-41

Gongylidioides keralaensis sp. n., ® paratype (38) and G. pectinatus sp. n., & holotype (39-41).
(38) Epigyne, ventral view. (39-41) Carapace, lateral, frontal and dorsal views, respectively. Not
to scale.

distally. Radix very small, surrounded by membranous tissue. Embolus long, curved
gradually. Convector (after Tanasevitch, 1998: 423) complicated in shape, it longest
lobe long, narrow and accompanying the embolus for its protection. Abdomen 1.30
long, 0.88 wide, laterally pale, dorsally grey, with a pale median stripe and several
transversal bands.

Female. Total length 2.65. Carapace 1.00 long, 0.80 wide, unmodified.
Chelicerae 0.50 long, unmodified. Leg I 3.90 long (1.05+0.30+1.00+0.95+0.60), IV
4.01 long (1.13+0.30+1.00+1.03+0.55). TmI 0.80. Abdomen 1.68 long, 1.13 wide.
Epigyne and vulva as shown in Figs 38, 114. Body and leg coloration, as well as
chaetotaxy, as in male.

TAXONOMIC REMARKS: The new species seems to be most similar to G. diellip-
ticus Song & Li, 2008 known from a female from Taiwan (Song & Li, 2008), but
differs clearly by the shape of the ventral epigynal plate.

DISTRIBUTION: Known from the type locality only.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 573

Gongylidioides pectinatus sp. n. Figs 39-46

HOLOTYPE: & , INDIA, Himachal Pradesh, 12 km E of Mandi, 750 m a.s.l., leaf litter:;
25.X.1988; leg. S. Vit [351].

PARATYPE: 1 à , INDIA, Uttar Pradesh, Kumaon, environs of Bhim Tal, eastern slope,
1500 m a.s.l., edge of dry secondary forest; 4.X.1979; lee. I. LGbl [1].

ETYMOLOGY: The specific name is a Latin adjective meaning “with a comb”,
referring to the peculiar shape of the convector.

DIAGNOSIS: The new species is characterized by the peculiar shape of the palpal
tibia, as well as by the presence of a fringed lobe in the embolic division.

DESCRIPTION: Male (holotype). Total length 1.50 (1.68 in paratype). Carapace
0.68 long, 0.53 wide, modified as shown in Figs 39-41, brown with blackish margin.
Eyes relatively large. Chelicerae 0.23 long, unmodified. Legs pale brown. Leg I 2.25
long (0.60+0.18+0.60+0.50+0.37), IV 2.15 long (0.60+0.18+0.55+0.52+0.30).
Chaetotaxy 2.2.1.1, length of spines about 1.5 diameter of segment. TmI 0.56 (0.38 in
paratype). AI metatarsi with a trichobothrium. Palp (Figs 42-46): Tibia conical, elon-
gated, with two small denticles terminally. Paracymbium long and narrow, with several
short spines proximally, and a few long spines distally. Distal suprategular apophysis
very large, flat, distally pointed, well-protruded forward. Radical part of embolic
division very small and membranous, embolus thin, long and curved. Convector rela-
tively large and complicated in shape, with several lobes, one with a comb-like frisge.
Abdomen 0.83 long, 0.55 wide, dark grey, with a pale median stripe.

Female unknown.

TAXONOMIC REMARKS: The new species is well distinguished from other

congeners by the small body size, by the peculiar shape of the palpal tibia, as well as
by the presence a lobe with a comb-like fringe on the convector.

DISTRIBUTION: Known from two localities in northern India.

Gorbothorax aff. ungibbus Tanasevitch, 1998 Figs 47-52

MATERIAL: 1 G , INDIA, Meghalaya, 15 km N of Darugiri, Garo-Hills, 400 m ass.
forest, sifting litter in ravine; 4.X1.1978; leg. C. Besuchet & I. Lôbl [40b].

DESCRIPTION: Male. Total length 1.63. Carapace 0.80 long, 0.63 wide, unmod-
ified, yellow. Chelicerae 0.30 long, unmodified. Legs pale yellow. Leg I 2.92 long
(0.75+0.25+0.75+0.70+0.47), IV 2.95 long (0.80+0.25+0.75+0.75+0.40). Chaetotaxy
and trichobothriotaxy unclear: spines mostly lost, but should be 2.2.1.1, all metatarsi
with a trichobothrium. Palp as shown in Figs 47-52. Abdomen 0.78 long, 0.53 wide,
white with ambiguous pale grey pattern.

REMARKS: The male shows strong similarities to G. ungibbus Tanasevitch,
1998, described from Nepal (Tanasevitch, 1998), but differs by some small details of
palp structure. Each form is known from a single male, so it is impossible to say row
if the differences are due to variability of characters or if these males belong to distinct
but closely related species.

DISTRIBUTION: Known from a single locality in northern India.

574 A. V. TANASEVITCH

FiGs 42-46

Gongylidioides pectinatus sp. n., 4 paratype. (42-43) Right palp, retrolateral and prolateral
views, respectively. (44) Palpal tibia, prolateral view. (45) Palpal tibia and paracymbium, retro-
lateral view. (45) Embolic division.

Halorates concavus sp. n. Figs 53-57

HOLOTYPE: 4 , PAKISTAN, Swat District, above Utrot, 2600 m a.s.., wet clearing in
Abies & Cedrus forest, sifting mosses & rotten litter; 13.V.1983; leg. C. Besuchet & I. Lôbl
[114].

PARATYPE: 1 d , Swat District, above Utrot, 2500-2600 m a.s.1., Abies & Cedrus forest,
sifting rotten wood litter; 14.VI.1983; leg. C. Besuchet & I. Lôbl [12c].

COMPARATIVE MATERIAL EXAMINED: Collinsia japonica, SMF 31674, 4 ®, NEPAL,
southern part of Annapurna Massive, Pass Gorapani, 2700-2800 m a.s.l.; 23.11.1974; leg J.
Martens. — SMF 31670, 2 G , southern part of Dhaulagiri Massive, Dhorpatan, 3000 m a.s..:

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 73

FicGs 47-52

Gorbothorax aff. ungibbus Tanasevitch, 1998. (47-48) Right palp, retrolateral and prolateral
views, respectively. (49-50) Palpal tibia, dorsal and prolateral views, respectively. (51) Distal
suprategular apophysis and embolus. (52) Distal suprategular apophysis and embolic division.

24.V.1973; leg J. Martens. — SMF 31671, 1 %, vestern part of Dhaulagiri Massive, trail from
Dhorpatan to Tarakot, 3100-3600 m a.s1.; 30.V.1973; leg J. Martens. —- SMF 31672, 2 9,
northern part of Dhaulagiri Massive, Dolpo, Tal Valley above Barbung Khola, Charka, 4300 m
a.s.l.; 24.-29.VI.1973; leg. J. Martens. — SMF 31673, 1 d, Thakkhola, Kali-Gandaki Valley,
Chadziou-Khola Valley near Ghasa, 2600 m a.s.1.; IX.1969; leg J. Martens. AII identified by J.
Wunderlich in 1979.

ETYMOLOGY: The specific name is a Latin adjective meaning “concave”,
“cupped” referring to the shape of the embolus.

576 A. V. TANASEVITCH

FiGs 53-57

Halorates concavus Sp. n., & holotype. (53-54) Right palp, retrolateral and ventral views,
respectively. (55) Palpal tibia, dorsal view. (56) Distal suprategular apophysis. (57) Embolic
division.

DIAGNOSIS: The new species is characterized by the peculiar shape of its
embolus.

DESCRIPTION: Male (holotype). Total length 2.03. Carapace 1.00 long, 0.80
wide, unmodified, reddish brown. Chelicerae 0.35 long, with small pointed frontal
tooth. Legs reddish brown. Leg I 2.54 long (0.70+0.25+0.58+0.58+0.43), IV 2.58 long
(0.70+0.23+0.65+0.60+0.40). Chaetotaxy 2.2.2.1, spines weak, their length about
1-1.5 diameter of segment. TmI 0.44. Metatarsus IV without trichobothrium. Palp

LINYPHIID SPECIES FROM PAKISTAN AND INDIA x tif;

(Figs 53-57): Tibia abruptly narrowed distally, ending with a tapering transparent
membranous process. Posterodorsal cymbial outgrowth keel-shaped, with a shallow
saddle. Distal suprategular apophysis flat, spatulate, with a narrow membrane distally.
Radical part of embolic division with two serrate lobes and a strongly sclerotized
tooth-shaped outgrowth; embolus flat and wide, with claw-shaped tip. Abdomen 1.13
long, 0.75 wide, dark grey.

Female unknown.

TAXONOMIC REMARKS: The new species appears to be most similar to H. cras-
sipalpis (Caporiacco, 1935) known from the Karakorum (Caporiacco, 1935; Thaler,
1987), but differs well by the larger size and by the unequal posterodorsal outgrowth
of the cymbium, as well as by the claw-like distal part of the embolus, and also by the
shape of the serrated lobes of the radical part of the embolic division. The new species
clearly differs from H. japonica (O1, 1964) (see Wunderlich, 1983), the only congener
that also occurs in the Himalayas, by the shorter and wider lobes of the embolic
division.

VARIABILITY. Details of the both palps in both males examined look identical.

DISTRIBUTION: Known from the type locality only.

Indophantes tonglu sp. n. Figs 58-63

HOLOTYPE: d , India, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m a.s.l.,
forest near top, sifting litter;, 18.X.1978; leg. C. Besuchet & I. Lôbl [19].

PARATYPE: 1 & , West Bengal, Darjeeling District, Tonglu, northern slope, 2700 m a.s.I.,
forest, sifting litter; 16.X.1978; leg. C. Besuchet & I. LGbl [17].

ETYMOLOGY: The specific epithet is a name in apposition taken from one of two
localities at which this species was found.

DIAGNOSIS: The new species can be easily distinguished by the peculiar shape
of the lamella characteristica, and by the embolus with an expanded thumb carrying a
well-sclerotized black tubercle.

DESCRIPTION: Male (holotype). Total length 1.75. Carapace 0.83 long, 0.68
wide, unmodified, pale brown, almost yellow, with a narrow grey margin. Chelicerae
0.43 long, unmodified. Legs pale yellow. Fel 0.95 long. Leg IV 3.38 long
(0.93+0.20+0.80+0.90+0.55). Chaetotaxy: Til: 2-1-1-0, Till: 2-0-1-0, III-IV: 2-0-0-0,
MtI-IV: 1-0-0-0. Metatarsus IV without trichobothrium. TmI 0.27. Palp (Figs 58-63):
Cymbium with a keel-shaped posterodorsal outgrowth. Distal part of paracymbium
with two tooth-like projections. Lamella characteristica relatively short, its upper
branch thin, 1ts lower one wide, bifurcated. Embolus large, 1ts thumb highly expanded
and carrying a well-sclerotized black tubercle.

Abdomen 0.85 long, 0.60 wide, grey.

Female unknown.

TAXONOMIC REMARKS: The new species seems to be similar to /. agamus
Tanasevitch & Saaristo, 2006, known from Nepal (Tanasevitch & Saaristo, 2006). Both
species can be easily distinguished by the shape of the lamella characteristica and
embolus.

DISTRIBUTION: Known from two localities in northern India.

578 A. V. TANASEVITCH

Fics 58-63

Indophantes tonglu sp. n., 4 holotype (58-59, 61-63) & & paratype (60). (58) Right palp, retro-
lateral view. (59-60) Paracymbium, different aspects. (61) Embolic division. (62) Embolus. (63)
Lamella characteristica.

Indophantes digitulus (Thaler, 1987)

MATERIAL: 2 © , PAKISTAN, Punjab, Murree, 1950 m a.s.l., leaf litter under Quercus:;
23.1V.1984; leg. S. Vit [PAK-84/20]. — 1 ©, Hazara District, Kaäghän Valley, NW of Mahandfri,
Kamalban Forest, 1800 m a.s.]., fern litter; 3.VIL.1985; leg. S. Vit [PAK-85/15]. - 2 G , Nathia
Gali, 2300 m a.s.1., under stones; 17.1V.1984; leg. S.Vit [PAK-84/4]. — 1 9, INDIA, Himachal
Pradesh, Kulu Valley, Chijoga (S of Manali), 1900 m a.s.1., in rotten stump of comiferous tree:
12.X.1988; leg. S. Vit [16]. — 1 ?, Kulu Valley, Vashisht Baths (N of Manali), bank of Beas
River, 1900 m a.s.1., fern litter; 13.X.1988; leg. S. Vit [20].—1 4,3 © , Chijoga (S-Manali), 1900
m a.s.]., wet ravine, from rodent burrow; 14.X.1988; leg. S. Vit [22]. —-3 &,2 9, Khajjiar, E of
Dalhousie, 1950 m a.s.I., Cedrus forest, fern litter; 21.X.1988; leg. S. Vit [30/88].

REMARKS: The species was originally described from Kashmir, India (Thaler,
1987, under Lepthyphantes Menge, 1866) and later recorded from Nepal (Tanasevitch,

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 579

1987; Tanasevitch & Saaristo, 2006). The species is here recorded from Pakistan for
the first time.

RANGE: Himalayan.

Maso sundevalli (Westring, 1851)

MATERIAL: 1 subad. G ,2 ® , PAKISTAN, Punjab, Murree, 1950 m a.s.1., leaf litter under
Quercus; 23.1V.1984; leg. S. Vit [PAK-84/20]. — 1 ®, 3 subad. ©, same, leaf litter under
Aesculus; 23.1V.1984; leg. S. Vit [PAK-84/22]. — 1 © , Hazara District, Käghän Valley, NW of
Mahandri, Kamalban Forest, 2200 m a.s.I., leaf litter under Viburnum; 3.VIL.1985; leg. S. Vit
[PAK-85/18].-— 1 © ,3 subad. © , Dunga Gali, 2300 m a.s.l., leaf litter in scree; 22.1V.1984; leg.
S. Vit [PAK-84/18].

REMARKS: The species is here recorded from the Himalayas and Pakistan for

the first time.

RANGE: Holarctic.

Microbathyphantes palmarius (Marples, 1955)
MATERIAL: 1 G , INDIA, New Dehli, coll. Heimer. — 1 & , Uttar Pradesh, Kumaon, Bhim
Tal, eastern slope, 1500 m a.s.I., edge of secondary forest, sifting, 4.X.1979, leg. I. Lôbl.
REMARKS: The locality in Uttar Pradesh is the northernmost point of the known
distribution of M. palmarius, Iying just at the border between the Palaearctic and the
Oriental regions. The species is here recorded from the Himalayas and India for the
first time.

RANGE: Oriental-Pacific.

Microlinyphia pusilla (Sundevall, 1830)

MATERIAL: 1 9 (MSNV), PAKISTAN, Karakorum, Baltistan, Katzaran, 2200 m assl.,
23.VIL.1976, leg. G. Osella. — 4 £ (MSNV),2 8 (ZMMU), Northern Areas, Skardu District,
Skardu, Pakova, 2300 m a.s.l., 27.VIL.1976, leg. G. Osella.

REMARKS: In the Himalayas this species was hitherto known only from the
Karakorum (Caporiacco, 1935; Helsdingen, 1970).

RANGE: Holarctic.

Oia sororia Wunderlich, 1973

MATERIAL: 1 d, INDIA, West Bengal, between Ghoom and Lopchu, 13 km from
Ghoom, northern slope, 2000 m a.s.1., 14.X.1978, leg. C. Besuchet & I. Lôbl [14].

REMARKS: This species was hitherto known only from Nepal (Wunderlich,
1973, 1983), and is here recorded from India for the first time.

RANGE: Himalayan.

Paracymboides gen. n.
TYPE SPECIES: Paracymboides tibialis sp. n.

ETYMOLOGY: The generic name refers to the peculiar shape of the para -
cymbium; gender masculine.

580 A. V. TANASEVITCH

DIAGNOSIS: The new genus is characterized by the highly modified palpal .
tibia, the narrow, long and mammoth-tusk-like paracymbium, the simple embolic
division with a long embolus, and the well-developed median membrane.

DESCRIPTION: Small-sized erigonines, total length 1.50-1.80 mm. Carapace
unmodified, sulci absent, eyes normal. Chaetotaxy 1.1.1.1.TmI 0.30-0.36. Metatarsus
IV without trichobothrium. Palpal tibia highly modified. Paracymbium narrow, very
long and strong curved. Median membrane well developed and protruded forward.
Embolic division very simple: radical part elongated, flat, without outgrowths or
processes; embolus thin, very long and shightly curved distally. Epigyne without cavity,
its ventral surface (= “ventral plate” sensu Millidge (1984)) hairy. Receptacles rela-
tively large, complex.

TAXONOMIC REMARKS: It is difficult to say anything about the closest relatives
of this genus at present. It 1s likely that they will be found among other SE-Asian
erigOnines.

SPECIES INCLUDED: Paracymboides tibialis sp. n. (the type species) and P.
aduncus Sp. n.

DISTRIBUTION: Southern India.

Paracymboides tibialis sp. n. Figs 64-67, 115

HOLOTYPE: d , INDIA, Tamil Nadu, Nilgiri Hills, Ootacamund, 2150 m a.s.l., sifting
litter under bushes in ravine; 21.X1.1972; leg. C. Besuchet & I. Lôbl [41].

PARATYPES: 2 4,6 9,2 d,2 ® (ZMMU)), from same locality, collected together with
the holotype.— 1 4,1 ©, Tamil Nadu, Nilgiri Hills, Coonoor, 1600 m a.s.1., forest, sifting litter;
22.X1.1972; leg. C. Besuchet & I. Lôbl [43]. - 3 © , same, 6 km E of Coonoor, forest, sifting
litter, 22.X1.1972; leg. C. Besuchet & I. Lôbl [42]. — 1 &, Kerala, NW of Anaimalai Hills,
Nelliampathi Hills, Kaïkatty, 900 m a.s.l., forest, sifting litter near brook; 30.X1.1972; leg. C.
Besuchet & I. Lôbl [58]. — 1 ©, Madras (= Chennai), Cardamom Hills, 6 km NE of Kumily,
700 m ass.l., forest, sifting litter; 3.X1.1972; leg. C. Besuchet & I. Lôbl [7].

ETYMOLOGY: The species name, an adjective, refers to the peculiar shape of the

male palpal tibia.

DIAGNOSIS: The new species can be easily recognized by the peculiar shape of
the male palpal tibia.

DESCRIPTION: Male (paratype). Total length 1.63. Carapace 0.75 long, 0.63
wide, pale reddish brown, unmodified. Chelicerae 0.25 long, unmodified. Legs pale
brown. Leg I 2.21 long (0.65+0.18+0.55+0.45+0.38), IV 2.19 long (0.65+0.18+
0.58+0.45+0.33). Chaetotaxy 1.1.1.1, spines weak, their length about same as diameter
of segment or a little longer. TmI 0.36. Metatarsus IV without trichobothrium. Palp
(Figs 64-66): Tibia extended in dorso-ventral directions, with a narrow retrolateral out-
growth curved upward. Paracymbium very long, narrow, curved upward almost to a
full circle. Distal suprategular apophysis massive, complex in shape. Median mem-
brane very long, protruded forward and covering distal part of embolus. Radical part
of embolic division elongated and lacking of outgrowths. Base of embolus wide, bent

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 581

LE à

FIiGs 64-67

Paracymboides tibialis sp. n., & ® paratypes from Ootacamund, Tamil Nadu. (64-65) Right
palp, retrolateral and ventral views, respectively. (66) Embolic division. (67) Epigyne, ventral
vieW.

180°, embolus very long, narrow, slightly curved distally. Abdomen 1.00 long, 0.65
wide, grey.

Female. Total length 1.75. Carapace 0.73 long, 0.60 wide, unmodified. Cheli
cerae 0.30 long, unmodified. Leg I 2.14 long (0.58+0.20+0.53+0.45+0.38), IV 1.71
long (0.48+0.20+0.38+0.35+0.30). TmI 0.34. Abdomen 1.20 long, 0.90 wide. Epigyne
and vulva as shown in Figs 67, 115. Body and leg coloration, as well as chaetotaxy, as
in male.

TAXONOMIC REMARKS: Paracymboides tibialis Sp. n. is similar to P. aduncus
Sp. n.: their embolic division is almost identical. These species can be easily distin-

582 A. V. TANASEVITCH

FiGs 68-72

Paracymboides aduncus sp. n., & holotype, ® paratype from Cardamom Hills, Muttapatti.
(68-69) Right palp, retrolateral and prolateral views, respectively. (70-71) Palpal tibia, dorsal and
prolateral views, respectively. (72) Epigyne, ventral view.

guished by the shape of the palpal tibia and of the distal suprategular apophysis. The
female P. tibialis sp. n. differs from that of P. aduncus sp. n. by the absence of the two-
humped outgrowth on the posterior side of the epigyne.

DISTRIBUTION: Southern India.

Paracymboides aduncus sp. n. Figs 68-72, 116
HOLOTYPE: 4 , INDIA, Tamil Nadu, Palni Hills, 23 km W of Kodaïkanal, Lake Berijam,
2150 m a.s.l., Rhododendron forest, sifting litter; 14.X1.1972; leg. C. Besuchet & I. Lbl [26].

PARATYPES: 1 &,1 ®, INDIA, Kerala, Cardamom Hills, 13 km NE of Munnar, 1900 m
a.s.l., forest, sifting litter; 26.X1.1972; leg. C. Besuchet & I. Lôbl [51].-— 1 % , Cardamom Hills,

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 583

near Munnar, Muttapatti, 1700 m a.s.l, forest, sifting litter under tree ferns; 24.X1.1972; leg. C.
Besuchet & I. Lôbl [1972/48].

ETYMOLOGY: The species name is a Latin adjective meaning “hook-shaped”,
referring to the shape of the palpal tibia outgrowth.

DIAGNOSIS: The new species can be easily recognizable by the peculiar shape
of the male palpal tibia.

DESCRIPTION: Male (holotype). Total length 1.55. Carapace 0.68 long, 0.55
wide, unmodified, pale reddish brown, with a narrow black margin. Chelicerae 0.23
long, unmodified. Legs pale brown. Leg I 2.08 long (0.58+0.18+0.50+0.45+0.37), IV
2.13 long (0.60+0.20+0.55+0.43+0.35). Chaetotaxy 1.1.1.1, spines weak, their length
a little more than diameter of segment. TmI 0.31. Metatarsus IV without tricho-
bothrium. Palp (Figs 68-71): Palpal tibia with a long and narrow outgrowth starting at
retrolateral side and running orthogonally of main palpal axis to prolateral side.
Paracymbium very long, narrow, strongly curved. Distal suprategular apophysis short,
wide and rounded. Median membrane very long, protruded forward and covering distal
part of embolus. Radical part of embolic division elongate and lacking outgrowths.
Base of embolus wide, bent 180°, embolus very long, narrow, slightly curved.
Abdomen 0.90 long, 0.58 wide, dark grey.

Female. Total length 1.58. Carapace 0.63 long, 0.53 wide. Chelicerae 0.18 long,
unmodified. Leg I 1.99 long (0.50+0.18+0.48+0.45+0.38), IV 2.17 long (0.63+0.20+
0.58+0.43+0.33). TmI 0.32. Epigyne and vulvae as shown in Figs 72, 116. Body and
leg coloration, as well as chaetotaxy, as in male.

TAXONOMIC REMARKS: Paracymboides aduncus sp. n. is similar to its only
known congener, P. tibialis sp. n.; see above.

DISTRIBUTION: Southern India.

Paragongylidiellum caliginosum Wunderlich, 1973

MATERIAL: 4 4,7 2:;:2@,2 $ (ZMMU), India, Madras (= Chennai), Anaimalai Hills,
18 km N of Valparai, 1250 m a.s.l., forest, sifting litter; 18.X1.1972; leg. C. Besuchet & I. Lôbl
[35].

REMARKS: This species was hitherto known only from Nepal (Wunderlich,
1973, 1983), and is here recorded from India for the first time.

DISTRIBUTION: Nepal Himalayas and southern India.

Pelecopsis indus sp. n. Figs 73-80

HOLOTYPE: à , INDIA, Uttar Pradesh, Garhwal, above Pauri, northern slope, 1900 m
a.s.1., Quercus dry forest, sifting leaf litter and moss; 28.X.1979; leg. I. Lôbl [28].

PARATYPES: 1 G,3 ©, from same locality, collected together with the holotype. — 1 à,
4 ©, Uttar Pradesh, Kumaon, Rangarh, 2000 m a.s.., forest in ravine, sifting leaf litter;
9.X.1979; leg. I. Lôbl [6b]. — 1 $ , Garhwal, 2 km E of Dhanolti, northern slope, 2250 m as...
near brook, Abies & Rhododendron forest, sifting leaf litter; 21.X.1979; leg. I. Lôbl [19].—1 9,
Kumaon, Naini Tal, 2000-2100 m a.s.I., forest in ravine, sifting leaf litter and moss: 8.X.1979;
leg. I. Lôbl [5b].-— 1 4, Himachal Pradesh, Kulu Valley, Naggar, 1850 m a.s.l., rotten stump of
Cedrus, under bark; 16.X.1988; leg. S. Vit [25]. — 1 4, Kulu Valley, Vashisht Baths (N of
Manali), 1900 m a.s..; 13.X.1988; leg. S. Vit [18]. —- 1 $, PAKISTAN, Swat District, Kalam,
2100 m a.s.., Quercus forest, sifting leaf litter; 12.V.1983; leg. C. Besuchet & I. Lôbl [9b]. —

584 A. V. TANASEVITCH

FiGs 73-80

Pelecopsis indus Sp. n., & & 9 paratypes from Pauri, Uttar Pradesh. (73-74) Male carapace,
lateral and dorsal views, respectively (not to scale). (75-76) Right palp, retrolateral and prolateral
views, respectively. (77) Palpal tibia, dorsal view. (78-80) Epigyne and vulva, ventral, dorsal
(inclined 90°) and anterodorsal views, respectively.

1 ©,S of Saidu Sharif, Murghazar, leaf litter under Platanus, 1300 m a.sl.; 8.V.1983; leg. C.
Besuchet & I. Lôbl [2b]. - 1 4, Ushu Valley, upper reaches of Kalam River, 2300 m ass.
Cedrus forest, sifting leaf litter under Corylus; 15.V.1983; leg. C. Besuchet & I. Lôbl [13b]. —

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 585

1 ©, Malam Jabba, 2300 m a.s.., Pinus forest, sifting under Pinus and Corylus; 9.V.1983; leg.
C. Besuchet & I. LGbl [4c].

COMPARATY VE MATERIAL EXAMINED: Pelecopsis minor, SMF 60195,3 G ,2 % paratypes,
Dundgovi Aimak, 20 km S of Somon Delgertsogt, 1480 m a.s..; 9.VI.-13.VIL.1967; leg. Z.
Kaszab, det. J. Wunderlich.

ETYMOLOGY: The specific name, a noun in apposition, refers to an inhabitant of
India.

DIAGNOSIS: The new species 1s characterized by the smooth surface of its cara-
pace (without pits), by the shape of the male palpal tibia, as well as by the peculiar
shape of the ventral plate of the epigyne.

DESCRIPTION: Male (paratype). Total length 1.65. Carapace 0.78 long, 0.58
wide, modified as shown in Figs 73-74; pale brown, with indistinct grey radial stripes.
Surface of carapace smooth, without pits. Chelicerae 0.33 long, unmodified. Legs
yellow. Leg I 2.16 long (0.60+0.20+0.50+0.48+0.38), IV 2.22 long (0.63+0.20+0.58+
0.48+0.33). Chaetotaxy 0.0.0.0. TmI 0.39. AÏI metatarsi with trichobothrium. Palp
(Figs 75-77): Tibia with a long, narrow, prolateral process. Cymbium without postero -
dorsal outgrowth. Distal suprategular apophysis relatively short, wedge-shaped.
Embolic division fusiform, with a narrow, long and curved membranous process
starting near base of embolus, running forward and covering end of embolus. Abdomen
0.90 long, 0.63 wide, grey, scutum absent.

Female. Total length 1.75. Carapace 0.88 long, 0.63 wide, unmodified. Cheli -
cerae 0.38 long, unmodified. Leg I 2.18 long (0.65+0.20+0.55+0.45+0.33), IV 2.31
long (0.70+0.20+0.63+0.45+0.33). Chaetotaxy 0.0.0.0. TmI 0.44. Abdomen 1.00 long,
0.75 wide. Epigyne and vulva as in Figs 78-80. Body and leg coloration, as well as
chaetotaxy, as in male.

TAXONOMIC REMARKS: The new species is similar to the Mongolian P. minor
Wunderlich, 1995, but differs by bigger body size of both sexes, by the presence a
trichobothrium on metatarsus IV, by the smooth carapace surface without pits, as well
as by some details of the genitalia.

RANGE: Himalayan.

Piniphantes himalayensis (Tanasevitch, 1987)

MATERIAL: 1 & , PAKISTAN, Swat District, above Utrot, 2800 m a.s.1., Abies & Cedrus
forest, sifting rotten coniferous litter; 13.V.1983; leg. C. Besuchet & I. Lôbl [11c].

REMARKS: This species was hitherto known from Kashmir (Thaler, 1987) and
from numerous localities in Nepal (Tanasevitch, 1987).

RANGE: Himalayan.

Porrhomma pygmaeum (Blackwall, 1834) Fig TE

MATERIAL: 1 © (MSNV), PAKISTAN, Northern Areas, Ghangche District, Ghangche,
Kaplu Ghwari, 2480 m a.s.1.; 26.VI.2008; leg. L. Latella.

REMARKS: The species is here recorded from the Himalayas and Pakistan for
the first time.

RANGE: Palaearctic.

586 A. V. TANASEVITCH

Prinerigone vagans (Audouin, 1826)

MATERIAL: 1 4 (MSNV), PAKISTAN, Karakorum, Baltistan, Shalabot, 1700 m a.s..:
15.VIIL.1976; leg. G. Osella. — 1 & (MSNV), Northern Areas, Skardu District, Skardu, Pakova,
2300 m a.s..; 27.VI1.1976; leg. G. Osella. — 1 4 (MSNV), Skardu, Lake Satpara, 2700 m as;
1.VIL.1976; leg. G. Osella. —- 3 4 (MSNV), Ghangche District, Khaplu, Sciaiak Channel, 2400
m a.s.l.; 10.VIL.1976; leg. G. Osella.

REMARKS: The species is here recorded from the Himalayas and Pakistan for
the first time.

RANGE: Old World.

Scotargus pilosus Simon, 1913

MATERIAL: 1 ®, PAKISTAN, Hazara District, Käghän Valley, 1450 m a.s.., Malkandi
Forest, rotten coniferous stub; 29.VI.1985; leg. S. Vit [PAK-85/3].-4 4,8 © , INDIA, Himachal
Pradesh, Kulu Valley, Naggar, 1850 m a.s.., litter;, 16.X.1988; leg. S. Vit [24]. — 3 ®, Uttar
Pradesh, Kumaon, Chaubattia near Ranikhet, 1800 m a.s.l., forest, sifting leaf litter and moss
under fern; 12.-13.X.1979; leg. I. Lôbl [10]. —- 2 4, 1 £ (ZMMU), Garhwal, 10 km E of
Dhanolti, 2450 m a.s.l., northern slope, Quercus forest, sifting litter; 21.X.1979; leg. I. Lôbl
[21b]. — 2 ©, Garhwal, 10 km W of Chamba, 2200 m a.s.., sifting mosses under bushes;
20.X.1979; leg. I. Lôb] [17].—1 4,4 £ (ZMMU), Garhwal, 4 km S of Bhatwari, 1400 m asl.,
timberline, degraded forest, sifting moss and fern litter; 23.X.1979; leg. I. Lôbl [22]. — 1 G,
Garhwal, above Pauri, northern slope, 1900 m a.s.., Quercus dry forest, sifting leaf litter and
moss; 28.X.1979; leo. I. Lôbl [28].- 1 & , Garhwal, 2 km E of Dhanolti, northern slope, 2250 m
a.s.l., near brook, Abies & Rhododendron forest, sifting leaf litter; 21.X.1979; leg. I. Lôbl [19].

REMARKS: In the Himalayas this species was hitherto known only from

Kashmir (Thaler, 1987).
RANGE: European-Ancient Mediterranean.

Tapinocyboides bengalensis sp. n. Figs 81-85
HOLOTYPE: & , INDIA, West Bengal, Darjeeling District, Tonglu, 3100 m a.s.l., near top,
under stones; 16.X.1978; leg. C. Besuchet & I. Lôbl [16a].
ETYMOLOGY: The species name, an adjective, refers to the Indian State where
the new species was found.

DIAGNOSIS: The new species is characterized by the peculiar shape of the split
palpal tibia.

DESCRIPTION: Male. Total length 1.40. Carapace 0.63 long, 0.53 wide, un -
modified, pale brown, sulci absent. Chelicerae 0.25 long, unmodified. Legs yellow.
Leg I 1.73 long (0.50+0.15+0.38+0.37+0.33), IV 1.72 long (0.52+0.17+0.43+0.35+
0.25). Chaetotaxy 1.1.1.1, length of spines about same as diameter of segment or a little
longer. Tml 0.32. Metatarsus IV without trichobothrium. Palp (Figs 81-85): Tibia
dorsally deeply divided by a narrow cleft into two parts. Paracymbium simple,
U-shaped. Distal suprategular apophysis weakly sclerotized, relatively short, wide and
rounded, with a pale thin tooth near it base. Median membrane well developed,
protruded forward, distally extended. Radical part of embolic division flat, elongated.
Embolus thin, long and coiled, with a narrow membranous edge. Abdomen 0.80 long,
0.55 wide, grey.

Female unknown.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 587

81, 82, 84, 85

FiGs 81-85

Tapinocyboides bengalensis sp. n., 4 holotype. (81-83) Right palp, retrolateral, prolateral and
dorso-prolateral views, respectively. (84) Palpal tibia, dorsal view. (85) Distal suprategular
apophysis and median membrane.

TAXONOMIC REMARKS: This new species is probably not congeneric with the
type species Tapinocyboides pygmaeus (Menge, 1869) because it has no “lamella”
sensu Merrett (1963) in the embolic division and no sulci on the male carapace.
Therefore so it is assigned to this genus only provisionally. In the absence of a female,
and/or other possible congeners, the taxonomic position of this species is unclear.

DISTRIBUTION: Known from the type locality only.

Tchatkalophantes baltistan sp. n. Figs 86-88

HOLOTYPE: 1 9 (MSNV), PAKISTAN, Karakorum, Baltistan, Shalabot, 1700 m a.s.l.:
15.VIIL.1976; leg. G. Osella.

588 A. V. TANASEVITCH

88

PMP

FIiGs 86-88

Tchatkalophantes baltistan sp. n.. ? holotype. (86-88) Epigyne, ventral, dorsal (upside down)
and lateral views, respectively.

ETYMOLOGY: The specific name is taken from the name of the country of
Origin.

DESCRIPTION: Female. Total length 1.96. Carapace 0.78 long, 0.60 wide, un-
modified, pale brown, almost yellow. Chelicerae 0.35 long, unmodified. Legs mostly
lost, yellow. Leg I 3.42 (0.93+0.23+0.88+0.83+0.55), Fel 0.90 long. Chaetotaxy: Fel:
0-1-0-0; Tif: 2-1-1-0, Till: 2-0-1-0, II-IV: 7; Mt: 1-0-0-0, II-IV: ? TmI 0.25. Abdomen
1.30 long, 0.90 wide, dorsally dark grey, with irregularly arranged small white spots.
Epigyne (Figs 86-88): Aperture wide. Proscape with a narrow base, stepwise widening
in middle part. bifurcated distally. Posterior median plate triangular.

Male unknown.

TAXONOMIC REMARKS: The new species is similar to the Mongolian T. hyper-
auritus (Loksa, 1965) and differs by a shallower depression in the distal part of the
proscape, as well as by the shape of the posterior median plate: in 7. baltistan Sp. n. it
wider than high vs higher than wide in T. hyperauritus.

DISTRIBUTION: Known from the type locality only.

Tenuiphantes tenuis (Blackwall, 1852)

MATERIAL: 2 9 (MSNV). PAKISTAN, Karakorum, Baltistan, Shalabot, 1700 m a.s.l.:
15.VIIL.1976; leg. G. Osella. — 1 4 (MSNV). Northern Areas, Gilgit District, Bagrot Valley, trap

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 589

Fics 89-96

Tiso incisus sp. n., 4 & © paratypes from Jutogh, Himachal Pradesh. (89) Right palp, retro-
lateral view. (90) Paracymbium. (91-92) Palpal tibia, dorsal and prolateral views, respectively.
(93) Distal suprategular apophysis. (94) Embolic division. (95) Epigyne, ventral view. (96)
Vulva, dorsal view.

#5; 25.X.-2.X1.2008; leg. L. Latella & R. Ahmed. — 1 © (MSNV), Bagrot Valley; 17.VI.2008:
leg. L. Latella.

REMARKS: The species is here recorded from the Himalayas and Pakistan for
the first time.

RANGE: European-Ancient Mediterranean.

590 A. V. TANASEVITCH

Tiso incisus Sp. n. Figs 89-96
HOLOTYPE: G , INDIA, Himachal Pradesh, Jutogh, 10 km W of Shimla, 2000 m as.
leaf litter near foot of rock: 29.X.1988; leg. S. Vit [37].

PARATYPES: 13 4,19 2;5 4,3 © (ZMMU), from same locality, collected together with
the holotype.— 1 © , Himachal Pradesh, Kulu Valley, S of Manali, Chijoga, 1900 m a.s.1., hollow
in Quercus; 14.X.1988; leg. S. Vit [21]. —- 1 4,2 ©, Uttar Pradesh, Garhwal, 10 km E of
Dhanolti, 2450 m a.s.., northern slope, Quercus forest, sifting litter; 21.X.1979; leg. I. LGbl
[21b].-1 © , PAKISTAN, Swat District, S of Saidu-Sharif, Marghuzar, 1300 m a.s.., sifting leaf
litter under Platanus; 8.V.1983; leg. C. Besuchet & I. Lôbl [2b]. — 1 4,3 ©, Malam Jabba,
2300 m a.s.l., Pinus forest, sifting litter under Pinus and Corylus; 9.V.1983; leg. C. Besuchet &
I. Lôbl [4c]. — 1 & , same, 2500-2600 m a.s.., under stones; 18.V.1983; leg. C. Besuchet & I.
Lôbl [17a].—1 4,5 © ,same, 2500-2600 m a.s.., Abies forest, sifting litter and moss; 18.V.1983;
leg. C. Besuchet & I. Lôbl [17b].— 1 4,3 ©, Kalam, 2100 m a.s.., Quercus forest, sifting leaf
litter; 12.V.1983; leg. C. Besuchet & I. Lôbl [9b].- 7 ©, Ushu Valley, upper reaches of Kalam
River, 2300 m a.s.1., Cedrus forest, sifting leaf litter under Corylus; 15.V.1983; leg. C. Besuchet
& I. Lôbl [13b].-—1 4,1 © , above Miandam, 2400-2500 m a.s.l., Abies forest, sifting litter and
moss; 17.V.1983; les. C. Besuchet & I. Lôbl [15b].-— 1 &,1 © , same, 2400-2500 m a.s.., Abies
forest, sifting litter and moss: 17.V.1983; leg. C. Besuchet & I. Lôbl [15b].

ETYMOLOGY: The species name means “incised”, “notched”, referring to the

shape of the embolus.

DIAGNOSIS: The new species is characterized by the peculiar shape of the palpal
tibia and the embolic division, as well as by the shape of the ventral plate of the
epigyne.

DESCRIPTION: Male. Total length 1.40. Carapace 0.75 long, 0.58 wide, pale
brown, unmodified, sulci absent. Chelicerae 0.23 long, unmodified. Legs pale brown.
Leg I 1.88 long (0.55+0.20+0.45+0.38+0.30), IV 2.06 long (0.60+0.20+0.55+0.43+
0.28). Chaetotaxy 1.1.1.1,spines weak, their length about same as diameter of segment
or a little longer. TmI 0.44. Metatarsus IV without trichobothrium. Palp (Figs 89-94):
Patella as long as cymbium, with a row of uniform short spines dorsally. Tibia strong-
ly modified: elongated, slightly sigmoid, with several outgrowths different in size and
shape. Paracymbium small and narrow, L-shaped. Distal suprategular apophysis long
and wide, flat, apically pointed and bent 90°. Radical part of embolic division relatively
large and flat. Embolus long, curved, with a notch near it base. Abdomen 0.78 long,
0.50 wide, pale grey.

Female. Total length 1.38. Carapace 0.68 long, 0.53 wide, unmodified. Cheli-
cerae 0.23 long, unmodified. Leg I 1.64 long (0.48+0.20+0.38+0.30+0.28), IV 1.76
long (0.50+0.20+0.45+0.35+0.26). TmI 0.43. Abdomen 0.80 long, 0.55 wide. Epigyne
and vulva (Figs 95-96): Ventral plate of epigyne long and narrow. Receptacles sphe -
rical, far apart from each other. Body and leg coloration, as well as chaetotaxy, as in
male.

TAXONOMIC REMARKS: The new species seems to be most similar to the
Holarctic T. aestivus (L. Koch, 1872) and can be easily distinguished by the structure
of the embolic division: in T. incisus sp. n. the loop of the embolus is almost orthogonal
to the main axis of the palp, whereas in 7. aestivus the loop lies almost in the same
plane; there are also other small differences in the shape of the palpal tibia and the
embolic division. The female differs by the shape of the ventral plate of the epigyne:
in 7. aestivus the ventral plate is triangular, in 7. incisus Sp. n. it is as a narrow trans-
versal stripe.

RANGE: Himalayan.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 591

100

FiGs 97-100

Tiso (?) indianus Sp. n., & paratype. (97-98) Right palp, retrolateral and prolateral views,
respectively. (99) Palpal tibia, dorsal view. (100) Palpal tibia and paracymbium, lateral view.

Tiso (?) indianus sp. n. Figs 97-100, 118
HOLOTYPE: G , INDIA, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m a.s..,
near top, forest, sifting litter, 18.X.1978; leg. C. Besuchet & I. Lôbl [19].
PARATYPE: 1 & , from same locality, collected together with the holotype.
ETYMOLOGY: The specific name is derived from the name of the country of
origin; adjective.
DIAGNOSIS: The new species is characterized by the peculiar shape of the
embolus and the palpal tibia.

DESCRIPTION: Male (paratype). Total length 1.40. Carapace 0.68 long, 0.50
wide, unmodified, greyish brown. Chelicerae 0.21 long, unmodified. Legs yellow. Leg
I 1.78 long (0.51+0.17+0.42+0.39+0.29), IV 1.81 long (0.53+0.15+0.45+0.39+0.29).
Chaetotaxy 1.1.1.1, length of spines about same as diameter of segment or a little
longer. TmI 0.42. Metatarsus IV without trichobothrium. Palp (Figs 97-100, 118): Tibia
with apical outgrowth narrowed at base. Paracymbium narrow, relatively long, hook-

592 A. V. TANASEVITCH

FiGs 101-104

Male carapace of Walckenaeria martensi Wunderlich, 1972, carapace, lateral view. (101-103)
Specimens from Tonglu, West Bengal. (104) Specimen from Thare Pati, Nepal.

shaped distally. Radical part of embolic division small, embolus very wide, flat,
crescent-shapes. Abdomen 0.75 long, 0.56 wide, grey.
Female unknown.

TAXONOMIC REMARKS!: In the shape of the paracymbium the species is similar
to the representatives of the genus Paracymboides gen. n., but the structure of the
embolic division is quite different and very peculiar, and this character does not allow
to place T. indianus sp. n. in that genus. The absence of a corresponding female or of
a related species makes the systematic position of the species unclear. Its placement in
the genus 7iso Simon, 1884 is thus only provisional.

DISTRIBUTION: Known from the type locality only.

Walckenaeria martensi Wunderlich, 1972 Figs 101-104
Walckenaeria nepalensis Wunderlich, 1972 syn. n.

TYPE MATERIAL EXAMINED: Walckenaeria martensi, SMF 25298/1, 4 holotype, NEPAL,
Khumbu, Everest Region at confluent of Imja- and Phunki-Drangka rivers, Betula forest, 3200-
3250 m a.sl.; 30.1X.-2.X.1970; leg. J. Martens. —- SMF 25299/7, 1 d,6 9 paratypes, same
locality, collected together with holotype.— SMF 25300,2 © paratypes, NEPAL, Khumbu, Pare,
Nangba-Tsangpo Valley, subalpine forest with Betula utilis and Rhododendron campanulatum,
3350 m a.s1.; 14.16.X.1970; leg. J. Martens. — SMF 25302/1, 1 subad. ® paratype, Gorapani,
Rhododendron forest along stream, 2850-2900 m a.s.l.; 10.-14.X11.1969:; leg. J. Martens. W.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 593

108

FiGs 105-108

Walckenaeria saetigera sp. n., 4 holotype (105-106), 4 paratype from Weiloi, Meghalaya (107)
and £ paratype from Shillong, Meghalaya (108). (105-107) carapace, lateral (105, 107) and dor-
sal (106) views, respectively. (108) Epigyne, ventral view (not to scale).

nepalensis, SMF 25303/1, 4 holotype, NEPAL, Kathmandu-Tal, Mt Phulchoki, Quercus semi-
carpifolia forest, 2600-2700 m a.sl.; 25.-30.1.1970; leg. J. Martens. —- SMF 25304/4, 4 9
paratypes, same locality, collected together with holotype.

OTHER MATERIAL: 2 à , INDIA, West Bengal, Darjeeling District, Tonglu, 3100 m a.s.,
near top, under stones;16.X.1978; leg. C. Besuchet & I. Lôbl [16a].-—1 4,3 9 , same, sifting in
dwarf forest under brushes on pasture; 16.X.1978; leg. C. Besuchet & I. Lôbl [16b]. New records
of W. martensi:1 ® , NEPAL, Bagmati Province, above Gul Bhanjyang, 2600 m a.s.1., northern
slope, old Quercus forest, sifting litter and rotten wood under trees; 6.1V.1981; leg. I. Lôbl & A.
Smetana [9]. —1 4,1 % , lower Thare Pati, 3300 m a.s.1., sifting litter under fern and Acer near
stream; 10.1V.1981; leg. I. Lôb]l & A. Smetana [18b].-— 1 © , lower Thare Pati, 3500 m a.s.I., Acer
forest, sifting litter;, 12.1V.1981; leg. I. Lôbl & A. Smetana [21].-—3 © , Malemchi, 2800 massl.,
sifting litter, 14.1V.1981; leg. I. Lôbl & A. Smetana [24]. - 3 © , above Shermathang, 2900 m
a.s.1., in faeces; 26.1V.1981; leg. I. Lôbl & A. Smetana [47b]. — 1 ©, Kathmandu District,
Phulcoki, 2600-2700 m a.s.l., dry forest, sifting litter and rotten wood along fallen trunks:
15.X.1983; leg. I. Lôbl & A. Smetana [36]. -— 1 4,1 © , Sankhuwasawa District, Kosi Province,
NE of Kuwapani, 2350 m a.s.l., sifting litter and humus near spring; 5.1V.1984; leg. I. Lôbl & A.
Smetana [5].-— 1 © , same, 2250 m ass. sifting litter near rotten trunk, 6.1V.1984; leg. I. Lôbl &
A. Smetana [6].-—1 ? ,NE of Mangmaya, 2300 m a.s.1., northern slope, dry forest, sifting rotten
wood and litter; 6.1V.1984; leg. I. Lôbl & A. Smetana [7].-—2 © , southern part of Mangsingma,
2200 m a.s.1., and rotten wood in ravine; 11.1V.1984; leg. I. Lôbl & A. Smetana [13]. -— 1 9,
Induwa Kola Valley, 2000 m a.s.l., sifting litter and rotten wood; 14.1V.1984; leg. I. Lôbl & A.
Smetana [18].— 1 ®, same, 2100 m a.s.., sifting litter; 17.1V.1984; leg. I. Lôbl & A. Smetana
271:

REMARKS: All three males of W. martensi from Tonglu, West Bengal have palps
identical to those of the type specimens, but the shapes of the cephalic elevations are

slightly different from each other (see Figs 101-104) and similar to that of the holotype

594 A. V. TANASEVITCH

109-111

112, 113

FiGs 109-113

Walckenaeria saetigera sp. n., & holotype. (109-110) Right palp, retrolateral and prolateral
views, respectively. (111) Palpal tibia, dorsal view. (112-113) Embolic division and distal
suprategular apophysis, different aspects.

Of W. nepalensis, described from a single male and several females from Nepal
(Wunderlich, 1972). A detailed comparison of the male palps of all available material
Of W. martensi and W. nepalensis (see above) shown that they belong to the same
species. Epigynes are very variable and I could not find identical ones in specimens
even from the same locality. W. martensi, Was described a few pages earlier, therefore
I decided that W. nepalensis becomes its junior synonym.

RANGE: Himalayan.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 595

FiGs 114-119

Gongylidioides keralaensis sp. n., ® paratype from Cardamom Hills, Muttapatti, India (114);
Paracymboides tibialis sp. n., ? paratype from Ootacamund, Tamil Nadu, India (115); P. adun-
cus Sp. n., ® paratype from Cardamom Hills, Muttapatti, India (116); Porrhomma pygmaeum
(Blackwall, 1834), © from Kaplu Ghwari, Pakistan (117); Tiso (?) indianus Sp. n., 4 paratype
from Tiger Hill, India (118); Walckenaeria saetigera Sp. n., ® paratype from Shillong,
Meghalaya, India (119). (114-117) Vulva. (118) Embolic division and distal suprategular apo -
physis. (119) Genital region, ventral view. Not to scale.

Walckenaeria saetigera sp. n. Figs 105-113, 119

HOLOTYPE: d, INDIA, Meghalaya, above Shillong, Khasi Hills, northern slope,
1850-1950 m a.s.1., primary forest near Shillong Peak, sifting litter; 25.X.1978; leg. C. Besuchet
& I. Lôbl [27].

596 A. V. TANASEVITCH

PARATYPES: 2 ©, from same locality, collected together with holotype. — 1 9,
Meghalaya, above Cherrapunjee, Khasi Hills, 1200 m a.s.1., forest, near foot of rock, sifting
litter; 26.X.1978; leg. C. Besuchet & I. Lôbl [28b]. — 1 4, Khasi Hills, Weïloi, 1700 masi,
sifting in forest; 27.X.1978; leg. C. Besuchet & I. Lôbl [31].

ETYMOLOGY: The species name is a Latin adjective meaning “carrying
bristles”, referring to the group of bristles on the male palpal tibia of this species.

DIAGNOSIS: The new species 1s characterized by the peculiar shape of the male
carapace and the shape of the palpal tibia. The epigyne is also well distinct from that
of congeners.

DESCRIPTION: Male (holotype). Total length 2.30. Carapace 1.08 long, 0.75
wide, modified as shown in Figs 105-107, pale orange-yellow. Chelicerae 0.30 long,
unmodified. Legs pale orange-yellow. Leg I 2.81 long (0.83+0.25+0.73+0.60+0.40),
IV 2.94 long (0.83+0.23+0.83+0.65+0.40). Chaetotaxy 2.2.1.1, spines very thin, their
length about same as diameter of segment or a little longer. TmI 0.53. AII metatarsi
with a trichobothrium. Palp (Figs 109-113): Tibia sickle-shaped, dorsal surface bearing
a group of short stout bristles. Paracymbium small, L-shaped. Distal suprategular
apophysis distally narrow, claw-shaped. Radical part of embolic division fusiform,
with a large conical outgrowth near base of embolus. Embolus relatively short, curved
to a semicircle. Abdomen 1.15 long, wide, 0.75, grey.

Female. Total length 2.33. Carapace 108 long, 0.80 wide, unmodified.
Chelicerae 0.35 long, unmodified. Leg I 3.11 long (0.90+0.30+0.83+0.65+0.43), IV
3.29 long (0.92+0.28+0.88+0.78+0.43). TmI 0.45. Abdomen 1.15 long, 0.78 wide.
Epigyne (Fig. 108, 119): Aperture fully covered by tapered ventral plate, with a trans-
versal wrinkle posteriorly. Receptacles oval, well visible on both sides of ventral plate.
Body and leg coloration, as well as chaetotaxy, as in male.

VARIABILITY: The shapes of the male carapaces in the holotype and the
paratype are slightly different (Fig. 105 cf. Fig. 107).

TAXONOMIC REMARKS: The epigyne bears some resemblance to that of the
representatives of the subgenus Wideria Simon, 1864, sensu Wunderlich (1972), but
the embolic division shows similarities to that of the subgenus Prosopotheca Simon,
1884.

DISTRIBUTION: Known from Meghalaya, northeastern India.

ACKNOWLEDGEMENTS

I am most grateful to all persons whose collections I have used for the present
study. I wish to thank Peter Jäger and Julia Altmann (SMF) for providing types and
comparative material. Special thanks to Leonardo Latella and Francesco Ballarin
(MSNV) who made available for me a small collection from Pakistan. I am also very
much obliged to Peter J. Schwendinger (MHNOG) for the opportunity to work on the
spider collections of the MHNG and for checking the manuscript, as well as to Peter
van Helsdingen (Leiden, Netherlands) for helpful comments. This study was supported
in part by the Russian Foundation for Basic Research, project # 09-04-01365-a.

LINYPHIID SPECIES FROM PAKISTAN AND INDIA 597

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REVUE SUISSE DE ZOOLOGIE

Tome 118 — Fascicule 3

GATT, Paul. Microphorella cassari sp. n., a new species of Microphorella
Becker (Diptera: Dolichopodidae) from Tumisia ................

TRAN, Dao Thi Anh, NGUYEN, Tao Thian, PHUNG, Trung My, LyY, Tri,
BÔHME, Wolfgang, and ZIEGLER, Thomas. Redescription of Rhaco-
phorus chuyangsinensis Orlov, Nguyen & Ho, 2008 (Anura:
Rhacophoridae) based on new collections from new south
Vietnamese provincial records: Lam Dong and Khanh Hoa .......

LIENHARD, Charles & ASHMOLE, N. Philip. The Psocoptera (Insecta:
Psocodea) of St Helena and Ascension Island (South Atlantic) with a
EN ECO TTONL OUR AITIER 20. 1 0... LA tu inoante mare

LIENHARD, Charles & BAZ, Arturo. Redescription of the genus Marce -
nendius Naväs (Psocodea: ‘Psocoptera’: Amphientomidae) with a
key to western Palaearctic amphientomids ....................

GONZALEZ-SOLIS, David & MARIAUX, Jean. An annotated list of the para-
sitic nematodes (Nematoda) of freshwater fishes from Paraguay
deposited in the Museum of Natural History of Geneva ..........

WEBER, Jean-Marc. Food habits of escaped Eurasian otters (Lutra lutra)
ina subDurban environment 1n SWIfZerland .…., sen sed

HLAVAC, Peter. Claude Besuchet, an eminent swiss coleopterists, 80 years
M ln des ar as an
SENGLET, Antoine. New species in the Zelotes tenuis-group and new of
little known species in other Zelotes groups (Gnaphosidae, Araneae)

TANASEVITCH, Andreï V. Linyphiid spiders (Araneae, Linyphiidae) from
D . . .. Pl à

Pages

401-412

413-421

423-449

451-466

467-484

485-489

491-512

513-559

561-598

REVUE SUISSE DE ZOOLOGIE

Volume 118 — Number 3

GATT, Paul. Microphorella cassari Sp. n., a new species of Microphorella
Becker (Diptera: Dolichopodidae) from Tunisia ................

TRAN, Dao Th1 Anh, NGUYEN, Tao Thian, PHUNG, Trung My, LY, Tri,
BÔHME, Wolfgang, and ZIEGLER, Thomas. Redescription of Rhaco -
phorus chuyangsinensis Orlov, Nguyen & Ho, 2008 (Anura:
Rhacophoridae) based on new collections from new south
Vietnamese provincial records: Lam Dong and Khanh Hoa .......

LIENHARD, Charles & ASHMOLE, N. Philip. The Psocoptera (Insecta:
Psocodea) of St Helena and Ascension Island (South Atlantic) with a
new record from'SOutR' AMC : STI CLONE RE

LIENHARD, Charles & BAZ, Arturo. Redescription of the genus Marce-
nendius Naväs (Psocodea: ‘Psocoptera’: Amphientomidae) with a
Key to Western Palkearctie AmphiemMOmIOS PEER

GONZALEZ-SOLIS, David & MARIAUX, Jean. An annotated list of the para-
sitic nematodes (Nematoda) of freshwater fishes from Paraguay
deposited in the Museum of Natural History of Geneva ..........

WEBER, Jean-Marc. Food habits of escaped Eurasian otters (Lutra lutra)
in à Suburban environment 10), SWILZCANON PP EERR

HLAVAË, Peter. Claude Besuchet, an eminent swiss coleopterists, 80 years
(6) ER ed de 2.

SENGLET, Antoine. New species in the Zelotes tenuis-group and new of
little known species in other Zelotes groups (Gnaphosidae, Araneae)

TANASEVITCH, Andrei V. Linyphiid spiders (Araneae, Linyphiidae) from
Pakistan and'Indià :7 2e IR RM A RE PR PER EE

Indexed in CURRENT CONTENTS, SCIENCE CITATION INDEX

Pages

401-412

413-421

423-449

451-466

467-484

485-489

491-512

513-559

561-598

PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENÈVE

CATALOGUE DES INVERTEBRES DE LA SUISSE, N9OS 1-17 (1908-1926) ........ série Fr.
(prix des fascicules sur demande)

REVUE DE PALÉOBIOLOGIE -_.............:......... Echange ou par fascicule Fr.

LE RHINOLOPHE (Bulletin du centre d’étude des chauves-souris) ....... par fascicule Fr.

THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND
DISTRIBUTION, WITH KEYS FOR DETERMINATION
at am mn er menu e ve aa ans tongue Fr.

CLASSIFICATION OF THE DIPLOPODA

ne la De manon min © con a mc Ê Une mia see 3 Fr.

LES OISEAUX NICHEURS DU CANTON DE GENÈVE
P. GÉROUDET, C. GUEX & M. MAIRE

RE CR COS 29.2... messe Er.

CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS
CONSERVES DANS LES COLLECTIONS NATIONALES SUISSES,
SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ
A LA CONNAISSANCE DES ECHINODERMES ACTUELS
D. = nm =» aaiu no done di esse mas ee os Fr.

RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE
(COTENTIN-BAIE DE SEINE, FRANCE)
NES EE MAREDA 2. 1991... 2.2... 0... Fr.

GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN:
SHELLS AND RADULAS

LIT TR LES 00 25 2 SA Ni 2 TA LS SSSR Fr.
O. SCHMIDT SPONGE CATALOGUE
RDESQUEYROUX-FAUNDEZ & SM STONE, 190 p., 1992 ...................... . Fr.

ATLAS DE RÉPARTITION DES AMPHIBIENS
ET REPTILES DU CANTON DE GENEVE
EE de ARILEN CN MARNE 28 p,299%..,...........:....1........ Fr.

THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS:
A DOCUMENTED FAUNAL LIST
2 LR veu uuns Fr.

NOTICE SUR LES COLLECTIONS MALACOLOGIQUES
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE
TL nn net dton à Fr.

PROCEEDINGS OF THE XIl!th INTERNATIONAL CONGRESS

OF ARACHNOLOGY, Geneva 1995 (ed. V. MAHNERT), 720 p. (2 vol.), 1996 ...... Fr.

CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE)
(Instrumenta Biodiversitatis X), 1. LOBL, xi + 190 p., 1997...................... Fr.

CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA)
DE LA REGION AFROTROPICALE

({nstrumenta Biodiversitatis W), H. G. DIRICKX, x +187 p., 1998 ................. Fr.

A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE
WEST PALAEARCTIC REGION
(Instrumenta Biodiversitatis WI), S. BOWESTEAD, 203 p., 1999 ................... Er.

THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE
SOKOTRA ARCHIPELAGO
(Instrumenta Biodiversitatis IV), B. SCHATTI & A. DESVOIGNES.

CT CL TITRE NN DR RER RDS Fr.

PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY
(Instrumenta Biodiversitatis V), C. LIENHARD & C. N. SMITHERS.
D QE om dhrnmcmmacmuer Fr.

REVISION DER PALAARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA
THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil)
(Instrumenta Biodiversitatis VI). G. SABELLA., CH. BÜCKLE, V. BRACHAT
D OR las ru eue be uno 2 Fr.

PHYLOGENY, TAXONOMY, AND BIOLOGY OF TEPHRITOID FLIES
(DIPTERA, TEPHRITOIDEA)
Proceedings of the “3rd Tephritoid Taxonomist’s Meeting, Geneva, 19.-24. July 2004”
(Instrumenta Biodiversitatis VII). B. MERZ, vi + 274 p., 2006 Fr.

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Volume 118 - Number 3 - 2011
Revue suisse de Zoologie: Instructions to Authors

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HANDLEY, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). /n: WENZEL, R. L. & TIPTON, V. J.

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