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Full text of "Revue suisse de zoologie."

REVUE SUISSE DE ZOOLOGIE 



TOME 1 18— 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 



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ANNALES 



de la 

SOCIÉTÉ SUISSE DE ZOOLOGIE 
et du 

MUSÉUM D'HISTOIRE NATURELLE 
de la Ville de Genève 



tome 1 1 8 
fascicule 3 
2011 



el GENÈVE SEPTEMBRE 2011 ISSN 0035 - 41 8 X 



REVUE SUISSE DE ZOOLOGIE 



TOME 1 18— 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 



Prix de l'abonnement: 



SUISSE Fr. 225.— UNION POSTALE Fr. 250.— 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 
à la rédaction de la Revue suisse de Zoologie, 
Muséum d'histoire naturelle, C.P. 6434, CH-121 1 Genève 6, Suisse 



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 3 rd 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 maie 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 - 
Médite rranean. 

INTRODUCTION 

Microphorella Becker is one of a small group of gênera, 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 
(Chvâla, 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 

Ail spécimens were collected by the author from coastal sand dunes or estuaries 
in two différent localities in the Tabarka embayment (northwest coast of Tunisia) in the 
spring of 2005 and 2007. Most spécimens 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 accepted29.1 1 .2010 



402 



P. GATT 



Dissected spécimens on which the drawings are based are stored in glycérine 
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 
vvere 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 internai 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 génital rotation and flexion, i.e. as 
they appear in the intact spécimen. 

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

as accessory sclerites of génital fork (sternite 9) 

C female cercus 

Ce maie right cercus 

ds dorsal sclerite of génital fork 

EA ejaculatory apodeme 

EPA epandrium 

fo foramen from segment 8 

HA hypandrium 

hslO female abdominal hemisternite 10 

ht 10 female abdominal hemitergite 10 

PH phallus 

S spermatheca 

sl-s8 abdominal sternites 1-8 

1 1 - 18 abdominal tergites 1-8 

vs ventral sclerite of génital fork (sternite 9) 

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

Spécimens are deposited as indicated by the following acronyms in brackets 
under 'material examined': 

IRSNB Institut Royal des Sciences Naturelles, Bruxelles, Belgium 

MHNG Muséum d'histoire naturelle, Genève 

NMWC National Muséum Wales, Cardiff 

PGS Private collection of P. Gatt, Sliema, Malta 

ZIN Zoological Institute of the Russian Academy of Sciences, St. Petersburg 
ZMHB Muséum fur Naturkunde, Berlin (Zoologisches Muséum der Humboldt 
Universitàt Berlin) 

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



A NEW M ICR OPHOR ELLA FROM TUNISIA 



403 



TAXONOMIC TREATMENT 

Microphorella cassari sp. n. Figs 1-13 

Material examined (13 6 6 14 9 9: Holotype, maie: "TUNISIA: Tabarka, / Oued 
Berkoukech, / dunes, 3 1 .iii.2007 / P. Gatf, "HOLOTYPUS / Microphorella I cassari sp. n. / 
Gatt." (IRSNB). The holotype is preserved in alcohol and is in perfect condition. - Paratypes, 9 
6 6 and 12 9 9 same data as holotype: 1 9 Tunisia, Tabarka: Oued Berkoukech, beach, 13. iv. 
2005, P. Gatt; 2 6 6 , Tunisia, Tabarka, Oued Bouterfess, beach, estuary, 14.iv. 2005, P. Gatt; 1 
6 and 1 9 , Tunisia, Tabarka, Oued Bouterfess, dunes, 31. iii.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 6 6 and 1 9 , same data as holo- 
type; 1 9 , Tunisia, Tabarka, Oued Berkoukech, estuary, 13.iv.2005, P. Gatt (PGS). 

Etymology: This species is 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: A small (1 .3 mm), brownish-grey microtrichose, sexually dimorphic 
species with white, lanceolate macrosetae and infuscated wings. Maie: antennal style 
long, strap-like, acuminate; fore and mid tibia, posteriorly, with long setae; mid fémur 
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 
Maie 

Length. Body 1.3 mm, wing 1.2 mm (spécimen 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, widening 
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 
latéral surface; sensory pit présent. 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 latéral verticals; postocular occipital setae 
uniserial above neck, becoming longer and irregularly multiserial below; several 
longer setae présent 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 






FlGS 1-3 

Microphorella cassari sp. n. (1) S head, viewed from above and in front. (2) S antenna, latéral 
view. (3) 6 , wing. Scale bars, Figs 1 and 3: 0.5 mm, Fig. 2: 0.2 mm. 

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



A NEW M ICR OPHOR ELLA FROM TUNISIA 



405 




Fig.4 

Microphorella cassari sp. n., â, 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 présent 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 ail 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. Ail tarsomeres with short, spine-like 



406 



P. GATT 




Fig.5 

Microphorella cassari sp. n., 6 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 ail 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 fémur 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 fémur 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 
fémur 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., â , 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 basai 
setae, anterior costal margin with a row of spine-like setae, longer and stronger at 
middle of wing; Se parallel to Rj, upturning to C before merging imperceptibly into 
membrane very close to humerai crossvein indistinct; R s originating opposite 



408 



P. GATT 




Fig.7 

Microphorella cassari sp. n., 6 hypopygium, viewed dorsally and from the left. Scale bar 
0.3 mm. Abbreviations explained in the text. 

humerai crossvein; Rj meeting C opposite tip of discal cell; R 2+ 3 sinuous, upcurved at 
its junction with C; R 4+5 sinuous, ending in C at tip of wing; crossvein R-M présent, 
sometimes indistinct; discal cell incompletely separated from second basai cell by 
incomplète crossvein BM-Cu, closed distally by crossvein DM-Cu and emitting 3 
veins to wing margin; base of M 2 complète; crossvein DM-Cu complète; CuA 2 curved, 
Al+CuA 2 absent. A 2 présent. Alula absent. Squama brown, short, with a fringe of long, 
pale hairs. Haltère 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 dise. 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 latéral margins, bare except for a covering of microtrichia. Sternites 2 and 3 
setulose on dise 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 M ICR OPHOR ELLA FROM TUNISIA 



409 




Figs 8-10 

Microphorella cassari sp. n. (8) S phallus and associated structures, latéral view. (9) 9 antenna, 
latéral view. (10) 9 abdomen, latéral 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 pôle 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, vving 1.3 mm (spécimen in alcohol). 

Resembling maie, 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 Unes on each side, appearing as a 
pair of parallel, longitudinal vittae extending from anterior edge of mesoscutum to 
prescutellar dépression. Ail 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, complète 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; génital 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 latéral surfaces, longest at tip. Spermatheca (Figs 10, 
12-13) tubular, réceptacle pigmented; middle part of spermathecal duct cylindrical, 
pigmented and with tracheated surface; minute dorsal and ventral sclerites (part of 
génital fork) posterior to réceptacle. 

Biology: Ail spécimens 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 maie is unique amongst previously described species of Microphorella , both fossil 
and extant. The costal vein of the maie bears several long, spine-like setae on the 
middle portion that also have not been recorded in other species of this genus. Among 



A NEW M ICR OPHOR ELLA FROM TUNISIA 




FlGS 11-13 

Microphorella cassari sp. n. (11) 9 tip of abdomen, dorsal view. (12) 9 sternite 8 with 
(internally) parts of génital fork and spermatheca, dorsal view. (13) 9 spermatheca and parts of 
génital fork, latéral view. Scale bars, Fig. 11: 0.3mm, Figs 12-13: 0.2mm. 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 ( Médite rranean Coastal River Ecosystems) 
Project co-ordinators, in particular Prof. Félicita 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 (All-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 

Chvâla, M. 1988. Revision of Palaearctic Microphoridae (Diptera) 3. Parathalassiinae 
{Parathalassius Mik and Microphorella Becker). Acta Entomologie a 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 région. Revue suisse de Zoologie 110 (4): 

669-684. 

Melander, A. L. 1928 (1927). Diptera, Fam. Empididae. In: Wytsman, P. (éd.). Gênera 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 ). In: 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 

Rajfles Bulletin ofZoology 52(1): 45-58 . 
Sinclair. B. J. 2000. Morphology and terminology of Diptera maie terminalia (pp. 53-74). In: 

Papp. 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 gênera of parathalassiine-like flies from South Africa (Diptera. 

Empodoidea). Bonner zoologische Beitràge 42(2): 187-216. 
Ulrich. H. 2003. How récent are the Empidoidea of Baltic amber? Studia Dipterologica 10: 

321-327. 



Revue suisse de Zoologie 118 (3): 413^21; 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 1 4 , Tao Thien NGUYEN 2 , Trung My PHUNG 3 , Tri LY 1 , 
Wolfgang BÔHME 4 , and Thomas ZIEGLER 5 

1 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.hcmuns@gmail.com 

2 Vietnam National Muséum of Nature, 18 Hoang Quoc Viet, Hanoi, Vietnam. 

3 9A Dong Khoi, Tarn 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 séries consisting of three adult maies 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 its type locality (Chu Yang S in National Park, Dak 
Lak Province, southern Vietnam, 1,600 m a.s.l.). We further add additional 
information on the natural history of R. 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 et al. (2008) 
based on a type séries consisting of three adult maies 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 et al. 2009). During récent 



Manuscript accepted 20.06.201 1 



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 nevv records from southern Vietnam. 

MATERIAL AND METHODS 

Spécimens 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 o 17'-108°42'E; 
600-2000 m a.s.l.) 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 o 02'-12°15'N, 108°57'-109°05'E; approximate 
1600 m a.s.l) in September 2010 (period of 10-15 September). 

Frogs were collected by hand from 19:00-23:00. After taking photographs, 
spécimens were anaesthetized, fixed in 80% ethanol for few hours, and subsequently 
preserved in 70% ethanol. Spécimens finally were deposited in the collections of the 
University of Science, Ho Chi Minh City, Vietnam (US); the Vietnam National 
Muséum of Nature (VNMN), Ha Noi, Vietnam and the Zoologisches Forschungs- 
museum Alexander Koenig (ZFMK), Bonn, Germany. Adults (S â): UNS 00500- 
00508, VNMN 965, ZFMK91 5 17-91522; Adults (9 9): UNS 00509, VNMN 969. 

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 
(H AL): 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 CHU YANGSINENSIS 415 



104° 108° 112 5 




104° 108° 112° 



FlG. 1 

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

(FFL); first toe length (FTL); width of dise on finger or toe: greatest width of terminal 
dise on finger or toe; third finger's dise width (TFPW); length of inner metatarsal 
tubercle (IMTL); length of nuptial pad (NPL); formula of vvebbing followed Glaw and 
Vences (2007). The sex of the spécimens was superficially determined based on the 



416 



D. T. A. TRAN ET AL 



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

RESULTS 

Redescription of Rhacophorus chuyangsinensis Orlov, Nguyen & Ho. 2008 

In the following \ve provide a detailed morphological description of R. chu - 
yangsinensis based on 15 adult maies 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 région flat. wider than internarial distance and as large as or little bit larger 
than width of upper eyelid: tympanum round, not raised above temporal région, 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<III: tips of fingers flat. en- 
larged into round dises with circummarginal grooves: dise of third finger as large as or 
little larger than the tympanum diameter; fingers incompletely webbed. formula 1(1) 
IIi(l)e(0.5) EHi(1.5)e(l) IV(1); narrow. smooth flap of skin présent 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 maies, located on latéral 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: dises of toes round with 
circummarginal grooves. smaller than those of fingers: relative lengths of toes 
I<II<V<in<IV: toes broadly webbed. formula 1(0) Oi(l)e(0.5) IlIid)e(O) 
IVi(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 présent. 

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. dises and webbings whitish-cream: 
the blue color surrounding the black patches on flanks and upper arms is only slightly 
discernible if at ail. 

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 CHU YANGSINENSIS 



417 



Table 1. Measurements (mean ± standard déviation, foilowed by minimum and maximum in 
parenthèses; 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 
spécimens. See the methods for abbreviations. 





Maies (n = 15) 


Females (n = 2) 


SVL 


37.9 ±2.6 (34.8^3.8) 


59.1 ±1.6 (58.0-60.2) 


HL 


14.7 ±0.7 (13.5-16.1) 


21.5 ±0.5 (21.2-21.9) 


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 


3.3 ±0.2 (2.9-3.5) 


4.6 ±0.7 (4.1-5.1) 


ED 


4.9 ±0.4 (4-5.4) 


5.9 ±0.1 (5.9-6.0) 


UEW 


3.8 ±0.6 (2.4-4.6) 


3.3 ±1.7 (2.1-4.5) 


IO 


4.5 ±0.8 (3.8-7. .9) 


8.5 ±3.5 (6.0-11.0) 


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 (2.0-4.2) 


4.1 ±0.9 (3.5^.8) 


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 (11.0-13.4) 


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 (27.3-29.0) 


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 (24.1-24.3) 


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 (2.2-4.3) 


5.6 ±2.3 (4.0-7.2) 


NPL 


3.3 ±0.3 (2.8-3.7) 





spots; thin, light stripes présent 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 vvith large, black patches that vary in size and shape; thèse patches are some- 
times surrounded by bright blue color and are more obvious in females (more détails 
are given in the discussion of the sexual dimorphism); webbings on fingers and toes 
yellow or orange, sometimes with black pattern at base; dises 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 maies (UNS00506-00508; 
ZFMK9 15 17-915 19) reddish brown, with two yellow dorsolateral stripes stretching 
from posterior corner of eye to groin and a médium third yellow stripe extending from 
behind the middle of the back towards cloaca; in such coloured spécimens, also the 
light snout stripes are more distinctly developed, and sometimes a light stripe between 
eyes is présent, forming a triangle on the dorsal surface of head (UNS00506 & 
ZFMK91519) (Fig. 2 E & F). 

Sexual dimorphism. Female size on average is 1 .6 times larger than that of 
maies. The females also differ from maies by their flank pattern, which consists of the 



418 



D. T. A. TRAN ET AL 




FlG. 2 

Différent color patterns of Rhacophorus chuyangsinensis in life. (A-B) female (VNMN965) at 
day and night time. (C-F) maies (C: UNS00500, D: UNS00508, E: ZFMK91518, and F: 
UNS00506). 



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



REDESCRIPTION OF RHACOPHORUS CHU YANGSINENSIS 



419 



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

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

A small rhacophorid species vvith 35.1-44.15 mm SVL in adult maies, and 
58.0-60.2 mm in adult females; body depressed; head wide and flat, approximately as 
long as wide; snout somevvhat 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 tvvo oblique ridges that reach the upper part of the round- 
ed choanas; dises of fingers and toes flat, large, round, with circummarginal grooves; 
dises on fingers larger than toe dises; fingers incompletely vvebbed (formula: 1(1) 
IIi(l)e(0.5) IIIi(1.5)e(l) IV(1)); toes extensively webbed (formula: 1(0) IIi(l)e(0.5) 
IIIi( l)e(0) IVi(0.5)e(0.5) V(0.5)); dermal fringe along the outside of the fourth finger 
and lower arm présent; similar ridge of skin présent 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 élévations 
between 1,320-1,600 m a.s.l. 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 thèse trees were up to 5 m distant from water sources (Fig. 3). On 
22 May 2010, we measured environmental températures 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 différences between the new collections and the 
type séries 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 tvvo of the three maies by 
Orlov et 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 séries (versus only half of finger's length according to Orlov et al. 2008); the 
tympanum diameter was nearly half (0.4-0.5) of the eye diameter in the new séries 
(versus tympanum diameter being 0.6-0.7 of eye diameter according to Orlov et al. 
2008). Based on our new records of R. chuyangsinensis we could also show that adult 
maies have a wider range in snout-vent length than it was noted previously (only 
42.9^4.2 mm after Orlov et al. 2008 versus 35.1^-3.8 mm in the new séries 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 maie R. 
chuyangsinensis are particularly noteworthy. Because thèse unusually patterned maies 
occurred in the same microhabitat and because we did not find significant morpho- 
logical différences 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 différent colour 
morphs. Orlov et al. (2008) could not document maie advertisement calls of R. 
chuyangsinensis during October 2007. Thus, thèse 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 CHU YANGSINENSIS 



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 Académie 
Exchange Service (DAAD). 

REFERENCES 

Glaw, F. & Vences, M. 2007. A field guide to the amphibians & reptiles of Madagascar. Third 
édition. 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) vvith 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. 



Revue suisse de Zoologie 118 (3): 423-449; septembre 201 1 



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

Charles LIENHARD 1 & N. Philip ASHMOLE 2 

1 Muséum d'histoire naturelle, c. p. 6434, CH-121 1 Genève 6, Svvitzerland. 

E-mail: charleslienhard@bluevvin.ch 
2 Kidston Mill, Peebles, EH45 8PH, U.K. 

The Psocoptera (Insecta: Psocodea) of St Helena and Ascension Island 
(South Atlantic) with a new record from South Africa. - Four new 
species are described: Cerobasis atlantica Lienhard sp. n. (Trogiidae) from 
St Helena, Sphaeropsocopsis insidarum Lienhard sp. n. (Sphaeropsocidae) 
from St Helena and Ascension Island, Indiopsocus 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 endémie 
Blaste basilewskyi Badonnel; this could be an example of sympatric spe- 
ciation. Helenatropos abrupta Lienhard, formerly supposed to be a St 
Helena endémie, is for the first time recorded from South Africa and its 
maie is 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 maie of the blind cave- 
dwelling St Helena endémie Sphaeropsocopsis myrrteae Lienhard & 
Ashmole is described for the first time; its génital 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 thèse 
islands. The number of species recorded from St Helena is raised to 23 (6 
endémies), that from Ascension Island to 1 3 (2 endémies). A checklist of the 
27 psocid species recorded from thèse islands is presented and a brief bio- 
geographical analysis is provided. 

Keywords: Trogiidae - Sphaeropsocidae - Psocidae - new species - new 
records - cave fauna - blind psocid - island endémies - 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 2 ) and in their origin as volcanic oceanic islands 
formed near the Mid- Atlantic Ridge, they differ dramatically in their âge: 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 
différent stages in the development of ecosystems in situations so remote that natural 



Manuscript accepted 20.04.201 1 



424 



C. LIENHARD & N. P. ASHMOLE 



colonization by plants and animais is a rare event and évolution 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 thèse 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 resuit of a séries of collecting opportunités in the past two décades. 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 et 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 
endémie 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 
endémie 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 et 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 gênera Cerobasis Kolbe, Sphaeropsocopsis Badonnel and Blaste Kolbe. Thèse 
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 in the 
checklist (see Appendix) but not treated in this paper. We also publish here for the first 
time the St Helena records of Lepinotus inquilinus and Liposcelis entomophila, already 
listed in the above mentioned report (Mendel et al., 2008), and one récent 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, 2000a). By adding a new 
species of the genus Indiopsocus 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 présent the descriptions of the new species and the new 
records, together with the first South African record of Helenatropos abrupîa. Habitus 
figures are given for the most interesting endémies, the subterranean and troglo- 
morphic Troglotroctes ashmoleorum (Ascension Island, Fig. 4a) and Sphaeropsocopsis 
myrîleae (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 Muséum, London, UK; MHNG = Muséum d'histoire 
naturelle, Geneva, Switzerland; NMSE = National Muséum of Scotland, Edinburgh, 
UK; SEHU = Systematic Entomology, Hokkaido University, Sapporo, Japan; UNAM 
= Universidad Nacional Autônoma 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); IO/D = shortest distance between compound eyes divided by ante- 
roposterior diameter of compound eye in dorsal view of head; P1-P4 = articles of 
maxillary palp; T = hindtibia (length); tl, t2, t3 = tarsomeres of hindtarsus (length, 
measured from condyle to condyle); V = width of head capsule on vertex. 

Bibliographical références 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, 3 9 (holotype and two paratypes), mentioned by 
Lienhard (2005a). 

New material: MHNG and BMNH, 8<5 , 119, 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 




FlG. 1 

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

Description of male: Colouration and gênerai morphology as described for 
the female by Lienhard (2005a). Sclerotized metanotal vvinglet-like lobes of maies 
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) thèse 
lobes are exceptionally well-separated medially]. Epiproct simple, paraproct nearly 
triangular (Fig. le), 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. la). Phallosome simple (Fig. lb), 
lacking conspicuous internai sclerotizations, parameres distally bifurcate, their 
posterior part with an anteriorly curved tip. Measurements (6 MNHG 7987, //m): BL 
= 1700; F = 330; T = 436; tl = 130; t2 = 47; t3 = 56. 

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



PSOCOPTERA OF ST HELENA AND ASCENSION 



427 



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

During an ecological research project in the Table Mountain National Park 
numerous spécimens of H. abrupto, were recently collected in South Africa (see 
material mentioned above). The South African females are identical to the St Helena 
spécimens, 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 (Aizoaceae) 
which was brought to St Helena from South Africa in the 19 th 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 séries of stri- 
king autapomorphies (see Lienhard, 2005a: 695). The présence of a forked sensillum 
on P4 in both sexes (see Lienhard, 2005a: fig. 4, 1998: fig. 26i), of a well-developed 
hypandrial brush (Fig. la) and of a simple phallosome with distally bifurcate para- 
meres (Fig. lb) 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 gênera of this family, which 
is not the purpose of this paper; thèse gênera are listed by Lienhard & Smithers (2002) 
and Li Fasheng (2002). 

Cerobasis atlantica Lienhard sp. n. Fig. 2 

Holotype: MHNG, 6, St Helena, Earwig Gully, Prosperous Bay Plain, S15°57.459' 
W5°39.059', ca 290-310m, a gully southeast of the Central Basin, 27.ix.2003, leg. P. & M. 
Ashmole (site PBP4, sample 79). 

Paratype: MHNG, 9 (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.ix.2003, leg. P. & M. Ashmole (site 
PBP3, sample 61). 

DESCRIPTION: Colouration: Body light yellowish white with a small dark brown 
médian patch on pronotum and some tiny brown spots on abdominal tergites, arranged 
in segmentai transversal rows. Compound eyes black, labrum dark brown, flagello - 
mères 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 latéral 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 latéral 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 internai spurs in apical 
half (one hindtibia of the female with 4 internai spurs of normal size and one additional 
short internai spur close to the apical spurs). Pretarsal claws lacking preapical tooth, 
with basai appendix and slightly enlarged membranous pulvillus. 



428 



C. LIENHARD & N. P. ASHMOLE 




FlG.2 

Cerobasis atlantica Lienhard sp. n., maie 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 pariétal gland, (g) 
Spermapore région. 

Hypandrial brush with about 40 acuminate setae. Phallosome as in Fig. 2d, 
lacking strongly sclerotized internai 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 
tvvo stouter setae about in middle and relatively short apical hairs. Région of spenna- 
pore eharacteristic (Fig. 2g). Both spermathecal pariétal 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 (//m): Maie holotype: BL = 1370; FW = 180: F = 300: T = 460: 
tl= 168: t2 = 52: t3 = 60; length of phallosome = 250. - Female allotype: BL = 1510; 
FW = 220: F = 350: T = 560; tl= 180: t2 = 56; t3 = 65. 

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

DISCUSSION: Cerobasis atlantica seems to be endémie 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 endémies 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 présence of 4 terminal spurs on 
hindtibia (3 in C. annulata), the absence of a sclerotized longitudinal internai structure 
near the apex of the paramere, the structure of the spermapore région 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 is 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 internai spurs on hindtibia and with more elongate 
external valvulae and a very eharacteristic spermapore région (see Lienhard, 1984: figs 
7 and 18). 

Cerobasis guestfalica (Kolbe, 1880) 

Materialexamined: BMNH, 1 9 .Ascension Island. Devil's Cauldron. S7°56' W14°19'. 
12.viii.2003. leg. H. Mendel (off Juniper us). 

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 maie from Mexico. 
Cerobasis cf. maya Garcia Aldrete. 1991. - Ashmole & Ashmole. 1997, 2000a. - Lienhard & 
Smithers." 2002. - Lienhard, 2004b. 

Holotype (examined): UNAM, 6 , Mexico. Yucatan Peninsula. Quintana Roo. Cancun, 
2.XÏ.1971 , on dead hanging fronds of coconut palm, leg. A. N. Garcia Aldrete. 

Newmaterial: MHNG. 1 S , Ascension Island, Command Hill. 17.-21 .iii. 1990, leg. N. 
P. & M. J. Ashmole, trapping on lava (grass cover low, small quantities of several différent 



430 



C. LIENHARD & N. P. ASHMOLE 




FlG.3 

Cerobasis maya Garcia Aldrete, maie holotype (a), maies from St Helena (b-g): (a) Phallosome, 
apical part, (b) Tip of left lacinia. (c) Tip of right lacinia (same spécimen), (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, 26, Ascension Island, South Gannet Flow, 23.- 
27.iii.1990, leg. N. P. & M. J. Ashmole, trapping off-lava (thistles, no moss or lichen) (sample 
0818 Asc). - BMNH, 1 â , 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 



431 



Description of male from Ascension Island: Colouration: Not very well 
preserved. Body yellovvish, frons medially with an approximately anchor-shaped 
brovvn patch, sometimes subdivided into smaller spots, several other brovvn spots on 
head, thorax and abdomen, laterally often fused to form larger patches or bands, 
vvinglets hyaline or with some brown pigment (as figured by Garcia Aldrete, 1991: 
fig. 2). Compound eyes black, basai flagellomeres apically brown (distal half of 
antennae lost in ail spécimens 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 latéral corners of metanotum, therefore pilosity covering almost ail 
the metanotum, except for 1/6 of its dorsal surface near latéral margin (Note: in many 
other Cerobasis species latéral parts of metanotum extensively covered by winglets 
and each latéral 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 internai spurs in apical half (3 internai spurs in 
one of 8 hindtibiae examined). Pretarsal claws lacking preapical tooth, with basai 
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 différent in closed (Fig. 3f) and open 
(Fig. 3g) position of the phallosome (see Discussion below). 

MEASUREMENTS (//m): Male holotype (data from Garcia Aldrete, 1991, except 
for length of phallosome): FW = 113; F = 287; T = 486; tl= 184; t2 = 50; t3 = 58; 
length of phallosome = 180. - Male from Ascension Island (MHNG 7 146): BL = 1260; 
FW = 120; F = 280; T = 460; tl= 172; t2 = 47; t3 = 57; length of phallosome = 165. 

DISCUSSION: The above mentioned material collected by Philip and Myrtle 
Ashmole has already been mentioned by thèse authors as Cerobasis cf. maya (Ashmole 
& Ashmole, 1997, 2000a; see also Lienhard & Smithers, 2002 and Lienhard, 2004b). 
Ail maies from Ascension Island are so similar to the only previously known spécimen 
of C. maya, its holotype, that there is no reason to consider them as belonging to a 
différent species or subspecies in spite of some slight différences concerning phallo - 
some morphology. The whole phallosome of the holotype is figured by Garcia Aldrete 
(1991: fig. 4), détails 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 maies 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 thèse parts in Fig. 3f is rather similar to 
that observed in the slide of the holotype (Fig. 3a). A careful analysis of thèse stuctures 



432 



C. LIENHARD & N. P. ASHMOLE 



in ail available maies showed that the différences between Fig. 3a and Fig. 3f are 
largely due to slightly différent positions after slide-mounting. The only significant 
différence between the holotype and the maies from Ascension Island is the présence, 
in the holotype, of a small field of scale-like sculpture on the parameres near the base 
of the internai process (Fig. 3a and Garcia Aldrete, 1991 : fig. 4); this sculpture is absent 
or only very weakly developed in the maies from Ascension Island (Fig. 3f, g). 
Hovvever, compared to usual interspecific différences in phallosome morphology in the 
genus Cerobasis, this extremely slight différence does not justify any taxonomic 
décision about spécifie or subspecific séparation of the Ascension population, 
especially in view of the low numbers of individuals available at présent. 

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, 2 9 , St Helena, Woodcot. S15°57.2' W5°42.7\ ca 489m, 
18.ii.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). 

LlPOSCELIDIDAE 

Liposcelis entomophila (Enderlein, 1907) 

Material examined: MHNG, 29, St Helena. Rupert's Battery Cave, ca 50m. 
23.ii.2006, leg. P. & M. Ashmole (sample 4444) and 4.xi.2006, leg. E. Thorpe (sample 
2821). 

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 et 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 
endémie by Mendel et 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, le?, 219 (most of them heavily damaged). 
Ascension Island, South Gannet Hill, 1 7. -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 T. ashmoleorum with the habitus of this 
typical Liposcelis species makes évident the troglomorphic habitus of the former (i. 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 corne to the 
surface, probably at night (Asmole & Ashmole, 2000b). 

T. ashmoleorum is 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, 9, St Helena, Rupert's Battery Cave, ca 50m, 25.xi-10.xii.2003, 
leg. N. P. & M. J. Ashmole, modified pitfall trap ("boot trap") (sample 1835). 

Paratypes: BMNH, 1 9 , 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, 1 9 , 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 (maie unknown): Head light brown, rest of body 
whitish brown to yellowish. Hindwings absent, forewings lost in ail spécimens, 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 aréoles (diameter of tubercles slightly smaller than diameter of the 
alveoli of the small vertical hairs). Antenna with 15 articles (mostly damaged, flagellar 
sensé 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 latéral one differentiated as a thick short sensé club (Fig. 5i). 
Lacinial tip bifurcate, inner tine shorter than outer tine, the latter with two small 
secondary denticles on its inner side (similar to S. myrtleae, see Lienhard & Ashmole, 
1999: fig. 5). Sculpture of thoracic tergites similar to that of vertex but aréoles absent 
or indistinct. Mesonotum not subdivided into lobes (Fig. 5d). Legs slender but not 
particularly long (index T/V = 1.2), no coxal organ présent, 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. 5e). Gonapophyses typical for 
the family (see Mockford, 2009), apex of external valvula broadly rounded, not bilobed 
(Fig. 5g). Région of spermapore with a weakly developed field of small microtrichia 
(Fig. 5f), spermathecal duct and sac not observed. Measurements (holotype, //m): BL 
= 880; V = 260; F+tr = 285; T = 315; tl= 114; t2 = 32; t3 = 45. 

ETYMOLOGY: The spécifie epithet, a féminine 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 




Fig. 5 

Sphaeropsocopsis insularum Lienhard sp. n., female holotype (c-d, g-i), 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 spécimen and différent position), (d) 
Dorsal view of mesothorax (with insertion points of forewings) and anterior part of wingless 
metathorax. (e) Subgenital plate, (f) Spermapore région, (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 spécimen 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 it consists of large irregularly lobate 
tubercles which are not arranged into aréoles (Badonnel, 1963). Unfortunately fore- 
wings are broken and lost in ail available spécimens of S. insularum; however, the 
insertion point of the forewing (Fig. 5d) 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 présence 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 maie genitalia it is impossible to décide if S. insularum is more closely related to 
this African species, as is S. myrtleae, the second St Helena species of the genus (see 
below), or to S. microps, which is only known from natural edaphic habitats in Chile 
(see Badonnel, 1963, 1967). S. reisi has eyes with 9 ommatidia (see Badonnel, 1971). 
However, eye réduction 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, 9 , St Helena, Rupert's Battery Cave, 13-17.iii.1995, leg. N. P. & M. 
J. Ashmole, modified pitfall trap ("boot trap") (sample 680 SH). 

New material: MHNG, 16 1 9 , and BMNH, 1 9 , St Helena, Rupert's Battery Cave, ca 
50m, 25.xi-10.xii.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 latéral sensé 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 (6 MHNG 7626, pm): BL = 1220; V = 250; F+tr = 304; 
T = 385; tl= 130; t2 = 39; t3 = 56. 

DISCUSSION: Based on two of the three females known at présent, the figure of 
the head (Fig. 6b) could be completed (see Lienhard & Ashmole, 1999: fig. 1, lacking 
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 



437 




Sphaeropsocopsis myrtleae Lienhard & Ashmole: (a) Habitus of female, dorsal view, antennae 
incomplète (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, maie. 



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 présence of an 
elongate fusiform P4 novv confirms this assignment (P4 subcylindrical in Badonnelia; 
see Badonnel. 1963). The newly discovered maie also confirais the initial generic 
assignment and allovvs a better understanding of the origin of this island endémie. The 
morphology of the phallosome of S. myrtleae is very similar to that of the African 
species S. reisi Badonnel, knovvn 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 présent. Even in forms with reduced compound eyes, as 
certain species of Liposcelis, at least two ommatidia are always présent (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 et 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) 

Materialexamined: BMNH. 1 9 . St Helena, Cuckhold's Point. S15°58' W5°42'. 771m. 
xii.2005-i.2006. leg. H. Mendel (Malaise trap). - BMNH. 1 9 (damaged, lacking abdomen), 
Ascension Island, Grazing Valley, S7°57' W14°21'. viii.2003. leg. H. Mendel (pitfall trap). - 
BMNH. 1 9 .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, Israël, Portugal and 
Tunisia). 

Peripsocidae 

Peripsocus leleupi Badonnel. 1976 

Materialexamined: BMNH and MHNG. 59. Ascension Island. Devil's Cauldron, 
S7°56' W14°19\ 12.viii.2003, leg. H. Mendel (off Juniperus). 

Comment: This species was previously considered as a St Helena endémie 
(Ashmole & Ashmole, 2000a); 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, 69, Ascension Island, Devil's Cauldron, 
S7°56' W14°19', 12.viii.2003, leg. H. Mendel (off Juniper as). 

Comment: This widespread pan-tropical waif (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, 6 , St Helena, Lot Summit, 454m, off St Helena rosemary Phylica 
polifolia, 29L2006, leg. P. & M. Ashmole (sample 4165). 

Paratypes: BMNH 1 6 , MHNG 1 9 (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 génital morphology. 
Hypandrium with a médian 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 région as in Fig. 7e. 

Measurements: Maie holotype: BL = 2.2 mm; IO/D = 2.1; FW = 2.5 mm; F = 
440 pm; T = 830 pm; tl= 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; tl= 210 pm; t2 = 140 pm. 

Etymology: The spécifie epithet refers to the island of St Helena, a British 
overseas territory in the South Atlantic Océan, 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 médian 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 différences 
concerning the sclerotized area surrounding the spermapore (see Fig. 7i 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 endémie gumwoods 
Commidendrum spp., although it has also been found on other plants (Badonnel, 1976 
and personal observations). The discovery of Blaste helenae on the endémie St Helena 
rosemary Phylica polifolia suggests a niche séparation between the two species. B. 



440 



C. LIENHARD & N. P. ASHMOLE 




Fig. 7 

Blaste helenae Lienhard sp. n. (a-d, maie holotype; e-f, female allotype): (a) Forewing. (b) 
Hindwing. (c) Phallosome. (d) Hypandrium, ventral view. (e) Spermapore région, (f) Subgenital 
plate. - Blaste basilewkyi Badonnel (g-i): (g) Hypandrium, ventral view. (h) Phallosome. (i) 
Spermapore région. 



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 érosion, which 
forms a striking feature of the landscape of Sandy Bay. Its nearly vertical sides are 
almost devoid of végétation but the summit - less than one tenth of a hectare in extent 
- provides a refuge for a few spécimens 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 détermine 
whether B. helenae is also présent on any of the other surviving spécimens. 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 resuit of sympatric speciation, possibly resulting from adap- 
tation to life on différent endémie plants, rather than of double invasion. 

Indiopsocus mendeli Lienhard sp. n. Figs 8-9 

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

Paratypes: BMNH, MHNG and SEHU, 19c? , 109 (one of them allotype, MHNG 
8018), same data as for holotype. - BMNH and MHNG, 11c?, 89, Ascension Island, Mt Red 
Hill, S7°58' W14°21', 14.viii.2003, leg. H. Mendel (off Tecoma stans). - BMNH, 1 6 , Ascension 
Island, Mt Red Hill, S7°58' W14°21', 14.viii.2003, leg. H. Mendel. - BMNH, 1 9 , Ascension 
Island, Devil's Ashpit, S7°57' W14°13\ 2.viii.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 médium brown. Maxillary palp with PI and P2 very light 
brown, P3 and P4 darker brown, apical half of P4 blackish brown. Forewing pattern 
slightly sexually dimorphic, with more extensive brown markings in female (Fig. 8f) 
than in maie (Fig. 8a), especially in basai half of the wing. Abdomen whitish, with 
some red-brown hypodermal pigment, in particular laterally, terminalia dark brown. In 
maie, 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 maie, distinctly 
smaller in female (see IO/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. 

Maie terminalia (Fig. 9): Posterior margin of clunium medially slightly 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 latéral 
protubérance, a pointed apical process and a small setose protubérance basally of the 
latter; trichobothria forming an arched, posteriorly open sensé cushion (Fig. 9a, d). 
Medio-distal protubérance 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 latéral 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, basai sclerotization 



442 



C. LIENHARD & N. P. ASHMOLE 




Fig.8 

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

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



PSOCOPTERA OF ST HELENA AND ASCENSION 



443 





Fig.9 

lndiopsocus mendeli Lienhard sp. n.. maie: (a) Abdominal apex, latéral view ( pilosity not shown. 
except for paraproctal trichobothria). (b) Phallosome. (c) Hypandrium. posterior view islightlv 
squashed). (d) Clunium. epiproct and paraprocts (dorsal view. pilosity partially omirted. para- 
procts in différent position). 



444 



C. LIENHARD & N. P. ASHMOLE 



MEASUREMENTS: Maie holotype: BL = 2.0 mm; IO/D = 0.9; FW = 2.75 mm; F 
= 590//m;T = 1200 //m; tl= 436 //m; t2 = 143 /<m. - Female allotype: BL = 2.1 mm; 
IO/D = 1.7; FW = 2.95 mm; F = 590 //m; T = 1240 pim; tl= 414 //m; t2 = 144 //m. 

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

DISCUSSION: The new species is closely related to Indiopsocus dentatus 
(Thornton & Woo, 1973), which is only known from the Galapagos Islands (Thornton 
& Woo, 1973; Lienhard & Smithers, 2002). The forewing pattern of the female of 
/. dentatus (see Thornton & Woo, 1973: fig. 77) is very similar to that figured here for 
the maie of /. mendeli (Fig. 8a), while in the female of the latter some additional dark 
markings are usually visible in the basai half of the wing (Fig. 8f). Female genitalic 
characters of both species are very similar (see Thornton & Woo, 1973: figs 78, 79; the 
spermapore région of /. dentatus is not known). However, thèse species are easy to dis- 
tinguish by the différent 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 is 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 gênerai distri- 
bution of the non-endemic species can be found in Lienhard & Smithers (2002) and 
more récent 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- 
falica (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 endémie 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 
thèse South Atlantic islands by natural dispersai: Stenocaecilius caboverdensis (St 
Helena, Ascension), Ectopsocus strauchi (St Helena, Ascension), Trichopsocus clarus 



PSOCOPTERA OF ST HELENA AND ASCENSION 



445 



(St Helena), Myopsocus eaîoni (St Helena). The case of the Mexican species Cerobasis 
maya, at présent only known from Ascension Island and the Yucatan Peninsula, is more 
puzzling, in view of the unfavourable conditions for trans-Atlantic dispersai of insects 
from west to east (Ashmole & Ashmole, 1997); anthropogenic distribution due to intro- 
duced plants or phoretic dispersai 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 dispersai or 
human-assisted transfer from one island to the other. 

The following six species can be considered as St Helena endémies (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 
endémie 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 endémies (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 endémie sister-species of unknown origin, Blaste 
basilewskyi and B. helenae, is here tentatively interpreted as a resuit of sympatric 
speciation (see description of B. helenae, above). 

Two species can be considered as endémies of Ascension Island, Troglotroctes 
ashmoleorum and Indiopsocus mendeli Lienhard sp. n. The latter is closely related to 
/. 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 /. 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, Fig. 4a) and Sphaeropsocopsis myrtleae (St Helena, Fig. 6a) are among the 
most interesting endémie arthropods of thèse South Atlantic islands, being the only 
known members of the suborder Troctomorpha with clear morphological adaptations 
to subterranean life. The adaptations presumably evolved after arrivai of the ancestral 
forms respectively on Ascension Island and St Helena. The case of Troglotroctes is 
especially interesting in view of the relatively récent 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 évo- 
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 pièce of évidence 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 stérile. Nearer the surface, the formation of soil 
and growth of végétation leads to faunal succession (Ashmole et 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 spécimens 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 
Terri tories Environment Programme. Grateful thanks are also due to ail those people 
who helped to make research on the islands so productive and enjoyable. 

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Online: http://www.ville-ge.ch/mhng/psocoptera/page/ps-coun.htm 
Lienhard. C. 2005a. Two new soil-dwelling psocids (Psocoptera: Trogiidae and Pachytroctidae) 
from the islands of St Helena and Madagascar. Revue suisse de Zoologie 112(3): 
689-701. 

Lienhard, C. 2005b. Additions and Corrections (Part 4) to Lienhard & Smithers. 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 7: 1-16. For 

online version see: http://www.psocodea.org/ 
Lienhard, C. 2006. Additions and Corrections (Part 5) to Lienhard & Smithers. 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 8: 1-18. For 

online version see: http://www.psocodea.org/ 
Lienhard. C. 2007. Additions and Corrections (Part 6) to Lienhard & Smithers, 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 9: 1-17. For 

online version see: http://www.psocodea.org/ 
Lienhard. C. 2008. Additions and Corrections (Part 7) to Lienhard & Smithers, 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 10: 1-18. For 

online version see: http://www.psocodea.org/ 
Lienhard, C. 2009. Additions and Corrections (Part 8) to Lienhard & Smithers. 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 11: 2-16. For 

online version see: http://www.psocodea.org/ 



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C. LIENHARD & N. P. ASHMOLE 



Lienhard, C. 2010. Additions and Corrections (Part 9) to Lienhard & Smithers, 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 12: 1-18. For 

online version see: http://www.psocodea.org/ 
Lienhard, C. 2011. Additions and Corrections (Part 10) to Lienhard & Smithers, 2002: 

"Psocoptera (Insecta) - World Catalogue and Bibliography". Psocid News 13: 1-18. For 

online version see: http://www.psocodea.org/ 
Lienhard, C. & Ashmole, N. P. 1999. Sphaeropsocopsis myrtleae sp. n., a blind subterranean 

psocid from St Helena (Psocoptera: Sphaeropsocidae). Revue suisse de Zoologie 106(4): 

905-912. 

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. A 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 endémie) 

St Helena 

Trogiomorpha 

Trogiidae Cerobasis annulata (Hagen, 1865) 

*Cerobasis atlantica Lienhard sp. n. 
Cerobasis guestfalica (Kolbe, 1880) 

Helenatropos abrupta Lienhard, 2005 
Lepinotus inquilinus Heyden, 1850 

Psyllipsocidae 

Psyllipsocus ramburii Selys- 
Longchamps, 1872 

Troctomorpha 

Liposcelididae Liposcelis bostrychophila Badonnel, 
1931 

Liposcelis entomophila (Enderlein, 
1907) 

Liposcelis spec. (species group A) 
Liposcelis spec. (species group C) 



Sphaeropsocidae Sphaeropsocopsis insularum Lienhard 
sp. n. 

* Sphaeropsocopsis myrtleae Lienh. 
& Ashm., 1999 



Ascension Island 



Cerobasis guestfalica 

Cerobasis maya Garcia Aldrete, 1991 



Psocathropos lachlani Ribaga, 1899 
Psyllipsocus ramburii 



Liposcelis bostrychophila 



Liposcelis spec. (species group C) 
*Troglotroctes ashmoleorum Lienh. 
1996 

Sphaeropsocopsis insularum 



PSOCOPTERA OF ST HELENA AND ASCENSION 



449 



PSOCOMORPHA 

Caeciliusidae 



Ectopsocidae 
Peripsocidae 



Trichopsocidae 
Psocidae 



*Stenocaecilius benoiti (Badonnel, 
1976) 

Stenocaecilius caboverdensis 
(Meinander, 1966) 

Ectopsocus briggsi Me Lac h 1 an, 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. 



Stenocaecilius c aboverdensis 



Ectops 



strauchi 



Myopsocidae Myopsocus eatoni McLachlan, 1880 



Peripsocus leleupi 
Peripsocus pauliani 



Indiopsocus mendeli Lienhard sp. n. 



Revue suisse de Zoologie 118 (3): 451-466; septembre 201 1 



Redescription of the genus Marcenendius Navâs 
(Psocodea: 'Psocoptera': Amphientomidae) 
with a key to western Palaearctic amphientomids 

Charles LIENHARD 1 & Arturo BAZ 2 

1 Muséum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. 

E-mail: charleslienhard@bluewin.ch 
2 Universidad de Alcalâ de Henares, Departamento de Zoologia y Antropologia 

Fisica, E-28871 Alcalâ de Henares (Madrid), Spain. 

Redescription of the genus Marcenendius Navâs (Psocodea: 'Psocop- 
tera': Amphientomidae) with a key to western Palaearctic amphien- 
tomids. - Based on recently collected spécimens from continental Spain and 
Mallorca island the type species of the genus Marcenendius Navâs, 1913, 
M. nostras Navâs, 1913, is redescribed and tentatively synonymized with 
the second known species of this genus, M. illustris Navâs, 1923. Ail thèse 
taxa were considered as enigmatic since their original description, almost 
one hundred years ago. Marcenendius is redefined to contain also the 
Macaronesian species M. fortunatus (Navâs, 1917) comb. nov. and the 
African species M. angolensis (Badonnel, 1955) comb. nov., both formerly 
assigned to Nephax Pearman, 1935. The diagnosis of the latter genus is 
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 présence 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 adhèrent dust particles. 

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

INTRODUCTION 

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



Manuscript accepted03.06.201 1 



452 



C. LIENHARD & A. BAZ 



Psocoptera recorded by Navâs. Due to the very superficial original descriptions and to 
the lack of new material thèse taxa were usually considered as nomina dubia and there- 
fore not keyed in comprehensive vvorks by subséquent 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 Navâs 1 original records 
(Acon Remâcha, 1980; Baz, 1989, 2007). 

Pearman (1935) described a second genus of Amphientomidae from Palestine 
(présent day Israël), Nephax Pearman, the type-species of which, N. sofadanus 
Pearman, vvas redescribed by Lienhard (1988). A second European species, initially 
described in the family Lepidopsocidae from the Canary islands (Tenerife) by Navâs 
(1917), vvas 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 posîalatus 
Lienhard, macropterous but morphologically closely related to the type species, vvas 
recently described from the United Arab Emirates (Lienhard, 2009). Some similarities 
betvveen 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 différences betvveen the Macaro- 
nesian species N.fortunatus (Navâs) and the two closely related species N. sofadanus 
and N. posîalatus suggested that it might be justified assigning them to two différent 
gênera. 

The récent discovery of a fully winged amphientomid species in the southeast 
of the Iberian peninsula and on the island of Mallorca enables us to résolve 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 
Navâs, M. illustris, is here tentatively considered as a junior synonym of M. nostras. 
The Macaronesian species Nephax fortunatus is 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 is 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 
Alcalâ 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 betvveen 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 latéral view; P2 = 
second article of maxillary palp; T = hindtibia (length); tl , t2, t3 = tarsomeres of hind- 
tarsus (length, measured from condyle to condyle). Abbreviations of wing veins are 
used according to Yoshizavva (2005). 



REDESCRIPriON OF MARCENENDIUS NAVÂS 



453 



TAXONOMIC TREATMENT 
Marcenendius Navâs, 1913 

Marcenendius Navàs, 1913: 334. Type species (by original désignation): Marcenendius nostras 
Navâs, 1913: 334. 

REVISED DIAGNOSIS: Belonging to Amphientomidae. Forewings densely 
covered with scales (Figs 1, 5c). Médian ocellus close to postclypeus, smaller than 
latéral ocelli, always présent in fully-winged forms, occasionally strongly reduced or 
absent in brachypterous forms; latéral ocelli widely separated, close to compound eyes. 
Antenna with 14 articles (i. e. 12 flagellomeres), rarely the tvvo 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 internai face of labrum and running parallel to each other in about 
1/3 of its width from latéral margin (Fig. 2e). Phallosome Y-shaped, with a slightly 
sclerotized longitudinal internai zone on each side in apical half (Fig. 3d). Internai 
T-shaped sclerite of female subgenital plate présent 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 Navâs, 1913; 
Marcenendius fortunatus (Navâs, 1917) comb. nov. from Nephax; Marcenendius 
angolensis (Badonnel, 1955) comb. nov. from Nephax. 

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

The venation of forewing and hindwing figured by Navâs (1913, 1923) 
corresponds to macropterous individuals and is very similar to that observed in the new 
material, except for the présence of only one anal vein in the figure given by Navâs 
(1913) for M. nostras. We interpret this as an inaccuracy of Navâs' figure (in Amphi- 
entomidae two anal veins are always présent; see Smithers, 1972). The shape of the 
apex of forewing and hindwing seems to be somewhat variable in Marcenendius, 
broadly rounded (Navâs, 1913: hindwing; Navas, 1923: forewing) or slightly acu - 
minate (Navâs, 1913: forewing; Navâs, 1923: hindwing). The wings of the new 
material are about intermediate in shape (Fig. 2a, b). In Navâs' 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 Al is usually very faint and curved 
towards CuP near vving margin. meeting the latter on the margin (nodulus); this 
situation could be confirmed on several slide-mounted forevvings (Fig. 2a). However, 
at low magnification Al seems to end in the membrane, almost parallel to CuP. Thus, 
the absence of a nodulus in Navâs' figures is probably due to inaccurate drawing (i. e. 
incorrect extrapolation of the faint distal part of Al). 

The présence in Spain (Alicante, Almeria, Malaga and Mallorca) of an am- 
phientomid species not belonging to the genus Nephax and shovving some characters 
mentioned by Navâs (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. no stras. Marcenendius is well-defined by a striking 
autapomorphy of nymphs, the présence of "corkscrew" hairs on dorsal side of thorax 
and abdomen (Fig. 6b), a kind of setae previously unknown in Psocoptera. Thèse hairs 
were présent in the nymphs from Alicante, Almeria and Mallorca (M. nostras) and also 
in nymphs from différent localities on Canary islands which belong to a second species 
of Marcenendius, M. forîunatus (Navâs) comb. nov. (see also discussion of this 
species. below). 

Meinander (1973) already considered the African species Nephax angolensis 
Badonnel as "evidently congeneric" with M. forîunatus (Navâs) when redescribing the 
latter as Nephax forîunatus. Because there is 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 lacinial tip. 

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

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

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

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

Material examined: Spain (Alicante): MHNG, 1 9 , Xixona, Penas Roset, under stones 
in a ravine (dry river bed). 15.vi.2003, leg. V. M. Ortuno: MHNG. 1 nymph. same locality, under 
stones. 23.V.2004. leg. V. M. Ortuno; UAH, 9 nymphs, same locality, under stones, 29.X.2009. 
leg. V. M. Ortuno. - Spain (Almeria): UAH, 19.5 nymphs. Maria. Umbria de la Virgen, under 
stones. 17.xii.2002. leg. V. M. Ortuno. - Spain (Malaga): UAH, le?, Velez-Malaga, Finca La 
Alegria. on flight interception trap. 15.X.2008, leg J. M. Vela. - Spain (Mallorca island): MHNG, 
3<5 , 1 9 . 2 nymphs, Calviâ, cave "Avenc de sa Finestreta". about 8m from the entrance inside the 
cave, on the wall, l.v.2010, leg. M. Vadell: UAH. 1 nymph, Calviâ, cave "Avenc des Eriço", 
about 15m from the entrance inside the cave, 27.iv.2007, leg. M. Vadell; UAH, 8 nymphs. 
Calviâ. cave "Cova de Na Boira", about 12m from the entrance inside the cave, 24.ii.2007, leg. 
M. Vadell: UAH, 7 nymphs, Calviâ, cave "Cova de s'Estora". about 7m from the entrance inside 
the cave. 28.iii.2010, leg. M. Vadell. 

REDESCRIPTION: Colouraîion: Body yellowish to brown, spécimens 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 NAVÂS 



455 




Fie. 1 

Marcenendius nostras Navâs, 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 yellovv; basai two thirds of labrum 
brown, apical part yellowish vvhite. Thorax laterally with a longitudinal zone of reddish 
brown hypodermal pigment, legs yellowish to médium brown, no transversal annu- 
lation visible on tibiae. Forewing membrane tinged with brown in basai 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 spécimens. 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 
maie 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 (IO/D 1 .9-2,2). Tip 
of lacinia as in Figs 2c and 3e, inner tine relatively close to lacinial apex (i. e. outer tine 
not much longer than its basai 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. 49i, j), Pearman's organ of hindcoxa incomplète 
(tympanum présent, 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 latéral 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 région 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 
9 ) or 9 (Mallorca 9 ) denticles, some of them minute; canal of spermatophore bent but 
not S-shaped (Fig. 2f). - Maie terminalia: Epiproct, paraproct and hypandrium simple, 
the latter comprising two sclerotized sternites. Phallosome as in Fig. 3d. 

Measurements: Female (9 Alicante/ 9 Mallorca): BL = 2.7/3.2 mm; FW = 
2.8/3.2 mm; HW = 2.3/2.6 mm; F = 710/775 pim; T = 1325/1370 //m; tl = 690/755 //m; 
t2 = 84/103 ]tm\ t3 = 130/132 //m; IO/D = 1 .9/2.2. - Maie (Mallorca S MHNG 8060): 
BL = 2.9 mm; FW = 3.3 mm; HW = 2.7 mm; F = 770 pim; T = 1380 pim: tl = 760 //m; 
t2= 100 /*m; t3= 133 //m; IO/D = 2.1. 

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

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



REDESCRIPTION OF MARCENENDIUS NAVÂS 



457 




Fig. 2 

Marcenendius nostras Navâs, from Alicante, female: (a) Forevving (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 région, (h) Gonapophyses. 



458 



C. LIENHARD & A. BAZ 




Fig. 3 

Marcenendius nostras Navâs. from Mallorca. female (a-c) and maie (d-e): (a) Hindmargin of 
subgenital plate with rudiment of T-shaped sclerite and dorsolateral sclerotized areas (interrupted 
lines). pilosit) not shown. (b) Gonapophyses. (c) Spermapore région, (d) Phallosome. (e) 
LaciniaJ tip. 

diagnostic characters separating M. nostras from M. illustris are indicated by Navâs 
(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 spécimens are somewhat darker and larger than the spécimens from 
continental Spain). \ve see no reason to think that Navâs' spécimen from Tarragona 
might represent another species of this genus. Thus. \ve tentatively propose the above 
mentioned synooymy. The species has been found under stones in open places 
(Alicante and Almeria spécimens) but also in caves, relatively near to their entrance 
(Mallorca spécimens). Nymphal camouflage has not been directly observed in this 
species. The présence of corkscrew hairs (see genus discussion) suggests a similar 
biology as described below for M. fortunatus. 



REDESCRIPTION OF MARCENENDIUS NAVÂS 



459 




FlG.4 

Marcenendius nostras Navâs. map of distribution. 

Marcenendius fortunatus (Navâs. 1917) comb. nov. Figs 6a-d; 5c. d 

Perientomum fortunatum Navâs, 1917: 20. 

Nephax fortunatus (Navâs, 1913): Meinander. 1973: 143 (redescription based on holotype. maie, 
figs); Lienhard, 1988: 369 (further description, maie and female. figs): Lienhard. 1998: 
141 (diagnosis, figs). 

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

DISCUSSION: Based on gênerai morphology of adults and nymphs. this brachy - 
pterous species is closely related to M. nostras (see key, below). Macropterous 
spécimens of M. fortunatus have never been observed, but the most significant mor- 
phological différence between this species and M. nostras is the strong réduction, 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 végétation, often also in stone vvalls 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 Navâs: (a) Biotope (dry river bed) of the Alicante spécimens 
(Penas Roset). (b) General view of Penas Roset. - Marcenendius fortunatus (Navâs): (c) Female 
from Fuerte ventura island (Canary Islands), near Lajares, 15. xi. 1998, habitus in dorsolateral 
view, body length 2.5 mm. (d) Biotope of this spécimen (stone wall built by humans). 
Photographs: V. M. Ortuno (a, b), C. Lienhard (c, d). 



REDESCRIPTION OF MARC EN EN Dl US NAVÂS 



461 




Fig.6 

Marc enendius for tunatus (Navâs), 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 présence of corkscrew hairs on thèse parts 



462 



C. LIENHARD & A. BAZ 



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

Nephax Pearman, 1935 

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

REVISED DIAGNOSIS: Belonging to Amphientomidae. Forewings densely 
covered vvith scales. Médian ocellus absent (even in fully-winged forms); latéral ocelli 
widely separated, close to compound eyes. Antenna with 13 articles (i. e. 11 flagel- 
lomeres). P2 with spur sensillum on inner side and some external macrochaetae in 
addition to the simple and relatively short gênerai 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 
internai sclerotizations. Internai 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. posta- 
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 is 
évident at présent. 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 internai 
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 différences in forewing venation between the macropterous species 
N. postalatus and the brachypterous species N. sofadanus are probably related to wing 
réduction; exactly the same différences 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 cross vein and the 
stigmapophysis is situated distally of the Rl-Rs 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 Rl-Rs bifurcation. We consider the striking similarity in forewing venation 



REDESCRIPTION OF MA RCENENDl U S NAVÂS 



463 



between thèse brachypterous forms not as a synapomorphy but as a case of conver- 
gence due to vving réduction. 

Nephax sofadanus Pearman, 1935 Fig. 6e-h 

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

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

DISTRIBUTION: N. sofadanus is an eastern Mediterranean species knovvn from 
the following countries. Israël (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 Crète 
and Dodecanese islands (Rhodes and Karpathos). Croatia: several localities near Split 
(leg. T. Radja, material in the MHNG). 

DISCUSSION: Ail spécimens of N. sofadanus knovvn at présent are brachypter- 
ous, they usually live under stones in open places incompletely covered by xerophytic 
shrub végétation (e. g. Greek "phrygana" végétation). Adults are rather well camou- 
flaged by their mottled vving pattern due to scales, vvhile nymphs are alvvays "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 
présent (see discussion of Marcenendius fortunatus, above). 

Nephax postalatus Lienhard, 2009 

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

Distribution: United Arab Emirates: Hatta and Wadi Maidaq (Lienhard, 

2009). 

DISCUSSION: No détails are knovvn about the biology of this macropterous 
species, because ail available spécimens vvere 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 (i. e. 11 flagellomeres). Médian 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 
gênerai 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 

Nephax Pearman 2 

Antenna with 14 articles (i. e. 12 flagellomeres), rarely the two apical- 
most flagellomeres fused. Médian ocellus usually présent, 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 Navâs 3 

2 Both sexes macropterous, hindwing well-developed, with complète 
venation, ail 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 Rl-Rs bifurcation. Head capsule 
slightly flattened (i. e. H/L>2). Spermapore plate with a forceps-like pair 

of short processes 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 Rl-Rs bifurcation. Head capsule almost semicircular in 
profile (i. e. H/L<2). Spermapore plate with a short médian appendix . . 
Nephax sofadanus Pearman 

3 Both sexes macropterous (Figs 1; 2a, b), forewing apically rounded or 
slightly acuminate, hindwing well-developed, with complète venation, 
ail 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 Rl-Rs bifurcation (Fig. 2a). T- 
shaped sclerite of female subgenital plate strongly reduced (Figs 2d, 3a) 
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 Rl-Rs bifurcation. T- 
shaped sclerite of female subgenital plate well-developed (Fig. 6d) .... 
Marcenendius fortunatus (Navâs) 

ACKNOWLEDGEMENTS 

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

REFERENCES 

Acon Remâcha, M. 1980. Contribuciôn al conocimiento de los Psocopteros de Espana 

Peninsular. Graellsia (Revista de Entomologos Ihericos) 34: 147-170. 
Badonnel, A. 1943. Psocoptères. Faune de France 42: 1-164. 

Badonnel, A. 1955. Psocoptères de TAngola. Publicacôes culturais da Companhia de Dia - 

mantes de Angola 26: 1-267. 
Baz, A. 1989. Los Psocopteros (Insecta: Psocoptera) del sistema ibérico méridional. 

Universidad de Alcala de Henares, Facultad de Ciencias, Madrid. 230 pp. (Doctoral 

thesis). 



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Baz, A. 2007. Psocoptera (p. 136). In: Domingo, J., Montagud, S. & Sendra, A. (eds). 
Invertebrados endémicos de la Comunitat Valenciana. Conselleria de Territori i 
Habitat ge. General itat Valenciana. 254 pp. 

Baz, A. & Zurita, N. 2001. Orden Psocoptera (pp. 179-180). In: Izquierdo, [., Martin, J. L., 
Zurita, N. & Arechavaleta, M. (eds). Lista de especies silvestres de Canarias (hongos, 
plantas y animales terrestres). Consejeria de Politica Territorial y Medio Ambiente 
Gobierno de Canarias, 437 pp. 

Baz, A. & Zurita, N. 2004. Orden Psocoptera (pp. 188-190). In: Izquierdo, I., Martin, J. L., 
Zurita, N. & Arechavaleta, M. (eds). Lista de especies silvestres de Canarias (hongos, 
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Territorial, Gobierno de Canarias, 499 pp. 

Domingo-Quero, T., Alonso-Zarazaga, M. A., Sânchez-Ruiz, A., Araujo Armero, R., 
Navas Sânchez, A., Sânchez Moreno, S., Garcia Becerra, R., Nebreda, M., 
Sânchez Ruiz, M., Fontal-Cazalla, F. & Nieves- Aldrey, J. L. 2003. Inventariando 
la biodiversidad en el Parque Nacional de La Caldera de Taburiente (La Palma, Islas 
Canarias, Espana): Novedades cientificas. Graellsia 59(2-3): 45-68. (Psoc: p. 62, det. 
A.. Baz). 

KÉLER, S. von 1966. Zur Mechanik der Nahrungsaufnahme bei Corrodentien. Zeitschrift fiir 

Parasitenkunde 27: 64-79. 
Ll Fasheng 1993. Psocoptera from National Chebaling Nature Reserve (Insecta: Psocoptera) 

(pp. 313-430). In: Collected Papers for Investigation in National Chebaling Nature 

Reserve. Science and Technology Publishing House of Guangdong Province. (In 

Chinese, with English summary). 
Lienhard, C. 1988. Contributions to a revision of the western Palaearctic psocids. V. The genus 

Nephax Pearman (Psocoptera: Amphientomidae). Mitteilungen der Schweizerischen 

Entomologischen Gesellschaft 61: 365-372. 
Lienhard, C. 1998. Psocoptères euro-méditerranéens. Faune de France 83: xx+517 pp. 
Lienhard, C. 2009. Order Psocoptera, Part 2 (pp. 49-60). In: Van Harten, A. (éd.). Arthropod 

fauna of the United Arab Emirates, vol. 2. Dar Al Ummah Printing, Abu Dhabi, UAE, 

786 pp. 

Lienhard, C. & Garcia, A. 2005. Psocoptera (p. 70). In: Arechavaleta, M., Zurita, N., Mar- 
rero, M. C. & Martin, J. L. (eds). Lista preliminar de especies silvestres de Cabo 
Verde: hongos, plantas y animales terrestres, 2005. Consejeria de Medio Ambiente y 
Ordenacion Territorial, Gobierno de Canarias, 155 pp. 

Lienhard, C. & Smithers, C. N. 2002. Psocoptera (Insecta): World catalogue and bibliography. 
Instrumenta Biodiversitatis 5: xli+745 pp. Muséum d'histoire naturelle, Genève. 

Meinander, M. 1973. The Psocoptera of the Canary Islands. Notulae Entomologicae 53: 
141-158. 

Meinander, M. 1979. Re vision of the European Psocoptera recorded by Navâs. Miscelanea 
Zoologica 5: 59-67 '. 

Navâs, L. 1913. Sôcidos (Ins. Neur.) de Espana, nuevos. Revista de la Real Academia de 

Ciencias exactas , fisicas y naturales de Madrid 12: 329-335. 
Navâs, L. 1917. Insecta nova. II Séries. Memorie délia Pontifica Accademia Romana dei Nuovi 

Lincei, ser. 2,3: 13-22. 

Navâs, L. 1923. Excursions entomologiques de l'Istiu de 1922. Arxius de l'Institut de Ciencias, 
Barcelona 8: 1-34. 

New, T. R. 1973. Some Psocoptera from Népal. Oriental Insects 7(1): 1-10. 
Pearman, J. V. 1935. Two remarkable Amphientomids (Psocoptera). Stylops 4(6): 134-137. 
Roesler, R. 1944. Die Gattungen der Copeognathen. Stettiner Entomologische Zeitung 105: 
117-166. 

Smithers, C. N. 1972. The classification and phylogeny of the Psocoptera. Australian Muséum 
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Smithers, C. N. 1990. Keys to the families and gênera of Psocoptera (Arthropoda: Insecta). 

Technical Reports of the Australian Muséum 2: 1-82. 
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Matsumurana, New Séries 62: 1-44. 



Revue suisse de Zoologie 118 (3): 467-484; septembre 201 1 



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

David GONZÂLEZ-SOLIS 1 & Jean MARI AUX 2 

1 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 
fîshes from Paraguay deposited in the Muséum of Natural History of 
Geneva. - The list comprises the évaluation of the material deposited in the 
Muséum d'Histoire naturelle, Geneva, whose taxonomic examination 
revealed the présence of 78 taxa of parasitic nematodes of freshwater fishes 
from Paraguay. Out of thèse, 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 
fréquent 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 région 
is poorly known and there still exist vast territories where parasitological surveys are 
lacking (Moravec, 1998). Paraguay represents one of thèse scarcely studied areas, 
since only few reports on fish parasites, based on a limited number of hosts and from 
very spécifie geographical areas, have been published so far (Masi Pallarés et al., 1973; 
Petter 1984, 1989, 1990, 1994, 1995a,b; Petter and Cassone, 1984; Petter and Dlouhy, 
1985; Petter and Morand, 1988). From thèse works, to date 28 nematodes have been 
reported from Paraguay, particularly from localities of the Paranâ and Paraguay Rivers 
and provinces bordering or close to Brazil (Concepcion, Alto Paranâ 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 .20 1 1 



468 



D. GONZÂLES-SOLI'S & J. MARIAUX 



MATERIALS AND METHODS 

The nematode spécimens used for this publication were recovered during the 
various expéditions of the MHNG in Paraguay (EMGP) between 1987 and 1996. 
Fishes belonging to 63 gênera 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 région 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 Paraguay an Alto 
Paranâ, which represents the most nematologically studied région 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 spécimens belonging to 92 species. Taxonomic 
références for hosts and parasites are Froese and Pauly (2011) and Moravec (1998) 
respectively. Each identified taxon is listed together with its host names, abbreviated 
localities and sampling date. Host species with an asterisk (*) represent new records. 
A list of localities is provided in the appendix. 

RESULTS 
Order Enoplida 

Family Capillariidae Railliet, 1915 

Capillariidae gen. sp. 

Spécimens: 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 naîtereri Steindachner, 1908 (= Serrasalmus 
natterert)* (VI 13/11/87, PC 05/01/96). 

REMARKS: Thèse spécimens were young fe maies or maies whose spécifie or 
generic identication was impossible. Capillariids from A. 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. naîtereri. 

Capillostrongyloides sentinosa (Travassos, 1927) 
Spécimens: 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. 

Spécimens: MHNG INVE 73325, 73345. 

HOST: Trachely opter us 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. 

Spécimen: 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 

Spécimens: MHNG INVE 73513. 
HOST: Pimelodella sp. (VI 13/1 1/87). 

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

Cosrnoxynema sp. 

Spécimens: MHNG INVE 73511. 7362 1 . 

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

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

Cosrnoxynema vianai Travassos, 1949 

Spécimens: MHNG INVE 73664, 73675. 73678, 73682, 73690, 74215. 

HOST: Cyphocharax modestus (Fernândez-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 

Spécimens: 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. 

Spécimen: MHNG INVE 73691 . 

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

Remarks: Only one maie available that could be a new species since the genus 
has ne ver been found in this fish family (Curimatidae). 

Ichthyouris brasiliensis Moravec, Kohn et Fernandes, 1992 

Spécimens: MHNG INVE 73402. 73415. 73426. 

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



470 



D. GONZÂLES-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 

Spécimens: MHNG INVE 73348, 73382, 73385, 73387, 73388. 

HOST: Rhinelepis aspera Spix & Agassiz, 1829 (CA 02/06/87, VI 14/1 1/87). 
Remarks: New locality record. 

Spinoxyuris sp. 

Spécimen: MHNG INVE 73685. 

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

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

Spinoxyuris oxy doras Petter, 1994 

Spécimens: MHNG 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 

Spécimens: 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 iheringi (Travassos, Artigas et Pereira, 1928) 

Spécimens: MHNG 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. 

Spécimens: MHNG INVE 73354, 73633. 

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

Remarks: Thèse larval nematodes probably belong to the species R. rondoni 
which parasitizes congeneric fishes. Unfortunately, their poor development prevents a 
spécifie identification. 

Rondonia rondoni Travassos, 1920 

Spécimens: 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, 7335 1 , 73590, 73653, 73654, 
74205, 74335, 74338, 74403, 74221 . 



NEMATODES FROM FRESHWATER FISHFS FROM PARAGUAY 



471 



HOSTS: Ageneiosus inermis Valenciennes, 1840 (= A. brevifilis)* (SA 06/1 1/95). 
Doradidae gen. sp. (ED 26/06/96, 17/10/96). Piaracîus 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). Pterodoras 
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 03/1 1/95). 
Serrasalmus nattereri* (ED 02/10/96). Sorubim lima (Bloch & Schneider, 1801)* (SA 
27/11/93). 

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

Family Cosmocercidae Railliet, 1916 

Raillietnema sp. 

Spécimen: 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 Di'az-Hungria, 1968 

Spécimen: MHNG INVE 73855. 

HOST: Piaracîus mesopotamicus* (SA 14/10/89). 

Remarks: First record in this fish species and geographical area. 

Spectatus spectatus Travassos, 1923 

Spécimen: MHNG INVE 73372. 

HOST: Piaracîus mesopotamicus* (VI 14/11/87). 

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

Family Quimperiidae Gendre, 1928 

Quimperiidae gen. sp. 

Spécimen: MHNG INVE 73553. 

HOST: Brycon orbignyanus (Valenciennes, 1850) (AA 07/10/89). 
Remarks: Poorly developed larva. 

Paraseuratum sp. 

Spécimens: 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 

Spécimens: MHNG INVE 73708, 73731, 74638. 



472 



D. GONZÂLES-SOLI'S & J. MARIAI X 



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

Touzeta ecuadoris Petter, 1987 

Spécimen: MHNG INVE 73710. 

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

Family Cucullanidae Muller. 1777 

Cuculianidae gen. sp. 

Spécimens: MHNG INVE 73365. 73484. 73637. 73658. 

HOSTS: Bryccm 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: Thèse specimenes were small and poorly developed larvae that 
could belong to the genus Cucullanus. 

CucuUanus sp. 

Spécimens: 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 pari (Valenciennes. 1835) (AT 28/10/89). Megalonema platanum 
(Giinther. 1880) (CA 29/05/87). Pimelodus sp. (R6 09/10/88). Pimelodus cf. albicans 
(ED 03/08/94). Pimelodus maculants (CA 02/06/87). Pimelodus ornants 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 z.ungaro (Steindachner. 1877) (= Paulicea luetkeni) (SA 27/11/93). 

Remarks: Larvae and young females that could be of the species CucuUanus 
pinnai pinnai or C. p. pterodorasi . 

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

Spécimens: 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 Paranâ River basin. 

Cucullanus (Cucullanus) pinnai pterodorasi Moravec, Kohn et Fernandes, 1997 

Spécimens: MHNG INVE 73319. 73583. 

HOST: Plagioscion ternetzi* (PI 08/10/91). Pterodoras 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 

Spécimens: 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. 

Spécimens: MHNG INVE 73640, 73642. 

HOST: Pimelodus cf. albicans (ED 03/08/94). 

Remarks: Thèse two young females might belong to D. pimelodi. 

Dichelyne pimelodi Moravec. Kohn et Fernandes, 1997 

Spécimens: 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) 

Spécimens: 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 (Paranâ River) but from différent province (Itaupu). 

Family Acanthocheilidae Wûlker, 1929 

Brevimulticaecum sp. 

Spécimen: 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. 

Spécimen: MHNG 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 is the first ani- 
sakid form occurring in thèse fish species. 

Contracaecum sp. type 1 

Spécimens: 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. GONZÂLES-SOLIS & J. MARIAUX 



73763, 73773, 73782, 73786, 73840, 73842, 73848, 73850, 73852, 73863, 73866, 73875, 73880, 
73885, 73890, 74217, 74218, 74222, 74295, 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 (Fernândez-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 (Gunther, 1874)* (VI 13/11/87). 

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

. Contracaecum sp. type 2 

Spécimens: MHNG INVE 73871, 73874,73450, 73414, 73441, 74657, 73719, 73397, 
73416, 73733, 73759, 73775, 73777, 73389, 73392, 73394, 73408, 73473, 73623, 73771 , 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 18/11787, 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. Thèse 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. 

Spécimens: MHNG INVE 73333, 73349, 73355, 73358, 73359, 73366, 73368, 73370, 
73391 , 73393, 73395, 73399, 73400, 7341 1 , 73417, 73425, 73442, 73443, 73444, 73449, 73453, 
73456, 73467, 73470, 73474, 73477, 73489, 73490, 73493, 73499, 73502, 73503, 73508, 73517, 
73521 , 73522, 73527, 73535, 73572, 73587, 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, 20/1 1/87). 
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 Isbrùcker & Nijssen, 1979* (EG 20/1 1/87). Lycengraulis 
grossidens (Eigenmann, 1907)* (PE 13/08/87). Markiana nigripinnis (Perugia, 1891)* 
(EG 20/11/87). Megalechis thoracata (Valenciennes, 1840) (= Hoplosternum thora - 
cataY (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. Thèse larvae could belong to 
différent 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 

Spécimens: 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. 

Spécimens: MHNG INVE 73374, 73375, 73378, 73379, 73381 , 73494, 74472. 

HOSTS: Doradidae gen. sp. (EF 15/02/96), 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) 

Spécimens: 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. GONZÂLES-SOLIS & J. MARIAUX 



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

Spécimens: MHNG INVE 73353, 73476, 73735, 74206, 74208, 74212, 74463, 74620, 

74623. 

HOSTS: Geophagus sp* (PI 05/01/96). 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. 

Spécimen: MHNG INVE 74181. 

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

Raphidascaroides brasiliensis Moravec et Thatcher, 1997 

Spécimens: 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). 
Platy doras costatus* (GN 05/1 1/91 , 07/1 1/91 , G Y 15/09/94). 

Remarks: New host and geographical records. 

Goezia brasiliensis Moravec, Kohn et Fernandes, 1994 

Spécimen: MHNG INVE 73361. 

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

• Porrocaecum sp. 

Spécimens: 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 thèse fish hosts and in South America. 

Order Spirurida Chitwood, 1933 
Familia Spiruridae 

Spiruridae gen. sp. 

Spécimen: MHNG INVE 73721. 

HOST: Hoplerythrinus unitaeniatus (ES 27/10/87). 

REMARKS: Very small and poorly developed larvae. 

Family Camallanidae Raillietet Henry, 1915 

Camallanidae gen. sp. 

Spécimen: 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. 

Spécimens: MHNG INVE 73627, 73872. 

HOSTS: Acestrorhynchus altus (SN 17/10/89). Pimelodus sp. (GN 07/1 1/91). 
Remarks: Small larvae without striations on the buccal capsule. 

Procamallanus {Procamallanus) sp. 

Spécimen: MHNG INVE 74269. 

HOSTS: Ageneiosus inermis* (SA 29/1 1/93). 

Remarks: New host record. 

Procamallanus (Procamallanus) annipetterae Kohn et Fernandes, 1988 

Spécimens: 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 

Spécimens: 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. 

Spécimens: 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 loricariids. 

Procamallanus (Spirocamallanus) cervicalatus (Petter, 1990) 

Spécimens: 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 

Spécimens: 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. GONZÂLES-SOLIS & J. MARIAUX 



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

Spécimens: MHNG INVE 73451, 73455, 73462, 73466, 73469, 73472, 73475, 73478, 
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 lacusîris (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 15/02/96). 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) 

Spécimen: MHNG INVE 73760. 

HOST: Hoplerythrinus unitaeniatus (AT 25/10/89). 

Remarks: New geographical record. 

Procamallanus (Spirocamallanus) paraensis Pinto et Noronha, 1976 

Spécimen: MHNG INVE 73604. 

HOST: Rhaphiodon vulpinus* (RS 09/10/91). 

Remarks: New host and geographical records. 

Procamallanus (Spirocamallanus) paraguayensis (Petter, 1990) 

Spécimens: MHNG INVE 73888, 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, Fâbio, Noronha et Rolas, 1974 

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

HOSTS: Pimelodella sp. (VI 13/11/87). Pimelodella grijfini 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 

Spécimens: MHNG INVE 73306, 73616. 

HOSTS: Iheringichthys 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. 

Spécimens: MHNG INVE 73861,73876,73882,73884,73887. 
HOSTS: Acestrorhynchus altus* (SN 17/10/89). Hoplosternum littorale* (SN 
\l/\0/S9).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 

Spécimens: 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 

Spécimens: MHNG INVE 73758, 73772, 73776, 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 

Spécimen: MHNG INVE 73720. 

HOST: Hoplerythrinus unitaeniatus (ES 27/10/87). 

Remarks : New geographical record. 

Travassosnema sp. 

Spécimen: MHNG INVE 73571 . 

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

REMARKS: Young female that probably belongs to the species Travassosnema 
travassosi paranaensis. 

Travassosnema travassosi paranaensis Moravec, Kohn et Fernandes, 1993 

Spécimens: 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. 

Spécimens: MHNG INVE 73566, 73580, 73581 . 
HOST: Crenicichla lepidota (AT 23-24/10/89). 

REMARKS: Maies in poor condition, impossible to détermine to the generic 

level. 



480 



D. GONZÂLES-SOLI'S & J. MARI AUX 



Family Gnathostomatidae Railliet, 1895 

Gnathostoma sp. 

Spécimen: 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. 

Spécimens: 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 imitaeniatus* (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). Trachely opter us galeatus* (GN 
05/11/91). Thporîheus paranensis* (VI 13/11/87). 

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

Rhabdochona acuminata (Molin, 1860) 

Spécimens: 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-Hungrfa, 1968 

Spécimen: MHNG INVE 73745. 

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

Family Cystidicolidae Skrjabin, 1946 

Cystidicoloides sp. 

Spécimens: 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. 



MAI MODES I ROM I RI.SHW \l I.R I ISHI S I ROM l'\R \ï 



481 



Spinitectus sp. 

Spécimen: MHNG INVE 73613. 

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

REMARKS: Females that could belong to S. multipapillaïus . 

Spinitectus asperus Travassos, Artigas et Pereira, 1928 

Spécimen: 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 

Spécimens: 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 

Spécimens: 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: Ail are new host records. 

Family Physalopteridae Railliet, 1893 

Heliconema sp. 

Spécimen: MHNG INVE 73652. 

HOST: Oxyrhopus melanogenis?* (SL 30/07/88). 
REMARKS: New host and geographical record. 

Pseudoproleptus sp. 

Spécimens: 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 is the first finding of thèse nematodes in Paraguay, therefore ail 
represent new host and geograpical records. 

Family Acuariidae Railliet, Henry et Sisoff, 1912 

Acuariidae gen. sp. 

Spécimens: 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. GONZÂLES-SOLIS & J. MARI AUX 



CONCLUSIONS 

The nematodes reported herein showed différent distribution range and hosts 
diversity. The species whose adult presented the broadest geographical and host ranges 
were R. rondoni (14 localities and 10 hosts) and Procamallanus {Spirocamallamis) 
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 juvénile 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 Paranâ 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 et 
al. 2009). Other factors contributing to this relatively poor parasite diversity are the 
low degree of host specificity of thèse 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 522). 

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 parâsitos del 

Nuevo Mundo. ECFACIM, Asunciôn, 689 pp. 
Masi-Pallarés, R., Benltez-Usher, C. A. & Vergara, G. 1973. Helminthes en peces y repti- 
les del Paraguay (lera parte). Revista Paraguaya de Microbiologia 8: 67-96. 
Moravec, F. 1998. Nematodes of freshwater fishes of the Neotropical région. Academia, Praha, 

Czech Republic, 464 pp. 
Moravec, F, Kohn, A. & Fernandes, B. M. M. 1992. Nematode parasites of fishes of the 

Paranâ River, Brazil. Part 1. Trichuroidea, Oxyuroidea and Cosmocercoidea. Folia 

Parasitologica 39: 327-353. 
Petter, A. 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, A. J. 1989. Nématodes de poissons du Paraguay. V. Cucullanidae. Description de deux 
espèces nouvelles et redéfinition du genre Neocucullanus Travassos et al. Revue suisse 
de Zoologie 96: 591-603. 



NEMATODES FROM FRESHWATER FISHES FROM PARAGUAY 



483 



Petter. A. J. 1990. Nématodes de poissons du Paraguay. VI. Description de deux nouvelles 

espèces du genre Spirocamalkmus et compléments à la description de Procamallanus 

annipetîerae Kohn & Fernandes. 1988. Revue suisse de Zoologie 97: 327-338. 
Petter. A. J. 1994. Nématodes de poissons du Paraguay. VII. Oxyuroidea: Spinoxyuris oxydants 

n.g.. n. sp. Revue suisse de Zoologie 101: 761-769. 
Petter, A. J. 1995a. Nématodes de poissons du Paraguay. VIII. Habronematoidea. Dracun- 

culoidea et Ascaridoidea. Revue suisse de Zoologie 102: 89-102. 
Petter. A. J. 1995b. Dichefyne moraveci n. sp.. parasite de Pseudoplatystoma fasciatum et notes 

sur les Cucullanidae du Paraguay. Revue suisse de Zoologie 102: 769-778. 
Petter. A. J. & Cassone. J. 1984. Nématodes de poissons du Paraguay. I. Ascaridoidea: 

Sprentascaris n. gen. Revue suisse de Zoologie 91: 617-634. 
Petter. A. 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. A. 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. A., da Silva. F. A. & Silya-Solza. 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. A. P.. Lacerda. A. C. F.. Yamada, F. H.. 
Moreira. L. H. A.. Ceschim. T. L. & Bellay. S. 2009. Diversity of parasites of fish 
from the Upper Paranâ River floodplain. Brazil. Brazilian Journal ofBiologx 69(suppl. 
2): 691-705. 

Travassos. L. 1927. Uma nova Capillaria parasita de peixes de âgua doce: Capillaha sentinosa 

n. sp. Boletin de Biologia 10: 215-217. 
Vicente, J. J. & Pinto, R. M. 1999. Nematôides do Brasil. Nematôides de peixes. Atualizacao: 

1985-1999. Revista Brasileira de Zoologia 16(3): 561-610. 



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





Locality 


Province 


Latitude 


Longitude 


AA 


Arroyo Allègre 


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 


Présidente Hayes 


-23.36 


-59.07 




(Trans Chaco Km 303) 








AP 


Arroy o Piraty i 


Canindeyu 


-24.07 


-54.30 


AR 


Arroyo Trementina 


Concepcion 


-22.82 


-56.70 


AT 


Arroyo Tapie uarai 


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) 


Présidente Hayes 


-25.07 


-57.95 


EG 


Estancia (Laguna) General Diaz 


Alto Paraguay 


-21.13 


-58.50 


EM 


Estancia Las Margaritas (Pilcomayo) 


Présidente 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 Paranâ 


-25.42 


-55.38 



484 



D. GONZÂLES-SOLIS & J. MARI AUX 



I V 


rdsU ^UIICcl 


V^CIllI al 


9S 1 9 

-ZJ . 1 Z 


S7 AS 


PF 


Piif^rtn Prif^lirQ 

r UCl LU HJCIlltl 


liapila 


-96 QO 


S S 1 7 

- j j . i / 


PT 
il 


Piquete-cué 


Central 


?S 1 9 

-ZZ) . 1 z 


- j / .DU 


PT 


r diicnuo Lopcz 


iviisiones 


-Z / .H-U 


S7 97 

- J / .Z / 


PO 


Pozo Arias 


Présidente Hayes 


9^ 


-OU.J)Z 


PV 


rllcIlO LyiU VcIUc 


ït"QT"M 1 Q 
lldpiid 


96 87 
-zo .o / 


-JJ . 1 J 


P7 


I UIUO Z^lIlIlO 


^onccpcion 


99 ^8 
-ZZ.jo 


S6 en 

-J0.7J 


IV II 


R-Par 638 


v^-uiiccpciun 


-93 83 

Z. J .OJ 


-S7 97 


IV 3 


lYlO OdldUO 


^.Clllldl 


9S 1 9 
-ZJ . 1 z 


S7 4 S 


SA 


San Antonio 


Central 


-25.43 


-57.55 


SL 


San Lorenzo 


Central 


-25.30 


-57.50 


SN 


San Lorenzo (10 Km SE) 


Neembucu 


-26.78 


-57.67 


SP 


Salto Pirareta 


Cordillera 


-25.55 


-56.90 


SU 


San Luis (1 lKm E) 


Concepcion 


-22.43 


-57.34 


VI 


Villeta 


Central 


-25.52 


-57.50 


YA 


Yabebyry 


Misiones 


-27.40 


-57.13 



Revue suisse de Zoologie 118 (3): 485-489; septembre 201 1 



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

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

Food habits of escaped Eurasian otters {Lutra lutra) in a suburban 
environment in Switzerland. - In 2005, a maie and a female otters escaped 
from the zoo of Bern, and settled in the nearby River Aar. The number of 
otters présent in the area increased to 5 individuals after the adult pair 
reproduced. A monitoring was launched in 2007 in order to examine how 
thèse 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 
maie 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 
thèse individuals was decided by the fédéral and cantonal authorities. In 2007, the maie 
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 maie, was also captured 
in 2007, leaving a young maie and a female in the wild. 

An extensive monitoring was launched to examine how thèse 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, évidences such as a generalized and drastic décline of 



Manuscript accepted 10.06.201 1 
* présent address: SFFN, Section Faune, Rue du ler-Mars 11, 2108 Couvet, Switzerland. 



486 



J.-M. WEBER 



fish catch in Svviss rivers, including the River Aar, since the 1990s suggest a possible 
détérioration of the potential trophic conditions for otters (Burkhardt-Holm et al., 
2002), but the exact conséquences 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 accidentai 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 fevv ponds adjoin the main river, and an undisrupted riparian forest is 
présent on both sides of it. More than 40 fish species occur in the région. In the absence 
of récent fish biomass counts (i.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 
fario), grayling {Thymallus thymallus), perch (Perça fluviatilis) and pike (Esox lucius), 
respectively, are the most frequently angled fish in the area (Fish Management Service 
of the Canton Bern, pers. corn.). Two species of crayfish, a potential alternative prey to 
otters, live also in the area (Stucky & Zaugg, 2005). The river flows in a dépression 
surrounded by the suburbs of Bern. Therefore, human (e.g. walkers, joggers, swimmers 
and bikers) présence is important year-round along the stream. A 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 ail otter faeces were collected. Faeces were 
dissected and analysed as described in Jenkins et al. (1979). Fish and amphibian 
remains, mostly scales and bones, were identified according to Conroy et al. (1993). 
The results of the analyses were expressed in relative frequency of occurrence (i.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 catégories 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 2 test = 5.1394, p > 0.05). 
Cyprinid exploitation tended to decrease in winter, although the trend was also not 
significant (Chi 2 test = 2.1997, p > 0.05). In contrast, pike occurred more frequently in 
the diet during winter than in other seasons (Chi 2 test = 12.9634. p < 0.01). Prédation 
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 




unueierrnincu ^ypnniuae 


Al 


1 7 1 
1 / . 1 




Minnow (Phoxinus phoxinus) 


LU 


A 1 
4. 1 




Bullhead (Cottus gobio) 




1 j.U 




Pike (Esox lucius) 


27 


I 1 A 

1 1 .0 




Perch (Perça fluviatilis) 


A 

4 


1 £. 

1 .6 




Undetermined Fish 


4 


1.6 




Total Fish 


225 


91.5 




Arthropods 


2 


0.8 




Amphibians 


11 


4.5 




Reptiles 


4 


1.6 




Birds 


2 


0.8 




Mammals 


1 


0.4 




Undetermined 


1 


0.4 




Total 


246 


100 




Table 2. Seasonal variation in the différent prey catégories 


(%) found in otters'spraints from the 


River Aar. Number of items counted in spraints are in parenthèses. 




Spring(105) Summer(18) 


Autumn (90) 


Winter (33) 


Salmonidae 38.1 44.4 




53.3 


30.3 


Cyprinidae 19.1 33.2 




24.4 


12.1 


Cottidae 13.3 11.1 




13.3 


12.1 


Esocidae 15.2 




3.4 


24.3 


Amphibia 6.7 




2.2 


6.0 


Otherprey 7.6 11.3 




3.4 


15.2 



DISCUSSION 

Both habitat type and latitude play a rôle in the composition of otter's diet in 
Europe (Jedrzejewska et al., 2001 , Clavero et al., 2003). According to thèse 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, i.e. according to prey availability and abundance (Kruuk, 2006). Food 
composition should, therefore, reflect vvhich 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 différences vvith our results 
vvere a much higher présence 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 et al., 2001, Brzezinski et al., 2006). 

Would the River Aar offer a suitable habitat for otters as suggested by Weber 
(1997)? A continued 5-yr présence 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 région. However, repro- 
duction may not be the sole criterion to ensure a long-term survival, as experienced 
vvith the reintroduced otters of the Schwarzwasser. Indeed, the introduced animais 
eventually disappeared from the area approximately 10 years after their release, despite 
successful reproduction (Weber et 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 décline of fish 
catch observed in the Swiss waters (Burkhardt-Holm et 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. corn.). Under 
thèse circumstances, accidentally released otters are possibly confronted to suboptimal 
trophic conditions, as far as their main prey, i.e. salmonids, are concerned. However, 
the fish farm présent 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 rôle as food supplier 
would have remained very local and time-limited, particularly if 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 et 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 thèse opportunistic observations gathered following the accidentai 
release of two otters, a perennial présence 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 Fédéral 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 référées for useful 
comments on the manuscript. 



DIET OF ESCAPED EURASIAN OTTERS 



489 



REFERENCES 

Clavero, M., Prenda, J. & Delibes, M. 2003. Throphic diversity of the otter (Lutra lutra L.) 
in temperate and Mediterranean freshwater habitats. Journal of Biogeography 28: 76 1 - 
769. 

Clavero, M., Hermoso, V., Brotons, L. & Delibes, M. 2011. Natural, human and spatial 
constraints to expanding populations of otters in the Iberian Peninsula. Journal of 
Biogeography. 

Brzezinski, M., Romanovski, J., Kopczynski, L. & Kurowicka, E. 2006. Habitat and 
seasonal variations in diet of otters, Lutra lutra in eastern Poland. Folia Zoologica 55: 
337- 348. 

Burkhardt-holm, P., Peter, A. & Segner, H. 2002. Décline of fish catch in Switzerland. 

Project Fishnet: a balance betvveen analysis and synthesis. Aquatic Sciences 64: 36-54. 
Conroy, J. W. H., Watt, J., Webb, J. B. & Jones, A. 1993. A guide to the identification of prey 

remains in otter spraint. An Occasional Publication of the Mammal Society, No 16. 
Jenkins, D., Walker, J. G. K. & McCowan, D. 1979. Analyses of otter (Lutra lutra) faeces 

from Deeside, N.E. Scotland. Journal ofZoology 187: 235-244. 
Jedrzejewska, B., Sidorovich, N. V., Pikulik, M. M. & Jedrzejewski, W. 2001. Feeding 

habits of the otter and the American mink in Bialowieza Primeval Forest (Poland) 

compared to other Eurasian populations. Ecography 24: 165-180. 
Kruuk, H. 2006. Otters: ecology, behaviour, and conservation. Oxford University Press, Oxford 

and New York, 265 pp. 

Lanzski, J., Kôrmendi, S., Hancz, C. & Martin,T.G. 2001. Examination of some factors 

affecting sélection of fish prey by otters (Lutra lutra) living by eutrophic fish ponds. 

Journal ofZoology 255: 97-103. 
Stucky, P. & Zaugg, B. 2010. Decapoda. Fauna Helvetica 15. CSCF, Neuchâtel, 56 pp. 
Weber, D. 1990a. Das Ende des Fischotters in der Schweiz. Schriftenreihe Umwelt Nr 128, 

Dokumentationsdienst Bundesamt fïir Umwelt, Bern, 101 pp. 
Weber, J.-M. 1990b. Seasonal exploitation of amphibians by otters (Lutra lutra) in north-east 

Scotland. Journal ofZoology 220: 641-651. 
Weber, D. 1992. Warum wurde der Versuch gestoppt, Otter in der Schweiz auszusetzen? In: 

Reuther, C. (éd.), Otterschutz in Deutschland. Habitat 7, Hankensbiittel, 176 pp. 
Weber, J.-M. 1997. Habitats potentiels de la loutre en Suisse: nouvelle évaluation. Rapport 

WWF-Suisse, Zurich, 10 pp. 
Weber, J.-M. 2008. Suivi des loutres de l'Aar. KORA Bericht 44, Mûri, 27 pp. 
Weber, J.-M. & Weber, D. 1991. The otter (Lutra lutra L.) in western Switzerland, 1989 

(pp. 53-55). In: Reuther, C. & Rôchert, R. (eds), Proceedings of the V. International 

Otter Colloquium. Habitat 6, Hankensbiittel, 344 pp. 
Weber, D., Weber, J.-M. & Mûller, H.-U. 1991. Fischotter (Lutra lutra L.) im Schwarz - 

wasser-Sense-Gebiet: Dokumentation eines gescheiterten Wiedereinburgerungsver- 

suchs. Mitteilungen der Naturforschenden Gesellschaft in Bern Al: 141-152. 



Revue suisse de Zoologie 118 (3): 491-512; septembre 201 1 



Claude Besuchet, an eminent swiss coleopterists, 80 years old 

Peter HLAVÂC 

Na doline 14, SK-040 14 Kosice, 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, I visited the Muséum 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 sphère 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 I made contact with ail actively work- 
ing Pselaphidologists and started to search for reprints of their papers. Thus in 1994 I 
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 ail 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") thèse 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 ail entomologists a little about your 80 years of 
life of which 63 were very much devoted mainly to your studies of Pselaphines. 

BlOGRAPHY 

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, Trimium 
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 thèse small, 
wonderfull créatures. 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 25.02.201 1 



492 



P. HLAVÂC 



In 1954 he graduated with the certificate for Physiology, Bacteriology, Parasitology, 
Hygiène 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 vvhere he had been vvorking for short time till 1956. But the 
name of Claude Besuchet vvill be forever connected to the Muséum d'Histoire naturelle 
de Genève, where he has been working at the department of entomology since l st 
January 1958 until his retirement at 31 st 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 conséquence 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 ail in ail 155 papers, from this 126 purely scientific taxonomic 
papers devoted to Staphylinidae (Pselaphinae), Scydmaenidae, Ptiliidae, 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 gênerai 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 thèse taxa are from the Palaearctic région (350), 
followed by the Oriental région (53), Afrotropical région (17), Neotropical région 
(10), Australian région (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 


332 


1 


1 


9 


14 


2 


1 360 


Scydmaenidae 


10 


16 








3 


1 


30 


Ptiliidae 


6 












6 


Leptinidae 


2 












2 


Cerylonidae 










35 




35 


Rhipiporidae 








1 






1 


Dryopidae 










1 




1 


TOTAL 


350 


17 


1 


10 


53 


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 Slipinski, 
one of most prominent specialists on Cucujoidea, visited Geneva muséum a few times. 
The resuit of this fruitfull coopération was two important revisions, Glyptolopus 
Erichson (1987b) and Axiocerylon Grouvelle (1988b). 



CLAUDE BESUCHET'S WORK 



493 




FlG. 1 

Claude Besuchet (left) and Ivan Lôbl (right) at the Natural History Muséum of Geneva. 



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

Rhipiporidae: Besuchet'paper (1956d) 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 Lepîinus (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 présence 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 Ptinella Motschulsky). Another two species were 
described in 1980e and last paper on this family of minute beetles is a list of Ptiliidae 
from Mongolia collected by Zoltân Kaszab (1969h). 

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



494 



P. HLAVÂC 



in ail in tvelve papers (Besuchet, 1958e, 1959b, 1961h, 1962c, 1971a,b, 1980d, 1981c, 
2004d, 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 gênera (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 région. Much less attention 
he paid to the tropical fauna, only 27 species were named by him from tropical régions, 
single species from the Nearctic région and 332 species from the Palearctic région. 

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 Gênera: 

1. Antrobythus Besuchet, 1985a: 511 (Staphylinidae: Pselaphinae) 

2. 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. Couloniella Besuchet, 1983a: 509 (Staphylinidae: Pselaphinae) 

7. Geoparnus Besuchet, 1978: 705 (Dryopidae) 

8. Nonveillera Pavicevic & Besuchet, 2003a: 279 (Staphylinidae: Pselaphinae) 

9. Pachacuti Besuchet, 1987a: 231 (Staphylinidae: Pselaphinae) 

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

11. Plitium Besuchet, 1971c: 329 (= Ptinella Motschulsky) (Ptiliidae) 

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) 

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

7. Acetalius pilosus Besuchet, 1985b: 763 

2. Afropselaphus breiti Besuchet, 1961a: 34 {Pselaphogenius) 

3. Afropselaphus canariensis Besuchet, 1968: 291 



CLAUDE BESUCHET'S WORK 



495 



4. Afropselaphus circassicus Besuchet, 1961a: 37 (Pselaphogenius) 

5. Afropselaphus fernandezi Besuchet, 1968: 290 

6. Afropselaphus g iianche Besuchet, 1970a: 123 

7. Afropselaphus maroccanus Besuchet, 1963b: 222 {Pselaphogenius) 

8. Afropselaphus spinipalpis Besuchet, 1968: 289 

9. Afropselaphus zacynthius Besuchet, 1961a: 35 {Pselaphogenius) 
10. Amauronyx auberti Besuchet, 1962b: 339 

77. Amauronyx caecus Besuchet, 1962b: 340 

72. Amauronyx caudatus Besuchet, 1999b: 798 

13. Amauronyx cobosi Besuchet, 1959a: 26 

14. Amauronyx franzi Besuchet, 1958e: 908 
75. Amauronyx mussardi Besuchet, 1963c: 229 

16. Amauronyx myops Besuchet, 1962b: 337 

17. Amaurops aubei binaghii Besuchet, 1980b: 615 

18. Antrobythus leclerci Besuchet, 1985a: 512 

19. Antrobythus perplexus Besuchet, 1993b: 223 

20. Bathybythus bleyi Besuchet, 1974b: 43 

27. Batrisodes bifossulatus Besuchet, 1988a: 436 

22. Batrisodes clypeatus Besuchet, 1981a: 290 

23. Batrisodes mitovi Besuchet & Bekchiev, 2007b: 75 

24. Batrisodes rousi Besuchet, 1981a: 292 

25. Batrisodes sulcaticeps Besuchet, 1981a: 289 

26. Batrisodes unisexualis Besuchet, 1988a: 433 

27. Batrisus taurus Besuchet, 2004e: 28 new name for tauricus Besuchet, 1979: 279 

28. Bibloplectus (s. str.) atomus Besuchet, 1958e: 906 

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

30. Bibloplectus (s.str.) beaumonti Besuchet, 1955a: 200 

31. Bibloplectus (s.str.) boveyi Besuchet, 1975a: 32 

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

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

34. Bibloplectus elongatus Besuchet, 1953: 231 (= B. strouhali Beier) 

35. Bibloplectus (s.str.) franzi Besuchet, 1964a: 411 

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

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

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

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

40. Bibloplectus linderi Besuchet, 1953: 228 (= tenebrosus Reitter) 

41. Bibloplectus (s.str.) machulkai Besuchet, 1955a: 187 

42. Bibloplectus minutus Besuchet, 1953: 229 ( = B. obtusus Guillebeau) 

43. Bibloplectus (s.str.) normandi Besuchet, 1955a: 196 

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

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

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

47. Bibloplectus pseudambiguus Besuchet, 1953: 226 (= B. spinosus Raffray) 

48. Bibloplectus (s.str.) subtilis Besuchet, 1975a: 35 



496 p HLAVÂC 

49. Bibloplectus (s.str.) iantulus Besuchet, 1975a: 35 

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

51 . Bibloplectus therondi Besuchet. 1953: 230 (= B. pusillus Denny) 

52. Bibloporus bicolor franzi Besuchet. 1958e: 903 

53. Bibloporus myops Besuchet, 1970b: 313 

54. Brachygluta abrupta septemtrionalis Besuchet. 1963a: 34 
(= B. abrupta Dodero) 

55. Brachygluta alpina Besuchet, 2004a: 109 

56. Brachygluta atlantica Besuchet, 2004a: 154 

57. Brachygluta exigua Besuchet. 1963a: 41 

58. Brachygluta exsculpta Besuchet. 1969e: 405 

59. Brachygluta franciscae Besuchet. 1963a: 35 

60. Brachygluta gnosiaca Besuchet. 2004a: 58 

61 . Brachygluta hanseni Besuchet. 1954c: 43 (= B. paludosa Peyron) 

62. Brachygluta hispana Besuchet. 1963a: 40 

63. Brachygluta jordanica Besuchet. 2004a: 78 

64. Brachygluta kurdica Besuchet. 2004a: 144 

65. Brachygluta lefebvrei meridionalis Besuchet. 1962b: 356 
( = B. lefebvrei lederi Saulcy) 

66. Brachygluta occidentalis Besuchet. 1963a: 35 

67. Brachygluta perissinottoi Besuchet. 1969e: 402 

68. Brachygluta pusilla Besuchet. 1958d: 335 

69. Brachygluta richteri Besuchet. 1961e: 1 

70. Brachygluta sengleti Besuchet. 1969e: 404 
77. Brachygluta tumidipes Besuchet. 1981b: 243 

72. Brachygluta ultima Besuchet, 2004a: 80 

73. Brachygluta vicaria Besuchet. 1963a: 43 

74. Bryaxis abkhasicus Besuchet & Kurbatov. 2007: 202 

75. Bryaxis adjaricus Besuchet & Kurbatov. 2007: 202 

76. Bryaxis adumbratus Besuchet & Kurbatov.2007: 172 

77. Bryaxis altivagus Besuchet. 1962b: 354 

78. Bryaxis arnoldii Besuchet. 1961g: 1830 

79. Bryaxis artvinensis Besuchet & Kurbatov. 2007: 179 

80. Bryaxis assingi Besuchet & Kurbatov. 2007: 178 

81 . Bryaxis atlanticus Besuchet. 1962b: 353 

82. Bryaxis badins Besuchet. 1961g: 1827 

83. Bryaxis balabanus Besuchet & Kurbatov. 2007: 188 

84. Bryaxis balneator Besuchet & Kurbatov. 2007: 193 

85. Bryaxis bergamascus breiti Besuchet. 1980d: 625 
(= B. bergamascus sorinensis Stolz) 

86. Bi-yaxis borckensis Besuchet & Kurbatov. 2007: 169 

87. Bryaxis brachati Besuchet. 1980b: 624 

88. Bryaxis corsus Besuchet. 1999b: 799 

89. Bryaxis credibilis Besuchet & Kurbatov. 2007: 182 

90. Bryaxis distinguendus Besuchet. 1961g: 1830 



CLAUDE BESUCHET' S WORK 



497 



91. Bryaxis effeminatus Besuchet, 1983b: 772 

92. Bryaxis egens Besuchet & Kurbatov, 2007: 196 

93. Bryaxis emendatus Besuchet & Kurbatov, 2007: 177 

94. Bryaxis euryscapus Besuchet & Kurbatov, 2007: 164 

95. Bryaxis festivus Besuchet, 1964a: 425 

96. Bryaxis focarilei Besuchet, 1980b: 623 

97. Bryaxis frustratus Besuchet, 1983b: 770 

98. Bryaxis gemellus Besuchet & Kurbatov, 2007: 202 

99. Bryaxis ghilarovi Besuchet, 1961g: 1829 

100. Bryaxis halbherri pacei Besuchet, 1983b: 777 

101 . Bryaxis herculinus Besuchet, 1962b: 351 

102. Bryaxis immodicus Besuchet & Kurbatov, 2007: 199 

103. Bryaxis ipsimus Besuchet & Kurbatov, 2007: 182 

104. Bryaxis jucundus Besuchet, 1961g: 1830 

105. Bryaxis karamane Besuchet, 1958e: 895 

106. Bryaxis khnzoriani Besuchet, 1964a: 423 

107. Bryaxis kovali Besuchet & Kurbatov, 2007: 193 

\08. Bryaxis krnegeri var. machulkai Besuchet, 1955: 277 (= B. kruegeri Machulka) 

109. Bryaxis kurnakovi Besuchet, 1960b: 398 

110. Bryaxis kuzmini Besuchet & Kurbatov, 2007: 194 

111 . Bryaxis laevipennis Besuchet & Kurbatov, 2007: 181 

112. Bryaxis lazistanicus Besuchet & Kurbatov, 2007: 198 

113. Bryaxis litoralis Besuchet & Kurbatov, 2007: 177 

114. Bryaxis longifrons Besuchet & Kurbatov, 2007: 173 

115. Bryaxis longulus inflatus Besuchet, 1983b: 778 

116. Bryaxis lurensis Besuchet, 2002: 212 

117. Bryaxis mirificus Besuchet, 1983b: 773 

118. Bryaxis monguzzii Besuchet, 1980b: 622 

119. Bryaxis multiplex Besuchet & Kurbatov, 2007: 199 

120. Bryaxis myops Besuchet & Kurbatov, 2007: 199 

121 . Bryaxis nebrodensis Besuchet, 1980b: 619 
722. Bryaxis nitidulus Besuchet, 1961g: 1829 

123. Bryaxis nivarius Besuchet & Kurbatov, 2007: 168 

124. Bryaxis obventicius Besuchet & Kurbatov, 2007: 174 
725. Bryaxis orcinus Besuchet & Kurbatov, 2007: 168 
126. Bryaxis osellai Besuchet & Kurbatov, 2007: 1 90 
727. Bryaxis oseticus Besuchet & Kurbatov, 2007: 196 
128. Bryaxis ossaeus Besuchet, 2008b: 245 

729. Bryaxis pachyscelis Besuchet & Kurbatov, 2007: 170 

130. Bryaxis pedemontanus Besuchet, 1958b: 74 

131 . Bryaxis polemon Besuchet & Kurbatov, 2007: 189 

132. Bryaxis ponîicus Besuchet & Kurbatov, 2007: 187 

133. Bryaxis porzenna var. ticinensis Besuchet, 1954: 436 (= B. porzenna Reitter) 

134. Bryaxis propinquus Besuchet & Kurbatov, 2007: 186 

135. Bryaxis pulchrotibialis Besuchet & Kurbatov, 2007: 172 



498 



P. HLAVÂC 



136. Bryaxis pygmaeus Besuchet & Kurbatov, 2007: 200 

137. Bryaxis rifensis Besuchet, 1962b: 351 

138. Bryaxis rivitlaris 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) 

14 3. Bryaxis schuelkei Besuchet & Kurbatov, 2007: 203 

144. Bryaxis silvicola Besuchet & Kurbatov, 2007: 187 

145. Bryaxis solarii Besuchet, 1958c: 8 (= B. rhinophorus W. Blattny & C. Blattny) 

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 troglodytes pierottii Besuchet, 1980b: 628 

152. Bryaxis tuberculiceps Nonveiller, Pavicevic & Besuchet, 2003b: 287 

153. Bryaxis viti Besuchet & Kurbatov, 2007: 163 

154. Bryaxis ypsilon Besuchet & Kurbatov, 2007: 169 

755. Bythinus confusus Besuchet, 1974c: 337 

756. Bythinus hauseri Besuchet, 1978a: 263 

757. Bythinus icariensis Besuchet, 1964a: 420 

158. Bythinus vicinus Besuchet, 1960a: 21 

159. Centrophthalmus klapperichi Besuchet, 1966b: 63 

160. Centrophthalmus mesopotamenus Besuchet, 1966b: 64 
767. Centrophthalmus sharpi Besuchet, 1966b: 61 

762. Centrophthalmus septentrionalis Besuchet, 1960a: 30 (= C.pici Jeannel) 

163. Centrotoma kaszabi Besuchet, 1969f: 301 

764. Claviger intermedius Besuchet, 1961c: 457 

765. Claviger pouzaui validus Besuchet, 1961c: 453 

766. Claviger saulcyi lucens Besuchet, 1961c: 455 

767. Colilodion incredibilis Besuchet, 1991: 503 
168. Colilodion mirus Besuchet, 1991: 506 
769. Colilodion inopinatus Besuchet, 1991: 507 
170. Colilodion concinnus Besuchet, 1991: 509 
777. Couloniella mirabilis Besuchet, 1983a: 510 

772. Decatocerus bicornis rotundatus Besuchet, 1961d: 95 
(= D. balearicus Jeannel, 1961) 

173. Decatocerus catalonicus Besuchet, 1961d: 94 

174. Decatocerus pityusensis Besuchet, 1958e: 909 

775. Desimia longicornis Besuchet, 1958d: 337 (= D. subtilipalpis Reitter) 

776. Dicentrius balcanicus balcanicus Besuchet, 1999c: 229 

777. Dicentrius balcanicus pirinensis Besuchet, 1999c: 229 
178. Dicentrius behnei Besuchet, 1999c: 228 

779. Dicentrius biroi Besuchet, 1999c: 230 



CLAUDE BESUCHET' S WORK 



180. Dicentrius discrepans Besuchet, 1999c: 231 

181 . Dicentrius ermischi Besuchet, 1999c: 231 

182. Dicentrius fodori Besuchet, 1999c: 228 

183. Dicentrius rousi Besuchet, 1999c: 231 

184. Dicentrius zerchei Besuchet, 1999c: 228 

185. Euplectus atlanticus Besuchet, 1962b: 346 

186. Euplectus caecus Besuchet, 1990: 161 

187. Euplectus canariensis Besuchet, 1968: 285 

188. Euplectus franzi Besuchet, 1968: 284 
\89. Euplectus frater Besuchet, 1964a: 415 

190. Euplectus insignis Besuchet, 1961a: 30 

191 . Euplectus kulzeri Besuchet, 1958d: 334 

192. Euplectus micropterus Besuchet. 1970a: 120 

193. Euplectus mussardi Besuchet, 1962b: 344 

194. Euplectus sarawakensis Besuchet, 1956c: 88 

195. Euplectus sexstriatus Besuchet, 1970a: 121 

196. Euplectus sparsus Besuchet, 1964a: 412 

197. Euplectus validus Besuchet, 1958e: 906 

198. Faronus andalusiacus Besuchet ,1969c: 114 

199. Faronus depressus Besuchet, 1960a: 15 

200. Faronus distinctus Besuchet, 1999b: 792 

201. Faronus festivus Besuchet, 1960a: 12 

202. Faronus festivus apterus Besuchet. 1960a: 13 (= F. festivus Besuchet) 

203. Faronus gracilis Besuchet, 1969c: 115 

204. Faronus insignis Besuchet, 1958e: 899 

205. Faronus lusitaniens Besuchet, 1969c: 111 

206. Faronus parallelus Besuchet. 1958e: 897 

207. Faronus parnassius Besuchet, 1969e: 397 

208. Faronus rifensis Besuchet, 1963c: 227 

209. Faronus sahlbergi Besuchet, 1960a: 11 (= F. parallelus Besuchet) 

210. Faronus testaceus Besuchet, 1962b: 336 

211. Faronus tingitanus Besuchet, 1962b: 334 
272. Faronus variabilis Besuchet, 1969c: 110 

213. Faronus venustus Besuchet, 1958d: 333 

214. Fustiger appendiculatus Besuchet, 1977b: 261 

215. Fustiger wittmeri Besuchet, 1977b: 264 

216. Geopselaphus affinis Besuchet, 1961f: 259 

217. Geopselaphus alticola Besuchet. 1 96 1 f: 260 

218. Geopselaphus balearicus Besuchet, 1969b: 104 

219. Geopselaphus depressus Besuchet, 1961f: 261 

220. Geopselaphus distinguendus Besuchet, 1961f: 254 
227. Geopselaphus espanoli Besuchet, 1969b: 99 

222. Geopselaphus formosus Besuchet, 1961f: 261 

223. Geopselaphus franzi Besuchet, 1 96 1 f: 260 

224. Geopselaphus frater Besuchet, 1969b: 100 



500 



P. HLAVÂC 



225. Geopselaphus jucundus Besuchet, 1961f: 256 

226. Geopselaphus lepidus Besuchet, 1969b: 102 

227. Geopselaphus longipalpis Besuchet, 1969b: 102 

228. Geopselaphus longulus Besuchet, 1961f: 250 

229. Geopselaphus mirandus Besuchet, 1961 f: 252 

230. Geopselaphus mussardi Besuchet, 1963c: 231 

231. Geopselaphus nitidus Besuchet, 1969b: 100 

232. Geopselaphus tingitanus Besuchet, 1962b: 364 

233. Glyphobythus fallax Besuchet, 1960c: 403 

234. Glyphobythus hervei Besuchet, 1960c: 404 

235. Halorabyxis gourvesi Besuchet, 1975c: 138 

236. Imirus outereloi Besuchet, 1980a: 56 

237. Lepioplectus perraulti Besuchet, 1993a: 340 

238. Lindeha picanyolae Besuchet, 1985a: 515 

239. Meliceha (Cyrtoplectus) italica Besuchet, 1966a: 56 

240. Mesoleptoehir rougemonti Besuchet, 1974a: 887 

241. Namunia cavernicola Besuchet, 1978b: 131 

242. Namunia lapidicola Besuchet, 1978b: 131 

243. Namunia terricola Besuchet, 1999b: 799 

244. Neopselaphus adisi Besuchet, 1982b: 801 

245. Neopselaphus armaîus Besuchet, 1982b: 801 

246. Neopselaphus curtipalpis Besuchet, 1987a: 237 

247. Neopselaphus degalieri Besuchet, 1982b: 806 

248. Neopselaphus filipalpis Besuchet, 1982b: 806 

249. Neopselaphus chalumeaui Besuchet, 1987a: 238 

250. Neopselaphus parki Besuchet, 1982b: 805 

251 . Neopselaphus tavakiliani Besuchet, 1982b: 804 

252. Nonveillera lepida Pavicevic & Besuchet, 2003a: 280 

253. Nonveillera romani Pavicevic & Besuchet, 2003a: 282 

254. Ocîomicrus denîifrons Besuchet, 1999b: 790 

255. Pachacuti huggerti Besuchet, 1987a: 235 

256. Panaphantus afer Besuchet, 1980c: 154 

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

258. Paratychus minutissimus Besuchet, 1960a: 26 

259. Plectophloeus binaghii Besuchet, 1964a: 417 

260. Plectophloeus erichsoni occidentalis Besuchet, 1969e: 399 

261 . Plectophloeus erichsoni orientalis Besuchet, 1969e: 400 

262. Plectophloeus nubigena bosnicus Besuchet, 1964a: 419 

263. Prionobythus genesti Besuchet, 1985a: 509 

264. Pselaphaulax carniolicus Besuchet & Sabella, 2000a: 263 

265. Pselaphaulax siculus Besuchet & Sabella, 1993c: 92 

266. Pselaphogenius laticeps Besuchet, 1961b: 262 

267. Pselaphogenius latinus Besuchet, 1980b: 630 

268. Pselaphogenius lepontinus Besuchet, 1980b: 629 



CLAUDE BESUCHET'S WORK 



501 



269. Pselaphogenius lucanicus Besuchet, 1964a: 435 

270. Pselaphogenius neapolitanus Besuchet, 1964a: 433 

271 . Pselaphogenius oriental i s Besuchet, 1961a: 39 

272. Pselaphostomus bergamascus Besuchet, 1980b: 628 

273. Pselaphostomus bussacensis estrellensis Besuchet, 1961b: 252 

274. Pselaphostomus franzi Besuchet, 1961b: 246 

275. Pselaphostomus intermedius Besuchet, 1961b: 249 

276. Pselaphostomus lusitaniens Besuchet, 1961b: 244 

277. Pselaphostomus pyrenaeus Besuchet, 1961b: 236 

278. Pselaphostomus stussineri vesulinus Besuchet, 1961a: 33 

279. Pselaphus mysius Besuchet, 1960b: 399 

280. Pselaphus xaymacus Besuchet, 1987a: 236 

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

282. Pygoxyon myops Besuchet, 1958e: 901 

283. Scotoplectus caspicus Besuchet, 1975b: 401 

284. Scotoplectus ponticus Besuchet, 1975b: 400 

285. Scotoplectus weiratheri Besuchet, 1975b: 399 

286. Seracamaurops fritschi Besuchet, 1986c: 4 

287. Syntectodes maldivicus Besuchet, 2008a: 73 

288. Tapas armifer Besuchet, 2008a: 77 

289. Tapas basseti Besuchet, 2008a: 76 

290. Tasmiger strumosus Besuchet, 2008a: 78 

291. Thelotia cebennica Besuchet, 1999b: 795 

292. Tremissus beaumonti Besuchet, 1982d: 320 

293. Tremissus inexspectatus Besuchet, 1982d: 319 

294. Triartiger nommai Besuchet, 2008a: 74 

295. Tribatus hauseri Besuchet, 1961a: 31 

296. Tribatus lopatini Besuchet, 1964a: 426 

297. Trimium atticum Besuchet, 1969e: 400 

298. Trimium illyricum Besuchet, 1969e: 402 

299. Trimium sardonm Besuchet, 1958c: 7 (= T. amplipenne Reitter) 

300. Trissemus bellax Besuchet, 1999b: 801 

301 . Trissemus holzschuhi Besuchet, 1999b: 805 

302. Trissemus maroccanus wittmeri Besuchet, 1981b: 245 

303. Trissemus micropterus Besuchet, 1970b: 314 

304. Trissemus mundulus Besuchet, 1961e: 2 

305. Trissemus sulcifrons Besuchet, 1999b: 803 

306. Trissemus trilobatus Besuchet, 1999b: 803 

307. Trogaster binaghii Besuchet, 1969d: 214 

308. Trogaster caprai Besuchet, 1969d: 216 

309. Trogaster doderoi Besuchet, 1969d: 213 

310. Trogaster gestroi Besuchet, 1969d: 215 

311. Trogaster solarii Besuchet, 1969d: 216 

312. Trogasteropsis coiffaiti Besuchet, 1977a: 291 

313. Tychobythinus atlanticus Besuchet, 1963b: 218 



502 



P. HLAVÂC 



314. Tychobythinus brachati Besuchet, 2008b: 246 

315. Tychobythinus curtii Besuchet, 1980b: 615 

316. Tychobythinus escalerai Besuchet, 1962b: 347 

317. Tychobythinus escolai Besuchet, 1974b: 49 

318. Tychobythinus espanoli Besuchet, 1974b: 53 

319. Tychobythinus listai Besuchet, 1985a: 514 

320. Tychobythinus muntani Besuchet, 1974b: 54 

321 . Tychobythinus naxius Besuchet, 1993b: 225 

322. Tychobythinus occidentalis Besuchet, 1962b: 349 

323. Tychobythinus rosai Besuchet, 1980b: 616 

324. Tychobythinus strinatii Besuchet, 1982a:50 

325. Tychobythinus urgellesi Besuchet, 1974b: 52 

326. Tychobythinus vignai Besuchet, 1978c: 69 

327. Tychomorphus franzi Besuchet, 1963b: 220 

328. Tychomorphus mussardi Besuchet, 1999a: 56 

329. Tychus affinis Besuchet, 1958e: 911 

330. Tychus altivagus Besuchet, 2011: 30 

331 . Tychus anatolicus Besuchet, 1964a: 429 

332. Tychus antalyanus Besuchet & Sabella, 1999e: 250 

333. Tychus asuniensis Besuchet, 1964a: 433 

334. Tychus atlanticus Besuchet & Sabella, 1999e: 239 

335. Tychus brachati Besuchet & Sabella, 1999d: 314 

336. Tychus caspicus Besuchet & Sabella, 1999e: 254 

337. Tychus coiffaiti Besuchet, 1958e: 913 

338. Tychus cordiger Besuchet, 1969e: 408 

339. Tychus distinguendus Besuchet, 1960a: 23 

340. Tychus epiroticus Besuchet, 1964a: 431 

341. Tychus georgicus Besuchet & Sabella, 1999e: 247 

342. Tychus holzschuhi Besuchet & Sabella, 1999d: 317 

343. Tychus judaeus Besuchet, 1964a: 428 

344. Tychus laminiger Besuchet, 1969e: 409 

345. Tychus latebrosus Besuchet, 201 1 : 26 

346. Tychus longicornis Besuchet, 1958e: 914 (= T. balcanicus Reitter) 

347. Tychus lusitanicus Besuchet & Sabella, 1999e: 237 

348. Tychus manicanus Besuchet & Sabella, 1999e: 238 

349. Tychus mundulus Besuchet, 1958e: 915 

350. Tychus paludivagus sicilianus Besuchet & Sabella, 1996: 111 

351 . Tychus pelopeius Besuchet & Sabella, 1999d: 312 

352. Tychus persicus Besuchet & Sabella, 1999e: 245 

353. Tychus ponticus Besuchet & Sabella, 1999e: 249 

354. Tychus remaudierei Besuchet, 1969e: 407 

355. Tychus rhodopeus Besuchet & Sabella, 1999d: 313 

356. Tychus sardous Besuchet, 1964a: 432 

357. Tychus sengleti Besuchet & Sabella, 1999d: 315 

358. Tychus striola andalusiacus Besuchet & Sabella, 1996: 107 



CLAUDE BESUCHET'S WORK 



503 



359. Tychus striola balearicus Besuchet & Sabella, 1996: 105 

360. Tychus viti Besuchet, 201 1 : 23 

361 . Tychus tingitanus Besuchet. 1962b: 364 

362. Tyrodes janetscheki Besuchet, 1970b: 316 

IV. List of species and subspecies of Scydmaenidae 

1. Cephennium (Cephennium) fraterculum Besuchet. 1971b: 278 

2. Cephennium (Cephennium) machulkai Besuchet, 1971b: 278 

3. Cephennium (Cephennium) paganetîii Besuchet, 1971b: 278 

4. Cephennium (Phennecium) galitense Besuchet, 1982e: 238 

5. Cephennium (Phennecium) solarii Besuchet. 1958e: 896 
new name for C. romanum Holdhaus, 1924: 21 

6. Cephennodes (s.str.) basilewskyi Besuchet, 1962c: 420 

7. Cephennodes (s.str.) indifferens Besuchet. 1962c: 422 

8. Cephennodes (s.str.) leleupi Besuchet, 1962c: 421 

9. Cephennodes (s.str.) marginaîus Besuchet, 1962c: 423 

10. Cephennomicrus fossulaîus Besuchet, 1961: 17 

1 1 . Cephennomicrus glaber Besuchet, 1961: 24 

12. Cephennomicrus impressus Besuchet, 1961 h: 15 

13. Cephennomicrus jucundus Besuchet, 1961 h: 23 

14. Cephennomicrus latipennis Besuchet. 1961 h: 21 

15. Cephennomicrus longicornis Besuchet, 1961 h: 18 

16. Cephennomicrus pauliani Besuchet. 1961 h: 19 

17. Cephennomicrus pusillus Besuchet. 1961 h: 24 

18. Cephennomicrus rugosicollis Besuchet, 1961 h: 19 

19. Cephennomicrus suturalis Besuchet, 1961 h: 22 

20. Cephennomicrus vadoni Besuchet, 1961 h: 16 

21. Cephennomicrus validus Besuchet. 1961h: 21 

22. Clidicus loebli Besuchet, 1971a: 254 

23. Clidicus mussardi Besuchet, 1971a: 255 

24. Clidicus quadricollis Besuchet, 1971a: 252 

25. Etelea tingitana Besuchet & Vit, 2004b: 341 

26. Leptocharis algericus Besuchet, 1958e: 916 

27. Nanophthalmus nonveilleri Besuchet & Vit, 2000b: 159 

28. Nanophthalmus serbicus Besuchet & Vit, 2000b: 159 

29. Neuraphes (Pararaphes) toumayeffi Besuchet. 1980d: 192 

30. Scydmaenus aelleni Besuchet, 1981c: 460 

31. Taurablepton asitawandas Besuchet, 1969g: 315 (Ablepton) 

32. Taurablepton rutash Besuchet, 1969g: 316 {Ablepton) 

V. List of species and subspecies of Ptiliidae 

1. Actidium reticulatum Besuchet. 1971c: 319 

2. Oligella intermedia Besuchet, 1971c: 320 

3. Ptiliolum stockmanni Besuchet, 1971c: 326 

4. Ptilium (Ptilium) cognatum Besuchet, 1971c: 323 



504 



P. HLAVÂC 



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 

VIL List of species and subspecies of Cerylonidae: 

1 . Axiocerylon baloghi Besuchet & Slipinski, 1988b: 908 

2. Axiocerylon bournei Besuchet & Slipinski, 1988b: 908 

3. Axiocerylon burckhardti Besuchet & Slipinski, 1988b: 908 

4. Axiocerylon decemcostatum Besuchet & Slipinski, 1988b: 922 

5. Axiocerylon ghanense Besuchet & Slipinski, 1988b: 919 

6. Axiocerylon gomyi Besuchet & Slipinski, 1988b: 920 

7. Axiocerylon hammondi Besuchet & Slipinski, 1988b: 916 

8. Axiocerylon humerale Besuchet & Slipinski, 1988b: 914 

9. Axiocerylon loebli Besuchet & Slipinski, 1988b: 923 

10. Axiocerylon luzonicum Besuchet & Slipinski, 1988b: 908 

1 1 . Axiocerylon minimum Besuchet & Slipinski, 1988b: 911 

12. Axiocerylon myops Besuchet & Slipinski, 1988b: 910 

13. Axiocerylon orousseti Besuchet & Slipinski, 1988b: 910 

14. Axiocerylon perkorum Besuchet & Slipinski, 1988b: 910 

15. Axiocerylon roberti Besuchet & Slipinski, 1988b: 916 

16. Axiocerylon solomonense Besuchet & Slipinski, 1988b: 913 

17. Axiocerylon triste Besuchet & Slipinski, 1988b: 911 

18. Axiocerylon variabile Besuchet & Slipinski, 1988b: 916 

19. Axiocerylon venustum Besuchet & Slipinski, 1988b: 919 

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 

31. Cautomus (Paracautomus) nitidus Besuchet, 1972: 136 

32. Cautomus (Paracautomus) reticulatus Besuchet, 1972: 133 

33. Glyptolopus amazonicus Besuchet & Slipinski, 1987b: 80 

34. Glyptolopus convexus Besuchet & Slipinski, 1987b: 78 

35. Glyptolopus peruanus Besuchet & Slipinski, 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: 
1. 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 Hlavâc 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 
Muséum for a period of two weeks identifying and checking what Claude regarded as 
ail the Leiodinae: Leiodini from Swiss institutional and private collections and sundry 
extra Swiss spécimens from non-Swiss muséums. 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 génial 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 it. I greatly enjoyed the hospitality Claude extended 
to me during my stay, which included practicalities of eating out in Geneva, visits to 
nature reserves in the Geneva area and the Jura as well as the Geneva Insekten Bôrse 
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 muséum 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. HLAVÂC 



5. Besuchet, C. 1954c. Une nouvelle espèce de Brachygluta du Danemark (Col. Pselaphidae). 

Entomologiste Meddelelser 27: 43-45. 

6. Besuchet, C. 1955a. Monographie des Bibloplectus et Pseudoplectus paléarctiques (Col. 

Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 28 (2): 
153-209. 

7. Besuchet, C. 1955b. Arcopagus picteti Tourn. et kriigeri Mach. (Col. Pselaphidae). 

Mitteilungen der Schweizerischen entomologischen Gesellschaft 28 (3): 274-278. 

8. Besuchet, C. 1955c. Bibloporus européens décrits par Jeannel (Col. Pselaphidae). Mit- 

teilungen der Schweizerischen entomologischen Gesellschaft 28 (3): 278. 

9. 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. 

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

Zoologie 63: 697-705. 

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

mologist 89: 88-90. 

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

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

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

Schweizerischen entomologischen Gesellschaft 29(3): 254. 

14. Besuchet, C. 1957. Contribution à l'étude des Rhipidiinae. Bulletin de la Société Vaudoise 

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

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

entomologischen Gesellschaft 31 (1): 65-69. 

16. 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. 

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

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

18. Besuchet, C. 1958d. Descriptions de quelques Psélaphides paléarctiques nouveaux 

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

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

(4): 891-919. 

20. Besuchet, C. 1959a. Quelques remarques concernant le genre Amauronyx Reitt. et 

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

21. Besuchet, C. 1959b. Coléoptères Psélaphides et Scydménides de la collection Cl. Rey. 

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

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

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

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

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

24. Besuchet, C. 1960c (1959). Petite revision du genre Glyphobythus Raffr. (Col. 

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

25. Besuchet, C. 1961a. Psélaphides paléarctiques. Espèces nouvelles et notes synonymiques 

(Coleoptera). Mitteilungen der Schweizerischen entomologischen Gesellschaft 34 (1): 
30-42. 

26. Besuchet, C. 1961b. Révision des Pselaphostomus et Pselaphogenius ibériques (Col. 

Pselaphidae). Eos, Madrid 37: 229-265. 



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27. Besuchet, C. 1961c. Révision des Claviger ibériques (Coleopt. Pselaphidae). Revue suisse 

de Zoologie 68: 443-460. 

28. Besuchet, C. 1 96 1 d. Révision du genre Decatocerus SAULCY (Col. Pselaphidae). 

Miscellanea Entomologica 1 (4): 91-97. 

29. Besuchet, C. 1961e. Deux Psélaphides nouveaux d'Iran (Coleoptera). Stuttgarter Beitràge 

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

30. Besuchet. C. 1 961 f (1960). Re vision du genre Geopselaphus Jeann. (Col. Pselaphidae). 

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

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

gicheskiy Zhurnal 40 ( 1 2): 1 827- 1 83 1 , in Russian. 

32. Besuchet, C. 1961 h. 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. 

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

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

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

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

35. Besuchet. C. 1962c. Coleoptera Scydmaenidae: Cephenniini. Mission zoologique de 

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

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

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

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

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

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

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

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

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

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

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

41. Besuchet, C. 1966b. Revision des Centrophthalmus paléarctiques. Mitteilungen der 

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

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

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

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

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

44. Besuchet, C. 1969a. Rybaxis SAULCY, 1876 (Insecta, Coleoptera): Proposed désignation 

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

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

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

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

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

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

Memorie délia Società Entomologica Italian 48: 211-218. 

48. Besuchet, C. 1969e. Psélaphides paléarctiques. Espèces nouvelles et notes synonymiques. 

III (Coleoptera). Revue suisse de Zoologie 76 (2): 397-420. 

49. Besuchet, C. 1969f. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der 

Mongolei. 180. Pselaphidae. Reichenbachia 11: 301-304. 



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

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

51. Besuchet. C. 1969h. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der 

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

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

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

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

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

54. Besuchet, C. 1971a. Les Clidicus de Ceylan (Col. Scydmaenidae). Mitteilungen der 

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

55. Besuchet, C. 1971b. 18.Familie: Scydmaenidae [1 . Tribus Euthiini, 2. Tribus Cephenniini] 

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

56. Besuchet, C. & Sundt. E. 1971c. In: Freude H.. Harde K. W. & Lohse G. A. (eds). Die 

Kàfer Mitteleuropas. Band 3. Adephaga 2. Palpicornia, Histeroidea. Staphylinoidea 1. 
Goecke & Evers, Krefeld, 365 pp. 

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

99-145. 

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

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

59. Besuchet. C. 1974b. Les Psélaphides cavernicoles de l'Espagne (Coleoptera Pselaphidae). 

Miscelanea zoologica 3 (4): 41-69. 

60. Besuchet. C. 1974c. 24. Familie: Pselaphidae (pp. 305-362). In: H. Freude, K. W. Harde, 

and G. A. Lohse (eds). Die Kàfer Mitteleuropas. Vol. 5, Staphylinidae II (Hypocyphtinae 
und Aleocharinae), Pselaphidae. Goecke & Evers, Krefeld, 381 pp. 

61. Besuchet. C. 1975a. Bihloplectus nouveaux de la région méditerranéenne orientale (Col. 

Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesellschaft 48 (1-2): 
31-36. 

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

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

63. Besuchet. C. 1975c. Un Halorabwis nouveau de la Polynésie française (Coleoptera 

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

64. Besuchet. C. 1976. Contribution à l'étude des Ptiliides paléarctiques (Coleoptera). 

Mitteilungen der Schweizerischen entomologischen Gesellschaft 49: 51-71. 

65. Besuchet. C. 1977a. Les Trogasteropsis de la Catalogne (Col. Pselaphidae). Nouvelle 

Revue d'Entomologie 7 (3): 291-293. 

66. Besuchet. C. 1977b. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen 

Muséums in Basel. Coleoptera: Fam. Pselaphidae Subf. Clavigerinae Gen. Fustiger 
BREND. mit Einschluss der indischen Arten. Entomologica Basiliensia 2: 261-267. 

67. Besuchet. C. 1978a. Un Bythinus cavernicole nouveau (B. hauseri) de la Grèce (Coleoptera 

Pselaphidae). Annales Musei Goulandris 4: 263-265. 

68. Besuchet. C. 1978b. Le genre Namunia Reitt. (Coleoptera. Pselaphidae). Revue Suisse de 

Zoologie 85 (1): 127-T33. 

69. Besuchet. C. 1978c. Le premier Psélaphide troglobie de la Turquie (Coleoptera). Quaderni 

di Speleologia, Circolo Speleologia Romano 3 (1978-79): 69-73. 

70. Besuchet. C. 1978d. Réintroduction en Suisse du Platypsyllus castoris Rits. (Coleoptera: 

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

71. 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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72. Besuchet, C. 1979. Description d'un nouveau Batrisus AUBÉ (Coleoptera, Pselaphidae). 

Revue suissse de Zoologie 86 (1): 279-283. 

73. Besuchet. C. 1980a. Le genre Imirus REITT. (Coleoptera, Pselaphidae). Nouvelle Revue 

d'Entomologie 10 (1): 51-58. 

74. Besuchet, C. 1980b. Contribution à l'étude des Coléoptères Psélaphides d'Italie et du 

Tessin. Revue suissse de Zoologie 87 (2): 611-635. 

75. Besuchet, C. 1980c. Le genre Panaphantus KIESW. (Col., Pselaphidae). Mitteilungen der 

Schweizerischen entomologischen Gesellschaft 53 (2-3): 151-155. 

76. 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. 

77. Besuchet, C. 1980e. Revision des Leptinus paléarctiques (Coleoptera: Leptinidae). Revue 

Suisse de Zoologie 87(1): 131-142. 

78. Besuchet, C. 1981a. Contribution à l'étude des Bathsodes paléarctiques (Coleoptera: 

Pselaphidae). Revue Suisse de Zoologie 88 (1): 275-296. 

79. Besuchet, C. 1981b. Insects of Saudi Arabia. Coleoptera: Fam. Pselaphidae. Fauna of 

SaudiArabia 3: 243-250. 

80. Besuchet, C. 1981c. Description d'un Coléoptère Scydménide cavernicole de la Nouvelle- 

Calédonie. Revue suisse de Zoologie 88(2): 459-461 . 

81. Besuchet, C. 1982a. Contribution à l'étude des Bythinini cavernicoles néarctiques (Cole- 

optera: Pselaphidae). Revue suisse de Zoologie 89 (1): 49-53. 

82. Besuchet, C. 1982b. Le genre Neopselaphus JEANN. (Coleoptera: Pselaphidae). Revue 

suisse de Zoologie 89 (3): 797-807. 

83. Besuchet, C. 1982c. Coléoptères rares des laisses de la Dranse (Haute-Savoie). Bulletin 

Romand d' Entomologie 1: 97-100. 

84. Besuchet, C. 1982d (1981). Un nouveau genre de Brachyglutini de la région méditer- 

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

85. Besuchet, C. 1982e. 238-240: In. Franz, H. Beitrag zur Kenntnis der Coleopterenfauna der 

Insel Galita. Sitzungsberichte der Ôsterreichischen Akademie der Wissenschaften, 
Mathematisch-Naturwissenschaftliche Klasse (Abt. I), 191 (5-10): 231-240. 

86. Besuchet, C. 1983a. Un nouveau genre de Pyxidicerini du sud de l'Inde (Coleoptera: 

Pselaphidae). Revue suisse de Zoologie 90 (2): 509-512. 

87. Besuchet, C. 1983b. Bryaxis nouveaux ou méconnus du Nord de l'Italie (Coleoptera: 

Pselaphidae). Revue suisse de Zoologie 90 (3): 769-780. 

88. Besuchet, C. 1983c. The species of Pselaphidae (Coleoptera) of the Hortobâgy National 

Park. In: The Fauna of the Hortobâgy National Park. Vol. 2. (Ed: Mahunka, S.) 
Akadémiai Kiadô, Budapest 197-198. 

89. Besuchet, C. 1985a. Bythinini cavernicoles nouveaux de France et d'Espagne (Coleoptera: 

Pselaphidae). Revue suisse de Zoologie 92 (2): 509-517. 

90. BesuchetT, C. 1985b. Le genre Acetalius SHARP (Coleoptera: Pselaphidae). Revue suisse 

de Zoologie 92 (3): 761-766. 

91. Besuchet, C. 1985c. 27. Coléoptères Psélaphides. 199: In Expédition Thai-Maros 85 - 

Rapport Spéléologique et Scientifique. Edit. Association Pyrénéenne de Spéléologie, 
Toulouse: 199. 

92. Besuchet, C. 1986a. Synonymes et homonyme nouveaux de quelques genres de Pséla- 

phides (Coleoptera). Revue suisse de Zoologie 93 (1): 257-264. 

93. Besuchet, C. 1986b. Coléoptères Psélaphides des îles du Cap Vert (Coleoptera: Psela- 

phidae). CFS - Courier, Forschurigsinstitut Senckenberg 81: 41-42. 

94. Besuchet, C. 1986c. Contribution à l'étude du genre Seracamaurops WINKLER. 1925 

(Coleoptera. Pselaphidae). Mitteilungen der Schweizerischen entomologischen Gesell- 
schaft 59: 459-463. 

95. Besuchet, C. 1987a. Pselaphini néotropicaux nouveaux (Coleoptera: Pselaphidae). 

Archives des Sciences (Genève) 40 (2): 231-240. 



510 



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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, A. 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). In: Lohse, G. A. & W. H. 

Lucht (eds). Die Kàfer 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 VIII. 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 

délie sottospecie maghrébine 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. 1999c. 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 délia 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. 1999e. Nouve specie di Tychus délia 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 espèces 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., Buckle, C, Brachat, V. & Besuchet, C. 2004a. Revision der Palà- 

arktischen Arten der Gattung Brachygluta Thomson, 1859 (Coleoptera, Staphylinidae) 
(1. Teil). Muséum d'histoire naturelle, Genève pp. 283. 



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117. Besuchet, C. & Vit, S. 2004b. Le genre Etelea Csiki, 1909 (Coeoptera: Scydmaenidae: 

Cephenniini). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 71: 
337-343. 

118. Lôbl I. & Besuchet, C. 2004c: Staphylinidae: Pselaphidae, pp. 272-329. In: I. Lôbl & A. 

Smetana (eds). Catalogue of Palearctic Coleoptera. Vol. 2. Stenstrup: Apollo Books, 
942 pp. 

119. Besuchet, C. 2004d: New nomenclatorial and taxonomic acts, and comments. Scyd- 

maenidae: Cephenniini (pp. 25-26). In: I. Lôbl & A. Smetana (eds). Catalogue of 
Palearctic Coleoptera. Vol. 2. Stenstrup: Apollo Books, 942 pp. 

120. Besuchet, C. 2004e: New nomenclatorial and taxonomic acts, and comments. Staphy- 

linidae: Pselaphinae (pp. 28-29). In: I. Lôbl & A. Smetana (eds). Catalogue of Palearctic 
Coleoptera. Vol. 2. Stenstrup: Apollo Books, 942 pp. 

121. Besuchet, C. 2004f: Scydmaenidae: Cephenniini (pp. 203-206). In: I. Lôbl & A. Smetana 

(eds). Catalogue of Palearctic Coleoptera. Vol. 2. Stenstrup: Apollo Books 942 pp. 

122. 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. 

123. Besuchet, C. & Bekchiev, R. 2007b: Description d'un nouveau Batrisodes de la Bulgarie 

(Coleoptera: Staphylinidae: Pselaphinae). Historia naturalis bulgarica 18: 75-80. 

124. Besuchet, C. 2008a. Synonymies et combinaisions nouvelles, revalidations et description 

de taxa nouveaux de Pselaphinae (Coleoptera: Staphylinidae). Mitteilungen der 
Schweizerischen Entomologischen Gesellschaft 81: 61-82. 

125. Besuchet, C. 2008b. Two new cavernicolous Bythinini from Greece (Coleoptera, Staphy- 

linidae, Pselapinae) (pp. 161-164). In: D. Pavicevic & 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: viii + 564 pp. 

126. Sabella, G., Besuchet, C. & Kurbatov, S. A. 201 1 . New species of Tychini from Turkey 

and Japan (Coleoptera: Staphylinidae: Pselaphinae). Zootaxa 2764: 22-34. 

II. Faunistics 

1. Besuchet, C. & Comellini, A. 1964b. Coléoptères endogés des environs de Genève. 

Mitteilungen der Schweizerischen entomologischen Gesellschaft 36 ((4) (1963)): 
313-320. 

2. Besuchet, C. & Comellini, A. 1965 (1964). Coléoptères endogés des environs de Genève. 

Proc. XIII Int. Congress ofthe Entomology, London 459-460. 

3. Besuchet, C. 1968. Répartition des insectes en Suisse. Influence des glaciations. Mit- 

teilungen der Schweizerischen entomologischen Gesellschaft 41 (1-4): 337-340. 

4. Besuchet, C. 1976. Coléoptères dans les laisses de la Dranse. Arve-Léman Nature 12: 1-6. 

5. Besuchet, C. 1977. Coléoptères endogés du Tessin. Mitteilungen der Schweizerischen 

entomologischen Gesellschaft 49: 305. 

6. Besuchet, C. 1982. Coléoptères rares des Laisses de la Dranse (Haute-Savoie). Bulletin 

Romand d'entomologie 1: 97-100. 

7. Besuchet, C. 1983. Coléoptères des Alpes suisses atteignant ou dépassant l'altitude de 

3000 m. Bulletin Romand d'entomologie 1: 167-176. 

8. Besuchet, C. 1985. Combien d'espèces de Coléoptères en Suisse. Bulletin Romand d'ento- 

mologie 3: 15-25. 

9. Besuchet, C. & Moeschler, P. 1986. Coléoptères cavernicoles dans des sources karstiques 

du Jura Neuchatelois (Suisse). Bulletin Romand d'entomologie 4: 155-158. 

10. 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. 



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11. Besuchet, C. 1993a. Ein neuer Kàferkatalog fur die Schweiz. Mitteilungen der 

Schweizerischen entomologischen Gesellschaft 43(3): 90-100. 

12. Besuchet, C. & Merkl, O. 1993b. Scydmaenidae, Ptiliidae and Pselaphidae (Coleoptera) 

from the Biikk National Park. In: The Fauna of the Biikk National Park. Hungarian 
naîural History Muséum, Budapest 99-103. 

13. Besuchet, C. 1994. Les insectes. Le Malagnou 2: 25-27. 

14. Besuchet, C. 1998. Des petits clandestins. Le Malagnou 4: 28-31 . 

III. Popular & Others 

1. Besuchet, C. 1957. Une technique nouvelle pour la préparation de l'édéage des 

Microcoléoptères. Mitteilungen der Schweizerischen entomologischen Gesellschaft 30: 
341-342. 

2. Besuchet, C. 1958. Un insecte parasite du castor. Bulletin Mensuel des Musées et 

Collections de la ville de Genève, 15me année, No. 10. 

3. Besuchet, C. 1962. Les myrmécophiles. Bulletin de la Société Vaudoise d'Entomologie, 3: 

13-16. 

4. Besuchet, C. 1971. Recherches entomologiques à Ceylan. La Revue "Musées de Genève" 

No. 116:3-6. 

5. Besuchet, C. 1973. Les Rhinogrades. La Revue "Musées de Genève" No. 137: 10-13. 

6. Besuchet, C. 1977. Les Atta, fourmis champignonnistes. La Revue "Musées de Genève" 

No. 172: 2-11. 

7. Besuchet, C. 1977. Les Atta, fourmis exotiques champignonnistes. Arve-Léman Nature, 

15: 11. 

8. Besuchet, C. 1978. Les Atta, Fourmis champignonnistes. L'Entomologiste, 34(6): 221-231. 

9. Besuchet, C. 1979. Que deviennent les Atta! La Revue "Musées de Genève" No. 196: 

15-18. 

10. Besuchet, C. 1979. Un insecte parasite du Castor. Arve-Léman Nature, 33: 15-17. 

11. WUEST, J. & Besuchet, C. 1981. La colonie d'Atta cephalotes du Muséum de Genève. 

Comptes rendus, Assemblée générale, Toulouse, Section française. Union internationale 
d'étude des Insectes sociaux, 131-132. 

12. 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. 

13. Besuchet, C. 1986. Les Animaux cavernicoles + Résultats d'une campagne de piègeage 

dans les Alpes vaudoises. Le Trou, Groupe Spéléo Lausanne, 44: 16-20. 

14. Besuchet, C. 1992. Vers un "Catalogue des Coléoptères de Suisse". Bulletin Romand 

d'Entomologie, 10: 49-55. 

15. Besuchet, C. & Rezbanyai-Reser, L. 1999. Angaben zur Verbreitung und Phànologie 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 teww/s-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 teww/s-group and new or little known species 
in other Zelotes groups (Gnaphosidae, Araneae). - Notes are given on 
mating mechanisms of Z. tennis and Z. semirufus in the Z. tenuis-gvowp. 
Eight species are recognized in the Z. tenuis-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 maie of Z. babunaensis, the maie 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 6 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. phshutovae and the types spécimens of Zelotes semi - 
rufus, Z. jïdvaster and Z. fulvopilosus, ail spider material was collected by myself. 
Measurements are in millimètres. 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 Muséum of Geneva (MHNG); the other material, if 
not indicated otherwise, remains in my collection. For détails on rearing and cryo- 
fixing of mated spiders, see Senglet (2004: 87). 

The typical leg spination according to Platnick & Shadab (1983) is: Femora; I, 
II dl 10, pOOl ; III, IV dl 10, pOl 1 , rOl 1 ; patella III rOlO; tibiae: III pl 1 1 , v222, rOl 1 ; IV 
plll,v222,rlll;metatarsi: I, II v200; III pl22, v221 , rll2; IV pl22, v220, rl22. Only 
différences to this pattern are given in the text. AME, ALE, PME, PLE and MOQ refer 



Manuscript accepted 03.06.20 1 1 



514 



A. SENGLET 



to anterior médian, anterior latéral, posterior médian, posterior latéral eyes and to the 
médian ocular quadrangle (eyes included). 

Terminology of génital structures follows Senglet (2004: 88-90). Additional 
comments on some structures are: 

Embolar radix: This structure (Figs 8-10; Senglet, 2004: 88, fig. la, ld) seems 
to be présent in many Zelotinae. In Trachyzelotes 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. Présent in différent 
Zelotinae gênera; originating from the prolateral base of the tegulum, it is usually 
connected to the embolus with a flexible joint. There is no connection to the embolus 
in the Zelotes thorelli-gxoxxp . 

Posterior sclerite of terminal apophysis: Where présent in Zelotinae, the termi- 
nal apophysis is a ventral sclerotized extension of the embolar base. The présence 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. thorelli-groups), the middle sector of the radix 
(Z. tenuis-group), or even the embolus (Z. baeticus-gvoup) . 

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. Présent only in conjunction with the posterior sclerite of the terminal 
apophysis, highly variable and often reduced, it seems that its basic function is 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 

1 Embolar radix with sclerotized connection to embolus; no dorsal apo- 
physis on embolar base Zelotes s. str., 2 

Embolar radix with a membranous connection to embolus; a dorsal 
apophysis présent 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 sclerite of terminal apophysis 

connected to posterior tegular base Z. subterraneus-group 

Différent 3 

3 Embolus not coiled 4 

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 

4 Elevated arched embolar base with a distad-directed embolus. Posterior 
sclerite of terminal apophysis connected to médian part of embolar 



NEW ZELOTES SPECIES 



515 



radix. Epigynum vvith single latéral folds; médian sector membranous . . 

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 latéral folds 

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 médian apophysis. Latéral folds of epigynum 
modified into large latéral pockets; médian part membranous (Figs 123, 
1 34) Z. baeticus-group 

Zelotes tenuis-group 

Définition: The maie 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 médian part of the radix in Z. subterra- 
neus and Z. longipe s-gvoups (Senglet, 2004: figs la, ld, 12-14), the connection is 
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 latéral epigynal folds enclosing a membranous cuticle. 
Except for Z. alpujarraensis, the type of ocular group shown in Figs 54-55 is shared 
by ail species of the Z. tenuis-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 insémination 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 médian 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 médian mem- 
branous part is found in Z. chaniaensis sp. n. and Z. fulvaster. Figures 5-11 show 
détails of the maie bulbus in copula. 



KEY TO THE SPECIES OF THE ZELOTES TENUIS-GROUP 

â 1 

9 8 

1 Apical margin of embolar base convex 3 

Apical margin of embolar base concave between distal tooth and 
embolus (Figs 12, 33) 2 



516 



A. SENGLET 




FlGS 1-4 

(1) Zelotes tenuis, epigynum in copula, maie palp extracted. (2-4) Zelotes semirufus, left maie 
palp extracted in copula. (2) Left maie palp, apical view (compare with Fig. 11). (3) Epigynum. 
(4) Broken left maie 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) Z.fulvaster 

3 A retrolateral projection on embolar base (Figs 61-62, 66-69) 7 

No retrolateral projection 4 

4 Retrolateral tibial apophysis ribbon-like, bifid (Fig. 20) Z.fuscorufus 

Retrolateral tibial apophysis tapering 5 

5 A small retrolaterally directed apical hook on embolar base (Fig. 53) . . 

Z. chaniaensis sp. n. 

A small triangular apical tooth on embolar base 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) Z. babunaensis 



NEW ZELOTES SPECIES 



517 




Figs 5-11 

Zelotes semirufus, left maie palp in copula, viewed from the female opisthosoma. (5) Anterior 
view. (6) Posterior view. (7) Left view. (8) Id., détail. (9) Right vievv. (10) Posterior view, détail. 
(11) Palp on female with vulva in dorsal view. Bold lines indicate female parts. Scale Unes 
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 

I Retrolateral embolar base projection slender, long, curved (Figs 66-69) 

Z. alpujarraensis sp. n. 

Retrolateral embolar base projection short, membranous (Figs 60-62) . . 
Z. manitchensis 

8 Epigynal plate much wider than long (Figs 21 , 63, 73 & 77) 14 

Epigynal plate longer than wide or slightly wider 9 

9 Latéral epigynal fold more or less curved, with a posterior rounded bend . . 10 
Latéral epigynal fold almost straight, with an acute posterior bend; 
latéral vulval pouch posteriorly widened; médian ducts connected on 
médian side of spermathecae (Figs 37-39, 75) 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 

II Short médian ducts leaving the médian epigynal cuticle undivided (Figs 

16-17) Z. semirufus 

Posterior part of médian cuticle divided by médian ducts and folds (Figs 
45-49) Z. tenuis 

12 Short médian ducts leaving the médian epigynal cuticle undivided (Figs 

70-71) Z. alpujarraensis sp. n. 

Posterior area of médian cuticle divided by médian ducts and folds 13 

13 Short curved médian ducts (Figs 57-58) Z. chaniaensis sp. n. 

Straight or slightly curved long médian ducts with an acute bend, close 

to the insémination pore (Figs 30-3 1 ) Z. babunaensis 

14 Short médian ducts (Fig. 21) Z.fuscorufus 

Long médian ducts (Figs 63-64) Z. manytchensis 



Zelotes semirufus (L. Koch, 1882) Figs 2-17, 74 

Prosthesima semirufa L. Koch, 1882: 636, pl. 20, fig. 15 (description of 9). 

Zelotes ruscinensis Simon, 1914: 157, 169, 219, figs 295, 346 (description of S) 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 Muséum (London); 1 9 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 6 (last moult 08.07.2002); 15.05.2002. - Estremadura, Caceres, west of 
Jarandilla (litter in végétation), 40°08'N 05°40'W; 1 $ (last moult 17.08.2009); 16.06.2009. - 
Andalusia, Cordoba, Almodovar del Rio (Brena dam), 37°50'N 05°04'W; 1 9 (last moult 
18.08.2009); 01 .06.2009. - Huelva, Puerto Gil /Aracena (cork-oak leaf litter), 37°53'N 06°29'W; 
1 9 (last moult 21.08.2009); 07.06.2009. - Huelva, Alajar /Aracena, 37°53'N 06°40'W; 6 S, 
5 9 (last moults of S 02 to 28.07.2009, of 9 12.07 to 23.08.2009); 09.06.2009. 

DlAGNOSIS: The maie 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 latéral pouches with folded cuticle, its narrow anterior epigynal 
margin and its widely separated médian ducts (Figs 12-17, 74). 



Description: See Senglet, 2004: 104. 



NEW ZELOTES SPECIES 



519 





Figs 12-21 

(12-17) Zelotes semirufus. (12-15) Left maie 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 maie palp. (18) Ventral 
view. (19) Prolateral view. (20) Tibia, retrolateral view. (21) Epigynum (on left) + vulva, dorsal 
view. Bold lines indicate epigynal folds. Scale Unes 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 S & ?). 

Zelotes fuscorufus. - Senglet, 2004: 102, figs lb-c, 32-34, 51-54. 

For redescription and previous literature, see Senglet (2004: 102) and Platnick (2010). 

DlAGNOSIS: The maie of Z. fuscorufus is easily distinguishable from maies 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). 



here removed from the synonymy of Z. tenuis 

Echemus babunaensis Drensky, 1929: 5, 59, pl. 1 , figs 3-4 (description of 9 ). 
Zelotes babunaensis. - Deltshev & Blagoev, 2001: 110 (transfer of 9 from Echemus). 
Zelotes tenuis. - Deltshev, 2003: 137, fig. 8, not figs 9-10 = Z. tenuis (synonymy of 9 , misiden- 

tification).-Chatzaki etal.,2003: 80, figs 108-110, 113-114 (misidentification), not figs 

105-107, 1 11-112 = 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 Neraïda, 
39°31'N 20°24'E; 1 S; 21.06.1998. - Thessalia, Larissa, Omolion-Tempé, 39°53'N 22°37*E; 
1 S (1 palpus lost), 1 9; 17.06.1970. - Sterea Hellas, Attiki, Marathon lake, 38°10'N 23°54'E; 

1 9 (with vulva in microvial); 11.07.1968. - Phthiotidas, Tràgana, 38°37'N 23°07'E; 1 S; 
13.07.1968. - Phthiotidas, Theologos, 38°39'N 23°12'E; 1 S , 2 9 (with vulva in microvial, last 
moults of $ 22.5, of 9 27.05 and 25.06.1998); 20.05.1998. - Eubea, Loutra Aidipsou, 38°53'N 
22°59'E; 1 â; 29.05.1983. - Peloponnesus, Laconia, east of Lira (in Pinus litter), 36°39'N 
22°58'E; 1 â, 1 9 (vulva in microvial, last moults of â 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 S (last moult 
05.06.2005); 01.10.2004. - Argolida, Palaia Epidauros, 37°39'N 23°09'E; 1 9 (vulva lost); 
05.06.1981. - Cyclades, Naxos, north-west of Sangri, 37°03'N 25°27'E; 1 S (last moult 
06.06.1998); 05.06.1998. - Cyclades, Koronido (650-750m), 37°09'N 25°37'E; 1 6, 2 9 (last 
moult of 1 9 28.06.1998); 08.06.1998. - Cyclades, Kato Potamia, 37°06'N 25°26'E; 2 6, 1 9 
(last moult of 9 12.06.1998); 10.06.1998. - Cyclades, Dimitra /Ano Sangri, 37°02'N 25°26'E; 

2 0,1 9 (last moults of 1 S 12.06, 9 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 vOOO; II v220. - â 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. tenuis) and 
width of pedipalpal tibia less than dorsal length of tibia (Fig. 28). Embolus short (Figs 
22-24). Scutum 1/5 of opisthosoma length. - 9 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. MOQ 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 latéral folds. Vulva (Figs 30-31): Long médian 
ducts, straight or slightly curved, with a sharp bend, close to the insémination pore. 



Zelotes babunaensis (Drensky, 1929), 



Figs 22-31,76 



NEW ZELOTES SPECIES 



521 




Figs 22-31 



Zelotes babunaensis. (22-28) Left maie palp. (22) Ventral view (médian apophysis expanded), 
from Crète. (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. 



522 



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. tennis, 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 9). 

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 9 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 (végétation), 40°41*N 23°05'E; 2 <?, 1 9 (last moults of S 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 6; 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 S (last 
moult 09.06.1998); 21.05.1998. - Same; 1 ô\ 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°47'E; 1 â, 1 9 (last moult of 9 28.06.2008); 11.06.2008. - Peloponnesos, Argolida, Palaia 
Epidauros,37°39'N23 o 09'E; 1 9 (last moult 29.07.1998); 14.05.1998. - Cyclades, Naxos, north 
of Aghios Prokopios (under végétation), 37°05'N 25°21'E; 2 6 (with palpus in microvial), 
1 9 (last moults of S 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, 
Falâvariân, 32°34'N 51°31'E; 1 S; 14.06.1974. - Fars, Khohkiluyeh, Arow, 30°35'N 50°43'E; 
1 ô\ 2 9 (with palpus and vulva in microvial); 24.05.1974. - Khohkiluyeh, Basht, 30°20'N 
51°15*E; 2 9; 25.05.1974. - Khohkiluyeh, Bishâpour, 29°47'N 51°35*E; 1 6; 28.05.1974. - 
Khohkiluyeh,Aliâbâd,30°01'N53°00'E; 1 9 ; 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 spécimens entirely tawny. - S from Naxos (in parenthèses 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). Médian apophysis with a large rounded concave retrolateral flap. Small hook- 
shaped, retrolaterally directed apical projection présent 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 vOOO; some large Iranian spécimens 
with additional retrolateral spines on metatarsi III & IV. - 9 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 



523 




Figs 32-39 

Zelotes fulvaster. (32-36) Left maie palp. (32) Prolateral view. (33) Ventral view. (34) Id., Iranian 
spécimen. (35) Retrolateral view. (36) Apical view. (37) Epigynum. (38) Vulva, dorsal view. (39) 
Id., Type "PM52" in MNHN Paris. Bold Unes indicate epigynal folds. Scale lines 0.2 mm. 



vvidened. Vulva (Figs 38-39): Latéral 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 vOOO. 

Zelotes tenuis (L. Koch, 1866) Figs 40-49, 72 

Drassus tenuis L. Koch, 1866: 101, pl. 4, figs 65-66 (description of S). 

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 9) syn. n. 



524 



A. SENGLET 



Zelotesfuscotestaceus.- 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 S and 9). - Chyzer 

& Kulczynski, 1897: 205, pl. 8, figs 2, 20. 
Zelotescircumspectus.- 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 3 and 9). 
Zelotes pallidus. - Platnick & Shadab, 1983: 185, figs 259-262 (description of S and 9). 

Material examined: GREECE, Macedonia, Thessaloniki, Aghios Vassilios, 40°40'N 
23°07'E; 1 SA 9 (last moults of 9 18-27.06.1998); 14.06.1998. -Thessalia, Larissa, Omôlion- 
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 9 (last moult 18.06.2005); 04.06.1978. - West of 
Malesina,38°37'N23°13'E; 1 S; 25.09.2004. - Same; 1 9 (last moult 12.06.2008); 08.06.2008. 
-SPAIN,Catalonia,Barcelona,Gelida,41 o 27'N0r5rE; 1 (with darker opisthosoma, 1 juv.); 
14.06.1971. - Levant / Murcia, Valencia, La Albufera, 39°21'N 00°19'W; 2 S; 16.06.1971. - 
Montroy, 39°20'N 00°35'W; 1 9; 22.06.1971 . - Alicante, Elche (under palm grove), 38°17'N 
00°42'W; 4 cT, 3 9 (last moults of 1 S 05.06.2002, of 9 11-14.06.2002); 15.05.2002. - Elda, 
38°30'N 00°47'W; 2 S\ 19.06.1971. - Murcia, Archena, 38°07'N 01°17'W; 1 9 ; 30.06.1971 . - 
Same; 2 S , 2 9 (last moults of S 26.05.2002, of 9 10 and 21 .07.2002); 17.05.2002. - Castilla 
/ Léon, Valladolid, Penafiel, 41°35'N 04°08'W; 1 S ; 23.06.2002. - Nueva Castilla / La Mancha, 
Madrid. Navalcarnero, 40°18'N 03°56'W; 2 S; 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 S; 14.06.1969. - Toledo, Cardiel de los Montes (rio Alberche), 40°02'N 04°39'W; 
1 (5,1 9; 15.06.1969. - Toledo, Urda /Consuegra, 39°25'N 03°42'W; 2 9; 12.08.1969. - 
Albacete, Hellin, 38°29'N 01°37'W; 1 6 ; 29.06.1971 . - Albacete, La Gineta (rio Jucar, fine leaf 
litter). 39°10'N 01°58'W; 6 S ; 16.05.2002. - Ciudad Real, Caracuel (laguna), 38°50'N 04°04'W; 
3 9; 04.08.1969. - Estremadura, Caceres, Jarandilla (rio Tietar. in litter on sand), 40°01'N 
05°37'W; 1 SA 9 (last moult of 1 9 03.08.2009); 15.06.2009. - Caceres, west of Jarandilla (in 
litter in végétation), 40°08'N 05°40'W; 2 6\ 1 9 (last moult of 9 22.06.2009); 16.06.2009. - 
Badajoz, south of Venta del Culebrin /Monesterio, 37°58'N 06°14'W; 1 S\ 19.06.1969. - 
Badajoz, Venta del Culebrin /Monesterio, 38°01'N 06°13'W; 11 ô\ 1 9; 19.06.1969. - Same; 

3 S (last moults of 2 S, 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 S (last moult of 1 S 11.06.2002); 09.06.2002. - Badajoz, south of 
Monesterio (under stones in végétation), 38°03'N 06°14'W; 10 S , 5 9 (last moults of 3 S 
6-12.06.2010, of 4 9 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 SA 9; 
25.05.2002. - Granada, La Râbita /Albunol, 36°45'N 03°10'W; 3 9; 09.07.1971. - Ugijar, 
36°59'N 03°04'W; 1 9; 15.07.1971. - Jaen, Canada de las Hazadillas (under Pinus), 37°39'N 
03°43'W; 3 ô\ 1 9 (last moults of S 03.06.2002, of 9 11.07.2002); 30.05.2002. - Cordoba, 
Palma del Rio; 1 6 \ 03.06.1967. - Cordoba. Palma del Rio, 37°43'N 05°18'W; 4 cT, 3 9; 
26.06.1969. - Cordoba, Almodovar del Rio (Brena dam). 37°50'N 05°04'W; 1 9 ; 28.06.1969. - 
Cordoba, Pantano de la Brena (evergreen oak litter), 37°51'N 05°04'W; 5 â , 8 9 (last moults of 

4 â 2-12.06.2002, of 9 02.6-16.07.2002); 01.06.2002. - Cordoba, Almodovar del Rio (Brena 
dam). 37°50'N 05°04'W; 1 S (last moult S 14.06.2009); 01.06.2009. - Cordoba, Penarroya, 
38°17'N 05°16'W; 1 S, 3 9 (with vulva in microvial); 30.06.1969. - Cordoba. Los Villares 
/Cordoba (Eucalyptus litter), 37°58'N 04°49'W; 1 S (last moult 02.06.2009); 31.05.2009. - 
Cordoba, Palma del Rio (Retortillo dam, leaf litter), 37°51'N 05°22'W; 1 S\ 03.06.2009. - 
Malaga. Ronda. 36°46'N 05°13'W; 1 9; 21.07.1969. - Malaga, Estepona, 36°25'N 05°11'W; 
1 9 ; 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 S\ 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 9 ; 24.06.1969. - Same, 37°43'N 05°53'W; 
8 S , 10 9 (with palpus and vulva in microvial, last moults of 3 S 09.6-9.07.2002, of 6 9 07.6- 
27.07.2002); 02.06.2002. - Sevilla. Cantillana. 37°37'N 05°50'W; 1 S\ 26.06.1969. - Sevilla, 
Alanis, 38°02'N 05°11'W; 1 S; 01.07.1969. - Sevilla, Sanlûcar la Mayor. 37°22'N 06°14'W; 



NEW ZELOTES SPECIBS 



525 




Figs 40-49 

Zelotes tenuis. (40-44) Left maie palp. (40) Greek spécimen, ventral view. (41) Id., retrolateral 
view. (42) Spanish spécimen, ventral view. (43) Id., retrolateral view. (44) Id., cleared, dorsal 
view. (45) Greek spécimen, epigynum. (46) Id., vulva, dorsal view. (47) Spanish spécimen, epi - 
gynum. (48) Id., vulva, dorsal view. (49) Id., variant. Bold lines indicate epigynal folds. Scale 
lines 0.2 mm. 



526 



A. SENGLET 



4 9; 17.07.1969. - Sevilla, Coripe (rio Guadalete), 36°58'N 05°26'W; 1 9; 18.07.1969. - 
Sevilla, east of Cazalla de la Sierra (deep leaf litter), 37°57'N 05°45'W; 2 6 , 3 9 (last moults of 

1 6 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 6, 3 9 (last moults of 2 6 5 and 12.06.2009, of 9 21.7-02.08.2009); 
05.06.2009. - Cadiz, Algodonales, 36°53'N 05°27'W, 1 9 , 19.07.1969. - Huelva, Santa Olalla 
(rio Cala), 37°55'N 06° 1 l'W, 2 9 , 04.07.1969. - Same, 2 6 , 05.06.2002. - Huelva, Zufre (rivera 
Huelva), 37°51*N 06°19'W, 1 9,04.07.1969.-Huelva,ZalamealaReal,37°41 , N06°39 , W, 1 ô\ 

2 9, 10.07. 1969. -Huelva, Moguer, 37°16'N 06°50'W, 1 9 , 11.07.1969. -Huelva, Torre de Oro 
/Mazagon, 37°05'N 06°43'W, 1 9 , 13.07.1969. - Huelva, Alajar /Aracena, 37°52'N 06°40'W; 
2 6 , 1 9 ; 07.07.1969. - Same (cork-oak leaf litter), 37°52'N 06°41'W; 1 6, 1 9 (last moult of 
9 08.08.2009); 09.06.2009. - Huelva, Puerto Gil /Aracena (cork-oak leaf litter), 37°53'N 
06°29'W; 10 6 (3 large maies, others small), 3 9 (last moults of 2 9 17.06 and 20.07.2009); 
07.06.2009. - Huelva, Fuenteheridos (deep leaf litter), 37°54'N 06°40'W; 3 6 , 3 9 (last moults 
of 1 6 14.06.2009, of 9 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 6; 02.06.1971. - Nord Corse, Above Ponte Leccia (under gravel, river); 4 6 , 10 9 ; 
20.06.1999. - Nord Corse, Above Barchetta (Golo river, under dry végétation); 2 6, 2 9; 
21.06.1999. - Nord Corse, Biguglia pond /Bastia; 1 9; 23.06.1999. - Sud Corse, Ajaccio 
(Gravone river); 1 6\ 01.06.1971. - Sud Corse, Portigliolo /Propriano (under végétation & 
stones); 1 9 ; 17.06.1999. - Sud Corse, Sartène, Acorane bridge; 3 6 , 2 9 (last moults of 9 20 
and 22.06.1999); 19.06.1999. - Provence / Côte d'azur, Var, Porquerolle (in litter), 43°00'N 
06°12'E;2 6,4 9 (last moults of 6 13.06.2001, of 9 19.07.2001); 23.06.2001 .- Languedoc, 
Hérault, Les Ouglou /Agde, (Etang de Thau), 43°20'N 03°33'E; 3 9 (last moult of 1 9 
08.07.2001); 26.06.2001. - Pyrénées Orientales, Arles sur Tech, 42°27'N 02°36*E; 1 9; 
01.07.2001. - ITALY, Sardinia, Sassari, Platamona /Porto Torres (damp leaf litter); 6 6 , 4 9 ; 
25.05.1999. -Sassari,Castelsardo; 8 6,5 9 (last moults of 2 6 5 and 20.06.1999, of 4 9 05.06- 
03.07.1999); 26.05.1999. - Sassari, Stagno di Calich /Alghero; 3 6 , 3 9 ; 29.05.1999. - Sassari, 
Porto di Vignola; 1 9 (last moult 27.07.1999); 12.06.1999. - Nuoro, lago alto de Flumendosa; 
8 6, S 9 (last moults of 6 11-27.06.1999, of 7 9 10.6-31.07.1999); 08.06.1999. - Nuoro, Cala 
Ginepro /Orosei (under Juncus); 1 6, 3 9 (last moults of 9 13-16.06.1999); 10.06.1999. - 
Cagliari, Quartu (laguna); 1 9; 15.09.1968. - Same; 6 6, 14 9 (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 
vOOO; II v220. - 6 from Greece (from Spain in parenthèses): 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 .08 (0 . 1 0) , ALE . 1 1 (0 . 1 4) , PME . 1 (0 . 1 4) , 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. 9 : From Greece (from 
Spain in parenthèses): 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 0.01 (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 



527 



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 
latéral 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 spécimen 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, Crète, Chania, Episkopi, 35°30'N 23°46'E; 6 (with palpus in 
microvial, last moult 04.01 .2000); 10.10.99. 

Paratype: Same locality and collecting date as for holotype; 1 9 (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 Crète. 

DlAGNOSlS: The maie differs from that of Z. tenuis by a hook-like distal embo- 
lar base tooth (Figs 50-53); the female with shorter médian ducts; anterior anchoring 
pockets wider than latéral folds and folded cuticle in médian 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. - â: 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 0.01 , 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. - 9 : 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. MOQ 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 endémie replacing Z. tenuis on 
that island. 

Zelotes manytehensis (Ponomarev & Tsvetkov, 2006) comb. n. Figs 59-64, 77 

Trachyzelotes manytehensis Ponomarev & Tsvetkov, 2006: 11, figs 18-19 (description of 6 
and 9). 

Material examined: IRAN, Khohkiluyeh, Dogonbadân, 30°22'N 50°47'E; 1 6,2 $ 
(with palpus and vulva in microvials); 21.05.1974. 

DlAGNOSlS: S pedipalp (Figs 59-62): The retrolateral loop of the embolar base, 
which reaches far below the level of terminal apophysis, and its membranous rétro- 



528 



A. SENGLET 




Figs 50-58 

Zelotes chaniaensis sp. n. (50-53) Left maie palp. (50) Prolateral view. (51) Ventral view. (52) 
Retrolateral view. (53) Embolar base, ventral view. (54) Maie, ocular group. (55-58) Female. 
(55) Ocular group. (56) Left chelicera, ventral view. (57) Epigynum. (58) Vulva, dorsal view. 
Bold Unes indicate epigynal folds. Scale lines 0.2 mm. 



NEW ZELOTES SPECIES 



529 




Figs 59-64 

Zelotes manytchensis. (59-62) Left maie palp. (59) Prolateral view. (60) Ventral view. (61) 
Retrolateral view. (62) Apex of palpai organ, cleared, dorsal-retrolateral view. (63) Epigynum. 
(64) Vulva, dorsal view. Bold lines indicate epigynal folds. Scale lines 0.2 mm. 



530 



A. SENGLET 



latéral projection distinguish Z. manytchensis from the other species of the tenuis- 
group. 9 : Epigynum (Figs 63, 77): latéral folds wide apart, with large convex latéral 
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. - S\ 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 0.01, 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 vOOO. - 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 0.01 , 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 latéral pouches. Vulva (Fig. 64) with almost straight 
médian 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; 6 ; 
26.05.2002. 

Paratypes: Same locality as for holotype; 1 6 , 3 9 (with palpus and vulva in microvial) 
(last moults of 2 9 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 III r 1 1 1 , IV r2 1 1 ; metatarsus II v220. - S 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 




Figs 65-71 

Zeloîes alpujarraensis sp. n. (65-69) Left maie palp. (65) Prolateral view. (66) Ventral view. (67) 
Retrolateral view. (68) Apex of palpai organ, cleared, dorsal view. (69) Id., apical view. (70) 
Epigynum. (71) Vulva, dorsal view. Bold Unes indicate epigynal folds. Scale Unes 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 médian ducts 
and large spermathecae. 

Remarks: The genitalia of this large and dark species correspond to those of 
the Z. tenuis-group, especially in the maie pedipalpal structure. The maie posterior 
ocular row differs from that of the Z. tenuis-group (Figs 54-55) in having smaller eyes 
of equal size separated by 0.7 of their diameter. 

Zelotes thorelli-group 

Définition: 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-GROUV 

1 Large embolar base projection (Figs 81, 89, 100). Epigynum with 

médian posterior notch thorelli-subgroup, 2 

Short embolar base projection (Figs 108, 115). Epigynum with médian 
posterior protrusion fulvopilosus-subgroup, 5 

2 Short posterior epigynal notch; short convolution of médian vulval duct 

(Figs 85-86). Slim transverse embolar base projection (Fig. 81) 4 

Médian posterior epigynal notch occupying 1/3 or more of epigynal 
plate; convolution of médian 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 

narrow (Figs 89, 95) Z. thorelli 

Embolar base projection narrow, ribbon-like; embolus with two narrow 
apophyses (Figs 98, 100-101). Posterior epigynal notch triangular 

(Fig. 102) Z. laconicus sp. n. 

4 Pedipalp widened, with narrow trans verse retrolaterad-directed embolar 
projection (Figs 81-83). Copulatory duct coiled in numerous narrow 

turns (Figs 86-87) 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 

few turns Z. lagrecai 

Copulatory duct coiled in numerous wide turns (Di Franco, 1994: fig. 9; 
Levy, 2009: figs 65-66). Maie unknown Z. pediculatus 

5 One-loop retrolateral embolar coil almost reaching base of bulbus. 
Latéral epigynal pouches elongated, in a posterior position (Figs 106- 

107, 109) Z.flagellans 

One-loop retrolateral embolar coil reaching mid-length of bulbus. Short 
latéral pouches extended laterally (Figs 113-114, 116) Z.fulvopilosus 



NEW ZELOTES SPECIES 



533 




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. pediculaîoides sp. n., 
Z. lagrecai and probably Z. pedicitlatus (maie unknown). 

Zelotes pediculatoides sp. n. Figs 80-87 

Holotype: SPAIN, Levant / Murcia, Archena, 38°07'N 01°17'W; S (last moult 
14.09.2002); 17.05.2002. 

Paratypes: Same locality as for holotype; 1 S (with palpus in microvial), 1 9 (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), 40°01'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 9; 
16.06.1971 . - Castellon, Villanueva de Alcolea, 40°14'N 00°03'E; 1 9 ; 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 végétation); 1 9; 18.05.2002. - Nueva 
Castilla /La Mancha, Madrid, Navalcarnero, 40°18'N 03°56'W; 1 9; 12.06.1969. - Madrid, 
AldeadelFresno(rioAlberche),40°19 , N04°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, II entirely scopulate. Posterior eye 
row straight to slightly recurved. - c? 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 0.01, PME-PME 0.05, 
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). Basai 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 S and 9). - 
Jézéquel, 1962: 525, fig. 5 (description of 9). 

Material examined: SPAIN, Catalonia, Gerona, Port de la Selva-Qadaquès (under 
Cistus),42 o 18'N03 o 13'E; 1 c?,3 9 (last moult of 6 26.09.2004): 23.05.2004. - Gerona, north 



NEW ZELOTES SPECIES 



535 




Figs 80-87 

Zelotes pediculatoides sp. n. (80-84) Left maie palp. (80) Prolateral view. (81) Ventral view. (82) 
Retrolateral view. (83) Apex of palpai 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. 



536 



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 9 ; 24.05.2004. - Castilla / Léon, Valladolid, 
Penafiel, 41°35'N 04°08*W; 1 9 (last moult 23.08.2002); 23.06.2002. - Avila, Arenas San 
Pedro; 1 â (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 S, 1 9 (last moult of â 11.09.2002); 
16.05.2002. - Estremadura, Caceres, Montànchez, 39°14'N 06°08'W; 1 9; 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 S, 2 9 (last moults of $ 09.09.2002, of 1 9 
04.10.2002); 19.05.2002. - Granada, Puebla de Don Fadrique (Pinus + rosemary litter), 38°00'N 
02°27'W; 1 9 (last moult 09.09.2002); 19.05.2002. - Granada, La Vidriera / Pinar pass, 38°03'N 
02°34'W; 1 â , 3 9 (last moult of Ô 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 <?, 1 9 (last moult of 9 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 â , 2 9 (last moult of 1 â 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, Canada de las Hazadillas (under Pinus), 37°39'N 03°43'W; 1 â , 3 9 
(last moults of â 25.09.2002, of 1 9 02.10.2002); 30.05.2002. - Jaen, Puente de la Sierra, 
37°41'N 03°46'W; 1 9; 31.05.2002. - Cordoba, Brena dam (in evergreen oak litter), 37°51'N 
05°04'W; 1 â (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 9 (with vulva in microvial); 07.07.1969. - Same; 2 â, 2 9 (last moults of 6 17- 
27.09.2002, of 1 9 23.10.2002); 07.06.2002. - Huelva, north of La Nava (schist litter), 38°00'N 
06°45'W; 1 9; 09.06.2002. - Huelva, Santa Olalla (rio Cala), 37°55'N 06°11'W; 1 9; 
04.07.1969. - FRANCE, Provence /Cote D'Azur, Basses-Alpes, La Pourcine /Limans, 43°59'N 
05°44'E; 2 9; 21.06.2001. - Basses-Alpes, Aubenas-des- Alpes, 43°57'N 05°48'E; 1 9; 
21.06.2001. - Languedoc, Gard, Vénéjan; 1 9 (with vulva in microvial); 01.10.1998. - Gard, 
Aude, Bedos pass /Mouthoumet, 42°58'N 02°34'E; 1 9; 30.05.2004. - Eastern Pyrénées, 
Banyuls sur Mer, 42°28'N 03°07*E; 1 6 (last moult 28.08.2001); 27.06.2001. - Eastern 
Pyrénées, Les Abeilles /Banyuls, 42°28'N 03°04'E; 1 ô\ 2 9 (last moult of 6 01.10.2001); 
28.06.2001 . - Eastern Pyrénées, Cerbère (under stones, in Cistus), 42°27'N 03°09'E; 2 â , 3 9 
(last moults of S 1-24.09.2004, of 1 9 29.09.2004); 22.05.2004. - Eastern Pyrénées, 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; 16,12 (last moult of S 17.09.2001); 08.07.2001. - Same (oak 
litter); 1 ô\ 1 9 (last moults of â 19.08.2004, of 9 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. - â : 
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 0.01 , 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. Médian 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 révolution, 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 



537 




Figs 88-96 

(88-95) Zelotes thorelli. (88-94) Left maie palp. (88) Prolateral view. (89) Ventral view. (90) 
Retrolateral view. (91) Apex of palpai 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 Unes 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 médian sector. 
Vulva (Fig. 94) characterized by coiled anterior part of médian 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 cT and 9). 

Material examined: SPAIN, Andalusia, Almeria, Adra (La Albufera), 36°45'N 
02°57'W; 1 9 (with vulva in microvial); 08.07.1971. 

Remark: Di Franco's (1994: fig. 6) drawing of the maie clearly shows that this 
species belongs to the Z. thorelli 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; 6; 04.10.2010. 

Paratypes: Same locality as for holotype ; 1 S, 2 ? (with vulva and palpus in 
microvial). - 1 9 , 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. - â 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 0.01, PME-PME 0.04, PME-PLE 0.04, ALE-PLE 0.06. MOQ length 0.21, front 
width 0.3 1 , 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 
tibial 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 internai 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 basai embolar apophysis also présent. Scutum occupying 40% of 
opisthosoma length. - 9 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 0.01, PME-PME 0.04, PME-PLE 
0.06, ALE-PLE 0.07. MOQ 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 médian notch reaching almost mid-length of 
plate. Vulva (Figs 103-104) characterized by uncoiled bag-like anterior part of médian 
ducts reaching anterior epigynal margin. 



NEW ZELOTES SPECIES 



539 





Figs 97-104 

Zelotes laconicus sp. n. (97-101) Left maie palp. (97) Prolateral view. (98) Ventral view. (99) 
Retrolateral view. (100) Apex of palpai 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 Unes 0.2 mm. 



540 



A. SENGLET 



Z. fulvopilosus-subgroup 

SPECIES INCLUDED: Z.flagellans, Z.fulvopilosus. 

Zelotes flagellons (L. Koch, 1 882) Figs 105- 1 1 1 

Prosthesima flagellans L. Koch. 1882: 635, pl. 20, fig. 14 (description of 6). 
Zelotes fiilxopilosus . - Machado, 1949: 15, fig. 10 (misidentification). 

Materialexamined: SPAIN,Andalusia,Granada,La Calahora (in pine forest). 37°10'N 
03°03'W; 2 d, 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. 
- 6 : 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 médian apo- 
physis. Intercalary sclerite présent. 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. Médian 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 r2 1 1 ; metatarsi III, IV v221. Tarsi 
and metatarsi I, II entirely scopulate. Scutum occupying almost 50% of opisthosoma 
length. - 9: 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 latéral and 
posterior folds. Deep latéral 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 révolutions. Médian ducts reaching anterior 
part of epigynal plate. Leg spination: Fémur IV pOOl, tibiae III, 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 spécimens examinated. 

Zelotes fulvopilosus (Simon, 1878) Figs 112-118 

Prosthesima fiilxopilosa Simon, 1878: 61, pl. 14, fig. 14 (description of cT and 9). 
Zelotes fulvopilosus. - Simon. 1914: 178, 214, figs 317-318. 375 (description of 6 and 9). - 
Jézéquel. 1962: 598. fig. 3. 

Type material: Muséum National Histoire Naturelle Paris, collection Simon; 1 <5, 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 
(understones.C/5rw5),42°18'N03°13'E; 1 9 (last moult 30.06.2004); 23.05.2004. - Gerona. Val 



NEW ZELOTES SPECIES 



541 




Figs 105-115 

(105-111) Zelotes flagellons. (105-108) Left maie palp. (105) Prolateral view. (106) Ventral 
view. (107) Retrolateral view. (108) Apex of palpai organ, cymbium discarded, prolateral view. 
(109) Epigynum. (110) Vulva, ventral view. (111) Id., dorsal view. (112-115) Zelotes fulvo - 
pilo s us, left maie palp (spécimen 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 6, \ 9; 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 9 (last moult 24.05.2002); 19.05.2002. - Granada, La Vidriera / Pinar pass, 38°03'N 
02°34'W; 2 6 , 3 9 (last moult of 1 9 31 .05.2002); 20.05.2002. - Granada, Prado de Zangarrilla 
(Sierra Nevada, under stones, pasture, 2000m) 37°07'N 03°26'W; 1 S\ 29.05.2002. -Jaen, Sierra 
de Cazorla (Fuente del Oso), 37°55'N 02°56'W; 4 S , 4 9 (with vulva and palpus in microvial, 
last moult of 1 9 27.05.2002); 21.05.2002. - Jaen, Sierra de Cazorla (Linarejas), 37°55'N 
02°55'W; 1 9; 22.05.2002. -Cordoba, Almodovar del Rio (Brena dam), 37°50'N 05°04'W; 1 9; 
01 .06.2009. - Sevilla, Alanis, 38°02'N 05°11'W; 1 S; 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 9 ; 07.07.1969. - Same; 2 9 ; 07.06.2002. - Same (cork-oak leaf litter); 3 9 ; 09.06.2009. - 
Huelva, Fuenteheridos (deep leaf litter), 37°54'N 06°40'W; 1 9; 12.06.2009. - FRANCE, 
Languedoc, Eastern Pyrénées, above Banyuls sur Mer (350m), 42°28'N 03°08'E; 1 9 (last moult 
07.06.2004); 23.05.2004. 

Description: Prosoma dark brown. Legs brown, with lightened tarsi. Opistho- 
soma black, covered with black bristles. - S from Spain, Fuente del Oso (in paren- 
thèses 6 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. MOQ 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 médian 
apophysis. Intercalary sclerite présent. 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, II in apical half. Leg spination: Tibiae III, IV p21 1 ; metatarsi III, 
VI v221. Scutum occupying 35% of opisthosoma length. - 9 from Spain, Fuente del 
Oso (in parenthèses 9 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. 
MOQ 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. Latéral epigynal pouches shallow, extended along the folds. Vulva 
(Figs 117-118): Médian ducts situated in posterior half of epigynal plate. Tarsi and 
metatarsi I, II entirely scopulate. Leg spination: Tibiae III p2 11, rl 1 1 , IV p2 11, r2 1 1 ; 
metatarsi III, VI v221. 

Zelotes baeticus-group 

Définition: Light coloured spiders. Maie palp: Retrolateral tibial apophysis 
with flattened and widened tip. A short intercalary sclerite présent. 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 basai 



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 médian apophysis. Epigynum: Latéral folds modi- 
fied into large latéral pockets; médian part membranous (Figs 123, 134). Cheliceral 
groove: Retromargin with large prolateral conical tooth and two smaller teeth behind; 
promarginal teeth (from entai to ectal): Médium, large, médium 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 

1 Prolateral apical protrusion of médian apophysis of bulbus large and 
conspicuous, bent retrolaterally (Figs 131, 133); embolus wide open; 

latéral epigynal pockets covering latéral folds Z. baeticus sp. n. 

Prolateral apical protrusion of médian apophysis reduced and cone- 
shaped (Figs 120, 127); embolus flattened 2 

2 Embolar base with terminal projection; embolus with ventral inden- 

tation; female unknown Z. cordubensis sp. n. 

Embolar base without terminal projection; embolus without ventral 

indentation; latéral epigynal pockets separated from latéral folds 

Z. hispaliensis sp. n. 

Zelotes hispaliensis sp . n . Figs 119-125 

Holotype: SPAIN, Andalusia, Huelva, Alajar /Aracena, 37°52'N 06°41'W; 6; 
07.06.2002. 

Paratypes: Same locality as for holotype; 3 6,59 (last moults of 1 S 09.06.2002, of 
2 9 20.06 and 07.07.2002). - Same; 2 S (with palpus in microvial); 07.07.1969. - Same (cork- 
oak leaf litter); 2 9 (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 6 ; 10.06.2009. 

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. - S paratypes (smaller S in paren- 
thèses): 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 0.01, 
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 médian apophysis forming a gutter. Apical protrusion 
on médian apophysis small, conical (Fig. 120). Embolar base projection absent. 



544 



A. SENGLET 




FlGS 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 maie palp. (119) Prolateral view. (120) 
Ventral view. (121) Retrolateral view. (122) Apex of palpai organ, cymbium discarded, retro- 
lateral-apical view. (123) Epigynum. (124) Vulva, ventral view. (125) Id., dorsal view. Bold Unes 
indicate epigynal folds. Scale Unes 0.2 mm. 



NEW ZELOTES SPECIES 



545 



Embolus with a prolateral protrusion (Fig. 122).Leg spination: Metatarsi I, II v220; III, 
IV v221, some large spécimens with an additional spine on fémur II pOl 1 . Tarsi and 
metatarsi I, II entirely scopulate. Scutum occupying 1/4 of opisthosoma length. - 9 
(smaller spécimen in parenthèses): 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 0.07, ALE 0.11, PME 0.07, PLE 0.08; AME-AME 0.07, AME-ALE 0.01 , 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): Latéral 
anchoring pockets not reaching posterior level of latéral folds; darker posterior médi- 
an marking due to médian vulval ducts. Vulva (Figs 124-125).Leg spination: Fémur II 
pOl 1 , metatarsi III v221 , rl22; IV rl22, some large spécimens with an additional spine 
on tibia III ri 1 1 . 

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; 6\ 31.05.2009. 

Paratype: Spain, Cordoba, Brena dam (evergreen oak litter), 37°51'N 05°04'W; 1 6; 
01.06.2002. 

Etymology: The species name, an adjective, is 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. - â holotype (paratype in parenthèses): 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 médian 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 indexa- 
tion. Tarsi I, II entirely scopulate; metatarsi in apical half. Leg spination: Fémur II 
pOll; 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 médian apophysis, and to Z. baeticus sp. n. due to the 
présence of a ventral indentation on the embolus and due to a terminal projection on 



546 



A. SENGLET 



the embolar base. The différences from the two sister species justify the description of 
a new species. The spécimens 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; 6 (last moult 12.06.2002); 09.06.2002. 

Paratypes: Same locality as for holotype; 1 S , 1 9 (last moults of â 27.06.2002, of 9 
09.06.2002). -2 6\ same; 14.06.2009. - Spain, Andalusia, Cordoba, road Posada-Villaviciosa 
(under stones in Pinus litter), 37°00'N 05°05'W; 1 â (last moult 19.06.2009); 02.06.2009. - 
Spain, Huelva, north of La Nava (schist litter), 38°00'N 06°45'W; 3 6 , 2 9 (last moults of 2 6 
16.06.2002, of 1 9 1 1 .08.2002); 09.06.2002. - Spain, Huelva, Alajar /Aracena (under stones and 
in végétation litter), 37°53'N 06°40'W; 1 S, 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 9 (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 is the adjective of Baetis, the Roman name of 
the river Guadalquivir. 

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 straight or slightly recurved. - $ paratype from Rio 
Sillo,: 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 0.01 , 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 médian apophysis triangular, forming a gutter. Médian apophysis with a large blunt 
apical protrusion (Figs 131, 133) corresponding to a rounded ventral indentation in 
basai half of terminal sector of embolus (Fig. 132). Embolar base with an apical pro- 
jection with a triangular dorsal flap. Tarsi I, II entirely scopulate, metatarsi I, II in 
apical half. Leg spination: Metatarsi I, II v220; III, IV v221 ; some large spécimens with 
an additional spine on fémur II pOll. Scutum occupying 1/5 of opisthosoma length. - 
9 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): Latéral epigynal pockets partially 
covering latéral folds, extended over posterior level of folds (Fig. 134). Vulva (Figs 
135-136). Tarsi and metatarsi I, II entirely scopulate. Leg spination: Tibia III rlll; 
metatarsi III, IV v221 ; some large spécimens with an additional spine on fémur II pOl 1 . 

Remarks: Z. baeticus seems to occur in a wide range of biotops: Below stones 
in Pinus, grass and schist litter. 



NEW ZELOTES SPECIES 



547 




Figs 126-136 

(126-128) Zelotes cordubensis sp. n., left maie palp. (126) Holotype, prolateral view. (127 Id., 
ventral view. (128) Paratype, cymbium discarded, retrolateral view. (129-136) Zelotes baeticus 
sp. n. (129-133) Left maie 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. latreillei (Simon). 



Zelotes atrocaeruleus (Simon, 1878) Figs 137-145 

Prosthesima atrocaerulea Simon, 1878: 73, pl. 14, fig. 16 (description of 6 and 9). 
Zelotes atrocoruleus . - Miller, 1967: 260, pl. 1 , figs 15-18, pl. 5, fig. 7 (description of 6 and 9). 
- Grimm, 1985: 238, figs 279, 292-293 (description of 6 and 9 ). 

Material examined: SWITZERLAND, Vaud, near Genolier, 46°26'N 06°14'E; 
I9(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 cT (with palpus in 
microvial), 1 9 (last moults ô* 1 and 11.07.2005, 9 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 6 (last moult 08.06.2005); 18.09.2004. - Phthiotidas, West of Malesina, 
38°37'N 23°13'E; 1 ô\ 1 9; 21.05.1968. - Same; 1 et, 2 $ (last moults 9 01.06.1998); 
21.05.1998. - Same; 1 6 (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 cT; 29.05.1983. - Peloponnesus, Elis, Andritsaina, 37°30'N 21°53'E; 2 ô\ 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 6 (with palpus in microvial, last moult 02.07.2005); 01.10.2004. 

REMARKS: The Greek â 6 (Figs 137-143) correspond to illustrations by Grimm 
(1985: fig. 279) and Miller (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 9 9 differ (Figs 144-145) by longer 
epigynum and glandular duct, and by a shorter médian duct. 

Description: See Grimm (1985: 238, figs 279, 292-293). 



Zelotes latreillei (Simon, 1878) Figs 146-147 

Prosthesima latreillei Simon, 1878: 62 (description of â and 9). 

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 6 and 9). - Grimm, 1985: 201, figs 2, 7, 9, 220- 
221,241,258-259 (description of 6 and 9). - Senglet, 2004: 117, figs 102-103 (9). 

Material examined: GREECE, Macedonia, Thessaloniki, west of Aghios Vassilios, 
40°40'N 23°05'E; 2 6; 08.09.2004. - Same; 1 S (last moult 03.08.2008); 31.05.2008. - 
FRANCE, Languedoc, Aude, Carcanet forest (in pasture), 42°41'N 02°08'E; 16,69 (with 
vulva in microvial, last moults 6 24.08.2001, 2£ 30.08.2001); 04.07.2001. - Aude, Carcanet 
forest (in pasture edge), 42°40'N 02°09'E; l 6,1 9 (last moults 6 15.08.2001, 9 15.09.2001); 
04.07.2001. - Eastern Pyrénées, above Les Fourquets (1800m), 42°26'N 02°25'E; 1 9; 
30.06.2001. - Eastern Pyrénées, Les Fourquets, 42°26'N 02°23'E; 1 6 (last moult 0.08.2001); 
30.06.2001 (1600m). - Eastern Pyrénées, Les Angles, /Matemale forest (1700m), 42°33'N 
02°04'E; 2 9; 03.07.2001. - PORTUGAL, Tras-Os-Montes, Vila Real, Cortico /Montalegre 
(900m),41°46 , N07°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 




Figs 137-147 

(137-145) Zelotes atrocaeruleus . (137-141) Left maie 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 maie palp, cymbium discarded. (146) Prolateral view. (147) Dorsal view. Bold 
lines indicate epigynal folds. Scale Unes 0.2 mm. 



550 



A. SENGLET 



Zelotes subterraneus-group 

Définition: Embolar base a simple trans verse 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 (CL. 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 / Léon, Avila, Penanegra pass (under stones), 40°25'N 
05°18'W; S (last moult 01.07.2009); 17.06.2009. 

Paratypes: Same locality as for holotype; 1 1 9 (with palpus and vulva in microvial, 
prosoma 6 partly destroyed, last moults S 20.07.2009, 9 04.08.2009). - Same locality as for 
holotype; 3 et, 1 9 (last moults 6 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 â (last moult 08.07.2009); 
18.06.2009. - Spain, Avila, Tornavaca pass (under stones), 40°16'N 05°40'W; 2 c?, 1 9 (last 
moults S 02.7 and 07.08.2009, 9 24.07.2009); 17.06.2009.- Spain, Badajoz, Rio Sillo (Higuera 
la Real), 38°06'N 06°41'W; 1 â, 1 9 (with palpus and vulva in microvial, last moults 6 
27.09.2009, 9 20.10.2009); 14.06.2009. 

Other material examined: SPAIN, Aragon /Navarra, Saragosse, Ariza, 48°18'N 
02°05'W; 3 9 (with vulva in microvial); 24.09.1967. -Salamanca, El Cabaco, 40°32'N 06°08'W; 
1 S; 12.08.1971. - Estremadura, Caceres, west of Jarandilla (litter in végétation), 40°08'N 
05°40'W; 1 9 (last moult 01.10.2009); 16.06.2009. - Andalusia, Granada, La Vidriera /Pinar 
pass, 38°03'N 02°34'W; 1 Ô (last moult 02.08.2002); 20.05.2002. - Granada, Puerto de la Ragua 
(2000m), 37°07'N 03°02'W; 1 S (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 â 1 â juv.; 
29.08.1969. - Vila Real, Cortico /Montalegre, 41°46'N 07°47'W; 1 S; 30.08.1969 (900m). - 
Beira Alta, Guarda, Maceira /Fornos de Algodres (Casai do Monte), 40°44'N 07°24'W, 1 6 , 1 9 
(with palpus and vulva in microvial); 09.08.1971. 

Etymology: The species name indicates a close relationship with Z. egregius 

Simon. 

DiAGNOSiS: cT: Close to Z. aeneus and Z. egregius (see Senglet, 2004: figs 
55-57) with respect to terminal apophysis and embolar base; differs from thèse two 
species by the absence of an acute prolateral ventral tooth on the embolar base and by 
the absence of a médian membranous flap on the embolus. - 9 : Differs from that of 
Z. egregius (see Senglet, 2004: figs 59-61) by the strongly retrolaterad-directed and 
longer glandular duct. Différent 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. - 6 paratype 
from Penanegra, (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 




Figs 148-156 

Zelotes egregioides sp. n. (148-152) Left maie palp. (148) Retrolateral view. (149) Ventral view. 
(150) Retrolateral view. (151) Embolus complex cleared, dorsal view. (152) Tip of embolus. 
(153) Vulva, short médian duct type, dorsal-lateral view. (154) Id., dorsal view. (155) Vulva, 
long médian duct type, dorsal view. (156) Epigynum. Bold Unes indicate epigynal folds. Scale 
lines 0.2 mm. 



552 



A. SENGLET 



interdistances: AME 0.06, ALE 0.11, PME 0.10, PLE 0.10; AME-AME 0.06, AME- 
ALE 0.01 , 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; III, IV v221. 
Tarsi I, II entirely scopulate, metatarsi in apical 3/4. - 9 paratypes from Penanegra: 
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 médian ducts variable (in Figs 153-154 short, in 155-156 long). Leg spination: 
Metatarsi III, IV v221. Tarsi and metatarsi I, II entirely scopulate. 

Remarks: Adult 6 â occur from July to September, 9 from July to October. 
The â examined by Machado (1941: fig. 10) could belong to Zeloîes egregius because 
of the présence of an acute prolateral tooth on the embolar base. 

Zelotes criniger-metellus-group 

Définition: 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 présence of a posterior sclerite on the terminal apophysis, the shape of the ocular 
group and the gênerai appearance, I leave this group in the genus Zelotes. 

KEY TO THE SPECIES OF THE Z. CRINIGER-METELLUS-GROUP 

1 Embolus elongated and coiled (Fig. 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 S). - Di Franco, 1987: 149, 
figs 8, 11 (â). 

Zelotes denisi Marinaro, 1967: 693, fig. 9 (description of 9), syn. n. - Di Franco, 1987: 152, 
figs 12-13(9).- Deltshev et al. 2006: 712, figs 12-13(9). 

Material examined: SPAIN, Andalusia. Cordoba. Brena dam, Almodovar del Rio (evergreen 
oak litter), 37°51'N 05°04'W; 4 6, 6 9 (with palpus, and vulva in microvial); 
01 .06.2002. - Cordoba. Almodovar del Rio (Brena dam), 37°50'N 05°04'W; 2 6 , 5 9 ; 
01.06.2009. - Sevilla, east of Cazalla de la Sierra (deep leaf litter), 37°57'N 05°45'W; 
1 9:04.06.2009. 

DESCRIPTION: Prosoma brown, adorned with fine adpressed hairs. Opisthosoma 
black, covered with médium adpressed hairs. Legs brown, with darker femora. 
Posterior médian eyes larger than posterior laterals in slightly recurved line. - 6 from 
Brena 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 



553 




Figs 157-164 

Zelotes criniger. (157-161) Left maie palp. (157) Prolateral view. (158) Ventral vievv. (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, II v220; III rl22. Tarsi I, II entirely sco- 
pulate. Scutum occupying 40% of opisthosoma length. - 9 from Brena 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 0.01 , PME-PME 0.02, PME-PLE 0.03, ALE-PLE 0.03. MOQ 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 latéral folds extended close to anterior 
anchoring pockets. Médian epigynal surface membranous. Additional latéral pockets 
présent. Vulva in dorsal view with large spiralled pouches almost reaching anterior 
anchoring pockets (Figs 163-164); sclerotized éléments présent in the posterior dorsal 
part. Leg spination: Metatarsi I v210; II v210; III rl22. 

Zelotes metellus-subgroup 

DEFINITION: Maie palp différent 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 présent. Embolar radix 
linked to embolus by membranous cuticle. Embolus short and wide. Retrolateral 
posterior lamina présent on tegulum. Epigynum: Médian 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 

1 6 : 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 elongated, not fused to epigynum 

(Fig. 170) Z. prishutovae 

â : Posterior apophyses of embolar base and embolar radix shorter, not 
visible on resting pedipalp; tip of embolus tapering (Figs 172, 175). 9 : 
Anterior sacs of vulva short, fused to cuticle of epigynum (Fig. 177) . . 
Z. metellus 

Zelotes prishutovae Ponomarev & Tsvetkov, 2006 Figs 165-170 

Zelotes prishutovae Ponomarev & Tsvetkov, 2006: 13, figs 25-26 (description S and 9). 
Camillina metellus. - Chatzaki, Thaler & Mylonas, 2003: 48, figs 3-7 (not figs 8-9 = Z. metellus). 



NEW ZELOTES SPECIES 



555 




Figs 165-170 

Zelotes prishutovae. (165-169) Left maie 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 Unes indicate epigynal folds. Scale lines 
0.2 mm. 



Material examined: GREECE. Peloponnesus, Laconia. south-west of Monemvasia, 
36°40'N 23°01'E; 2 â; 11 .05.1998. - Crète Lassithi 2 km west of Istro: 1 9 ; 02.06.1997 (les. 
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. - S : 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, II entirely scopulate, metatarsi I, II in apical third. Leg spination: 
Metatarsi I vOOO; III, IV v221. Scutum occupying 1/5 of opisthosoma length. - 9: 
Vulva (Fig. 170): Inflated and elongated glandular ducts with scattered pores. 

Zelotes metellus Roewer, 1 928 Figs 171-178 

Zelotes metellus Roewer 1928: 1 10, pl. 1 , figs 16-17 (description of 9 ). 

Camillina metellus. - Chatzaki, Thaler & Mylonas, 2003: 48, figs 8-9 (not 6 in figs 3-7 = 
Z. prishutovae) . 

Zelotes sumchi Levy, 1998: 151, figs 128-130 (description of S) syn. n. - Levy, 2009: 41, figs 
87-88 (description of 9). 

Material examined: IRAN, Fars, Firouzâbâd, 28°52'N 52°32'E; 1 â,l 9,1 immature 
<?; 06.06.1974. - Kavâr, 29°12'N 52°37'E; 1 S; 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. - â from Firouzâbâd (in parenthèses 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 0.01, 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, II entirely scopulate, metatarsi I, II in apical two thirds. 
Leg spination: Tibia III rl 1 1 ; metatarsus I vOOO. Scutum occupying 1/5 of opisthosoma 
length. - 9: 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): Latéral anterior vulval sacs fused to epigynal 
cuticle, visible on epigynum. Posterior eye row slightly recurved. Tarsi and metatarsi 

1, II scopulate. 

REMARKS: Chatzaki et al. (2003: 48) transferred this species to Camillina on the 
basis of a bifid terminal apophysis on the maie 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 is not bifid. 



NEW ZELOTES SPECIES 



557 




Figs 171-178 

Zelotes metellus. (171-176) Left maie 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 Muséum of Natural 
History in Paris, and to Mrs Janet Beccaloni of the Natural History Muséum in London 
for making type material available; to Jôrg 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 Muséum of 
Natural History) for producing the Scanning Electron Microscope pictures; to the 
Geneva Muséum of Natural History for covering the charges of thèse pictures. 

REFERENCES 

CHATZAKI, M. Thaler. K. & Mylonas, M. 2003. Ground spiders (Gnaphosidae: Araneae) from 

Crète and adjacent areas of Greece. Taxonomy and distribution. III. Zelotes and allied 

gênera. Revue suisse de Zoologie 110(1) 45-89. 
Chyzer. C. & Kulczynski, W. 1897. Araneae hungariae. 2. Ediîio Academiae Scientiarum 

Hungaricae, Budapest.^. 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). 

Miîîeilungen aus dem Muséum fur Naîurkunde 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 française 

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 i iougo-zapadna Makedonia. Spisanie na 

Bàlgarska akademija na naukitë, Sofia. 39: 1-76. 
Grimm. U. 1985. Die Gnaphosidae Mitteleuropas (Arachnida. Araneae). Abhandlungen des 

Natunvissenschaftlichen Vereins Hamburg 26: 1-318. 
JÉZÉQLEL. 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, Niirnberg, 

pp. 1-304, pis 1-12. 

Koch. L. 1882. Zoologische Ergebnisse von Exkursionen auf den Balearen. II: Arachniden und 
Myriapoden. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 31: 
625-678. 

Levy. G. 1998. The ground-spider gênera Setaphis, Trachyzelotes. Zelotes. and Drassyllus 
(Araneae: Gnaphosidae) in Israël. Israël Journal ofZoology 44: 93-158. 

Levy. G. 2009. New ground-spider gênera and species with annexed checklist of the Gnapho- 
sidae (Araneae) of Israël. Zootaxa 2066: 1-49. 



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Machado, A. de B. 1941. Araignées nouvelles pour la faune portugaise (II). Memorias e 
Estudos do Museu Zoologico da Universidade de Coimbra 1 17: i-xvi, 1-60. 

Machado, A. de B. 1949. Araignées nouvelles pour la faune portugaise (III). 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 iiber 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 Muséum of Natural History 174: 
97-192. 

Platnick, N. I. 2011. The World Spider Catalog, Version 11.5. American Muséum of Natural 
History, online at: http://research. amnh. org/entomology/spiders/catalog. html. 

Ponomarev, A. 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 Streifzùge in Attika, Morea, und besonders auf der Insel Kreta. 
XI. Araneae. Abhandlungen herausgegeben vom Naturwissenschaftlichen Verein zu 
Bremen 27: 92-123, pis 1-2. 

Senglet, A. 2004. Copulatory mechanisms in Zelotes, Drassyllus and Trachyzelotes (Araneae, 
Gnaphosidae), with additional faunistic and taxonomic data on species from southvvest 
Europe. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 11: 87-119. 

Simon, E. 1878. Les arachnides de France, 4. Roret, Paris, 334 pp. 

Simon, E. 1914. Les arachnides de France, 6, Ire partie: Synopsis générale et catalogue des 
espèces françaises de l'ordre des Araneae. Roret, Paris, 308 pp. 



Revue suisse de Zoologie 118 (3): 561-598; septembre 201 1 



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 ton g lu 
sp. n., Pelecopsis indus sp. n., Tapinocyboides bengalensis sp. n., Tchat- 
kalophanîes baltistan sp. n., Tiso incisus sp. n., T. (?) indianus sp. n., 
Walckenaeria saetigera sp. n. A new genus, Paracymboides gen. n., is 
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, i.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 régions: the northern, mountain terri tories, be- 
long to the Palaearctic région, but the main territories lie in the Oriental région. Ail 29 
linyphiid species known from Pakistan were described or recorded from the Palaearctic 
part, the Karakorum mountain System (Caporiacco, 1934, 1935). The linyphiid 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 cornes 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 24.06.201 1 



562 



A. V. TANASEVITCH 



MATERI AL AND METHODS 

This paper deals with linyphiids collected in Pakistan and India and kept at the 
Muséum d'histoire naturelle, Geneva, Svvitzerland (MHNG). A small additional 
material vvas also available from the Museo Civico di Storia Naturale di Verona, Italy. 
Unless otherwise stated, ail material is deposited in the MHNG. Some paratypes and 
non-type spécimens are in the collection of the Zoological Muséum 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) vvhich refers 
to the number of dorsal spines on tibiae I-IV. In Micronetinae, the chaetotaxy is given 
in a différent 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 séquence of leg segment measurements is as follows: fémur + 
patella + tibia + metatarsus + tarsus. Ail measurements are given in mm. Ail scale Unes 
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 



c 


Convector (according to Tanasevitch, 1998: 423) 


CAT 


Personal collection of Andrei Tanasevitch 


D 


Duct 


DE 


Duct entrance 


DSA 


Distal suprategular apophysis 


E 


Embolus 


ED 


Embolie division 


EP 


Embolus proper 


L 


Lamella characteristica 


MED 


Membrane of embolie division 


MHNG 


Muséum d'histoire naturelle, Geneva, Svvitzerland 


MM 


Médian membrane 


MSNV 


Museo Civico di Storia Naturale di Verona, Italy 


MT 


Membranous tissue 


PS 


Proscape 


PMP 


Posterior médian plate 


R 


Radix 


S 


Suprategulum 


SMF 


Senckenberg Muséum, Frankfurt a. M., Germany 


TA 


Terminal apophysis 


Th 


Thumb of embolus 


ZMMU Zoological Muséum of the Moscow State University, Moscow, Russia 



LINYPHI1D SPECIES FROM PAKISTAN AND INDIA 



563 



RESULTS 

Acartauchenius himalayensis sp. n. Figs 1-8 

Holotype: S (MSNV), PAKISTAN. Northern Areas, Gilgit District, Bagrot Valley, 
2500 m a.s.l.; 20.VI.2008; leg. L. Latella. 

ETYMOLOGY: The new species is named after the mountain System where it 
occurs; Latin adjective. 

DlAGNOSIS: A. himalayensis sp. n. is characterized by the peculiar shape of the 
maie carapace and palpai tibia. 

DESCRIPTION: Maie. Total length 1.58. Carapace 0.70 long, 0.55 wide, pale or- 
ange-brown, modified as shown in Figs 1-2. Head élévation carrying several strong 
curved spines directed backvvard. 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. Tml 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 A. asiaticus (Tanasevitch, 
1989) and A. monoceros (Tanasevitch, 1989) known from Turkménistan and 
Uzbekistan, respectively (Tanasevitch, 1989), but it differs clearly by the the peculiar 
shape of the maie carapace and palpai tibia. 

Distribution: Known from the type locality only. 

Agyneta nigripes (Simon, 1884) Figs 9-15 

Material:2 6,4 9 (MSNV), PAKISTAN, Northern Areas, Gilgit District, Passu,Passu 
Glacier (in the glacier), 2635 m a.s.l.; 30.X.2008; leg. L. Latella. 

Comparative material examined: Agyneta nigripes, more than 100 6 & 9 , AUS- 
TRIA, Mt Glockner, 1900-2580 m a.s.l.; 1978-1980; leg. K. Thaler. -2 6,3 9 (CAT), RUS- 
SIA, Arkhangelsk Area, 30 km W of Tobseda Village, 4 km S of Lake Peschanka-To (68°42'N, 
53°10'E); VII. 1984; leg. A. Tanasevitch. - 1 6 (CAT), Murmansk Area, Kola Peninsula, 
Dal'niye Zelentsy (69°07'N, 36°04'E); 4.-26.VII.2009; leg. A. Babenko. - 8 6 , 10 9 (ZMMU), 
Taymyrskiy Autonomous Région, Efremova River, meadovv; 7.VII.-1 VIII. 2004; leg. D. Osipov. 
-2 6,2 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 différent from those of conspecific spécimens 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 spécimen. 

Remarks: This is the southernmost known locality of A. nigripes. The species 
is here recorded from the Himalayas and Pakistan for the first time. 

Range: Holarctic arcto-alpine. 



564 



A. V. TANASEVITCH 




FlGS 1-8 

Acartauchenius himalayensis sp. n., S maie holotype. (1-2) Carapace, latéral and dorsal views, 
respectively (not to scale). (3-5) Right palp, retrolateral, prolateral and proventral views, 
respectively. (6) Palpai tibia, dorsal view. (7) Distal suprategular apophysis and embolus. (8) 
Embolus. 



Agyneta pakistanica sp. n. Figs 16-25 

Holotype: 6 , PAKISTAN, Punjab, environs of Islamabad, ca 550 m a.s.l., Lake Rawal, 
on bank near water; 26.IV.1984; leg. S. Vit [PAK-84/25]. 

Paratype: 1 S , from same locality, collected together with the holotype. 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



565 




Figs 9-15 

Agyneta nigripes (Simon, 1884), 6 spécimen from Passu Glacier, Pakistan. (9) Right palp, retro- 
lateral view. (10) Cymbium , prolateral view. (11) Palpai tibia, dorsal view. (12) Palpai tibia and 
paracymbium, retrolateral view. (13) Embolie division. (14-15) Lamella characteristica, différent 
aspects. 

ETYMOLOGY: The spécifie name is an adjective taken from the type locality. 

DiAGNOSlS: The new species is characterized by the peculiar shape of the palpai 
tibia and of the lamella characteristica. 

Description: Maie (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: ail tibiae with two dorsal spines. Tml 0.25. Metatarsus IV without 



566 



A. V. TANASEVITCH 




Figs 16-25 

Agyneta pakistanica sp. n., 6 paratype. (16) Right palp , retrolateral view. (17) Cymbium, 
prolateral view. (18) Palpai tibia, dorsal view. (19-21) Palpai tibia and paracymbium, latéral 
view, différent aspects. (22) Embolie division. (23) Embolus. (24-25) Lamella characteristica, 
différent 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. 



L1NYPHIID SPECIES FROM PAKISTAN AND INDIA 



567 



Female unknown. 

Distribution: Known from the type locality only. 

Anguliphantes nepalensis (Tanasevitch, 1987) 

Material: 1 9 , PAKISTAN, Hazara District, Kâghân Valley, 1450 m a.s.l., Malkandi 
Forest, near foot of rock; 29 .VI. 1985; leg. S. Vit [PAK-85/2]. -16, INDIA, Uttar Pradesh, 
Garhwal, above Pauri, 1900 m a.s.l., dry Quercus forest on N slope, sifting leaf litter and moss; 
28.X. 1979; leg. I. Lôbl [281. -19, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m 
a.s.l., near top, sifting litter in forest; 18.X.1978; leg. C. Besuchet & I. Lôbl [19]. 

Comparative material examined: 1 â, NEPAL, Goropani Forest between Kali 
Gandaki Valley and Pokhara Valley, Punhill near Goropani, 3050-3100 m a.s.l., 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 génital 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 is consistent enough and is a good, 
well- visible character to distinguish this species. 

REMARKS: A. nepalensis was previously known from numerous localities in 
Népal (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: et , INDIA, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m a.s.l., 
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 A. nepalensis (Tanasevitch, 1987). 

DiAGNOSlS: The new species is characterized by the peculiar shape of the 
lamella characteristica. 

Description: Maie. 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 its distal segments mostly lost, Fel & II 1 .23 long. Chaetotaxy unclear, but probably 
equal to that of A. nepalensis, i.e. Fe I: 0-1-0-0: Ti I: 2-1-1-0, II: 2-0-1-0, III-IV: 
2-0-0-0; Mt I-III: 1-0-0-0. Tmll 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 is very similar to A. nepalensis, but 
can be easily distinguished by the shape of the distal part of the lamella characteristica. 
See also above, under Remarks to A. nepalensis. 

Distribution: Known from the type locality only. 



568 



A. V. TANASEVITCH 




Figs 26-31 

Anguliphantes nepalensoides sp. n., â holotype. (26) Right palp, retrolateral view. (27) Palpai 
tibia and paracymbium, dorsal view. (28) Embolie division. (29) Embolus. (30-31) Lamella 
characteristica, différent aspects. 



Archaraeoncus prospiciens (Thorell, 1875) 

Material: 1 9 (MSNV), PAKISTAN, Northern Areas, Gilgit District, Bagrot Valley 
(36 o 01'36.7"N, 74°33'57.6"E), 2700 m a.s.l.; 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.l.; 1 .XI.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 9 , PAKISTAN, Hazara District, Kâghân Valley, 2000 m a.s.l., 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.IV.1984; leg. S. 
Vit [PAK-84/24]. - 1 9 , N of Lake Rawal, 3.IV.1986, leg. S. Vit. - 1 9 (ZMMU), Ghnwool 
Valley, Malkandi Forest, 1600 m a.s.l., near water, among stones; 30.VI.1985; leg. S. Vit 
[PAK85/7] . - 1 S , Swat District, Malam Jabba, 2500-2600 m a.s.l., Abies forest, sifting leaf litter 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



569 



and moss; 18.V.1983; leg. C. Besuchet & [. Lôbl [17b]. - 1 9 , S of Saidu Sharif, Murghazar, 
sifting leaf litter under Platanus, 1300 m a.s.l.; 8.V.1983; leg. C. Besuchet & I. Lôbl [2b]. - 1 9 , 
above Miandam, 2400-2500 m a.s.l., Abies forest, sifting leaf litter and moss; 17.V.1983; leg. C. 
Besuchet & I. Lôbl [ 15b]. -1 6, INDIA, Himachal Pradesh, Kulu Valley, Chijoga (S of Manali), 
1900 m a.s.l., from rodent burrow; 14.X.1988; leg. S. Vit [22]. -39, Uttar Pradesh, Garhwal, 
4 km S of Bhatwari, 1400 m a.s.l.; 23.X.1979; leg. I. Lôbl [1979/22]. - 4 6, 2 9, Madras 
(= Chennai), Palni Hills, 10 km W of Kodaikanal, 2300 m a.s.l., timberline, degraded forest, 
sifting moss under fern; 13.XI.1972; leg. C. Besuchet & I. Lôbl [1972/25a]. - 1 <?,3 9 , Palni 
Hills, 10 km NW of Kodaikanal, 2150 m a.s.l., edge of Rhododendron forest with fern, sifting 
litter near river; 15.XI.1972; leg. C. Besuchet & I. Lôbl [1972/271. - 2 6,1 9 (ZMMU), 
Kodaikanal, 2100 m a.s.l., forest below town, sifting litter; 11. XI. 1972; leg. C. Besuchet & I. 
Lôbl [1972/22]. -2 6,5 9 , 10 km W of Kodaikanal, 2350 m a.s.l., degraded forest, under 
stones; 13.XI.1972; leg. C. Besuchet & I. Lôbl [1972/25b]. - 1 6 , 1 9 , 23 km W of Kodaikanal, 
Lake Berijam, 2150 m a.s.l., Rhododendron forest, sifting litter; 14.XI.1972; leg. C. Besuchet & 
I. Lôbl [1972/26]. - 1 9 , Nilgiri, Ootacamund, 2150 m a.s.l., sifting litter under bushes in ravine; 
21 .XI. 1972; leg. C. Besuchet & I. Lôbl [1972/41a]. - 1 9 , Meghalaya, Khasi Hills, Mawphlang, 
1800 m a.s.l., 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.l, forest with fern, sifting litter; 
24.XI.1972; leg. C. Besuchet & I. Lôbl [48]. 

Comparative material examined: Caviphantes pseudosaxetorum, SMF 29677, 1 9 
paratype, NEPAL, Thakkhola, from Kali Gandaki Valley to main Himalayan Mt. Ridge, 
Chadziou Khola Valley near Ghasa, 2330 m a.s.l., 31.X.1969, leg. J. Martens. 

Distribution: The species was originally described from the Népal Himalayas 
(Wunderlich, 1979). Later, it was found in Japan (Ono et al., 1991), China (Gao et al., 
1992), and just recently in Lebanon (Tanasevitch, 2011). The species is here recorded 
from Pakistan and India for the first time. 

Range: South Palaearctic-Oriental. 

Ceratinella wideri (Thorell , 1871) 

Material: 1 6 , PAKISTAN, Hazara District, Kâghân Valley, Naran, Lake Saiful Muluk, 
3100 m a.s.l., litter under Saxifraga; 4.-5 .VII. 1985; leg. S. Vit [PAK-85/20]. - 1 6 , Swat District, 
above Utrot, 2500 m a.s.l., Abies & Cedrus forest, moss and leaf litter among fallen trunks; 
13.V.1983; leg. C. Besuchet & I. Lôbl [lie]. -19, Malam Jabba, 2500-2600 m a.s.l., Abies 
forest, sifting leaf litter and moss; 18.V.1983; leg. C. Besuchet & I. Lôbl [17b]. 

REMARKS: The discovery of both sexes of this species in the Pakistan 
Himalayas makes its détermination reliable. The species is here recorded from the 
Himalayas and Pakistan for the first time. 

Range: Palaearctic. 

Erigone dentipalpis (Wider, 1 834) 

Material: 1 6 (MSNV), PAKISTAN, Karakorum, Baltistan, Shalabot, 1700 m a.s.l.; 
15.VIII.1976; leg. G. Osella. - 2 9 (MSNV), Katzaran, 2200 m a.s.l.; 23.VII.1976; leg. G. 
Osella. - 2 9 (MSNV), Northern Areas, Gilgit District, Ghangche, Kaplu Ghwari; 27.VI.2008; 
leg. L. Latella. - 2 9 (MSNV), Skardu District, 2300 m a.s.l.; 27.VI.2008; leg. G. Osella. - 1 ô* 
(MSNV), Skardu, Pakova, 2300 m a.s.l.; 1. VII. 1976; leg. G. Osella. - 1 6 (MSNV), Naltar 
Valley (36°12'34.4"N, 74°08'20.6"E), 3000 m a.s.l.; 1.XI.2008; leg. L. Latella. - 1 9, Swat 
District, Saidu Sharif, 1000 m a.s.l., river bank, under stones and cow-dungs; 1 1 .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 Bôsenberg & Strand, 1906 

Material: 2 S , PAKISTAN, Punjab, environs of Islamabad, Lake Rawal, ca 550 m 
a.s.l., on bank near water; 26.IV.1984; leg. S. Vit [PAK-84/25]. 

REMARKS: In the Himalayas the species was hitherto known only from Népal 
(Wunderlich, 1983) and it is here recorded from Pakistan for the first time. 

Gongylidiellum confusum Thaler, 1 987 

Type material examined: Gongylidiellum confusum, SMF 33728, 4 9 paratypes, 
INDIA, Kashmir, Pahalgam, coniferous forest, 2400 m a.s.l.; 14.V.1976; leg. J. Martens. - SMF 
33730, 2 6 paratypes, Sonamarg, Nichinai-Tal, 3100-3200 m a.s.l.; 9.VI.1976; leg. J. Martens. 

Other material: 4 9 (ZMMU), PAKISTAN, Hazara District, Ghnwool Valley, 
Malkandi Forest, 1600 m a.s.l., near water, among stones; 30.VI.1985; leg. S. Vit [PAK-85/7]. - 
1 â , 1 9 , Swat District, above Utrot, 2500 m a.s.l., Abies & Cedrus forest, sifting moss and leaf 
litter among fallen trunks; 13.V.1983; leg. C. Besuchet & I. Lôbl [lie]. - 1 9 , same, 2500-2600 
m a.s.l.. sifting rotten Abies vvood; 14.VI.1983; leg. C. Besuchet & I. Lôbl [12c]. -19, same, 
2600 m a.s.l. , wet meadow, sifting mosses & rotten wood; 13.V.1983; leg. C. Besuchet & I. Lôbl 
[1 ld]. - 1 9 . Chitral, Madaglasht. 2700 m a.s.l., sifting under Salix near river; 26.V.1983; leg. 
C. Besuchet & I. Lôbl [27b]. - 2 9 , Chitral, above Bumburet, valley to Pass Ustui, 2700 m a.s.l., 
sifting leaf litter under Viburnum; 25 .V.l 983; leg. C. Besuchet & I. Lôbl [25c]. -19, INDIA, 
Uttar Pradesh, Garhwal, 2 km E of Dhanolti. northern slope, 2250 m a.s.l., near source, sifting 
mosses and leaf litter under Rhododendron and Abies; 21. X. 1979; leg. I. Lôbl [19]. - 1 6 , 2 9 
(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]. - 1 6 (ZMMU), Himachal Pradesh, 10 km W of Simla, 
Jutogh. 2000 m a.s.l., 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 3 1 70 1 , 1 o\2 9 paratypes, 
NEPAL. Thaksang above Tukche, coniferous forest, 3150-3400 m a.s.l.; 5.-10.VII.1970; leg. J. 
Martens. 

Other material: 8 9 , 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 cî,6 9, Tonglu, 3 100 m a.s.l., near top, under stones; 1 6. X. 1978; leg. C. Besuchet & 
I. Lôbl [16a]. - 1 â , 2 9 (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 â, 2 9 (ZMMU), same, 
2700 m a.s.l., forest, sifting near litter; 16.X.1978; leg. C. Besuchet & I. Lôbl [17]. 

REMARKS: The species was originally described from Népal (Wunderlich, 
1983) and is here recorded from India for the first time. 

Range: Himalayan. 

Gongylidioides keralaensis sp. n. Figs 32-38, 1 14 

Holotype: S , INDIA, Kerala, Cardamom Hills, near Munnar, Muttapatti, 1700 m a.s.l, 
forest, sifting litter under tree ferns; 24 .XI. 1972; leg. C. Besuchet & I. Lôbl [48]. 

Paratypes: 2 9 , 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., 6 holotype. (32-33) Right palp, retrolateral and prolateral 
views, respectively. (34) Palpai tibia, dorsal view. (35) Distal suprategular apophysis. (36-37) 
Embolie division, différent aspects. 



DiAGNOSlS: The new species is characterized by the peculiar shape of the maie 
palpai tibia, as well as by the shape of the ventral epigynal plate. 

Description: Maie. Total length 2.35. Carapace 1.13 long, 0.85 wide, pale 
brown, unmodified, with blurred grey médian 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. Tml 0.74. Ail 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 



572 



A. V. TANASEVITCH 




Figs 38-41 

Gongylidioides keralaensis sp. n., 9 paratype (38) and G.pectinatus sp. n., S holotype (39-41). 
(38) Epigyne, ventral view. (39-41) Carapace, latéral, 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 médian 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). Tml 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 maie. 

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: d, INDIA, Himachal Pradesh, 12 km E of Mandi, 750 m a.s.l., leaf litter; 
25.X. 1988; leg. S. Vit [35]. 

Paratype: 1 6 , INDIA, Uttar Pradesh, Kumaon, environs of Bhim Tal, eastern slope, 
1500 m a.s.l., edge of dry secondary forest; 4.X.1979; leg. I. Lôbl [1]. 

ETYMOLOGY: The spécifie name is a Latin adjective meaning "with a comb", 
referring to the peculiar shape of the convector. 

DlAGNOSIS: The new species is characterized by the peculiar shape of the palpai 
tibia, as well as by the présence of a fringed lobe in the embolie division. 

Description: Maie (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. Tml 0.56 (0.38 in 
paratype). Ail 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 embolie 
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 frh.«ge. 
Abdomen 0.83 long, 0.55 wide, dark grey, with a pale médian 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 palpai tibia, as well as 
by the présence 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 S , INDIA, Meghalaya, 15 km N of Darugiri, Garo-Hills, 400 m a.s.l., 
forest, sifting litter in ravine; 4.XI.1978; leg. C. Besuchet & I. Lôbl [40b]. 

Description: Maie. 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 , ail 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 maie shows strong similarities to G. ungibbus Tanasevitch, 
1998, described from Népal (Tanasevitch, 1998), but differs by some small détails of 
palp structure. Each form is known from a single maie, so it is impossible to say now 
if the différences are due to variability of characters or if thèse maies belong to distinct 
but closely related species. 

Distribution: Known from a single locality in northern India. 



574 



A. V. TANASEVITCH 




Figs 42-46 

Gongylidioides pectinaîus sp. n., 6 paratype. (42-43) Right palp, retrolateral and prolateral 
views, respectively. (44) Palpai tibia, prolateral view. (45) Palpai tibia and paracymbium, retro- 
lateral view. (45) Embolie division. 

Halorates concavus sp. n. Figs 53-57 

Holotype: ô\ PAKISTAN, Swat District, above Utrot, 2600 m a.s.L, wet clearing in 
Abies & Cedrus forest. sifting mosses & rotten litter: 13.V.1983: leg. C. Besuchet & I. Lôbl 
[lld]. 

Paratype: 1 6 . Swat District, above Utrot, 2500-2600 m a.s.L, Abies & Cedrus forest, 
sifting rotten wood litter; 14.VI.1983; leg. C. Besuchet & I. Lôbl [12c]. 

Comparative material examined: Collinsia japonica, SMF 31674, 4 9, NEPAL, 
southern part of Annapurna Massive, Pass Gorapani. 2700-2800 m a.s.L; 23.11.1974; leg J. 
Martens. - SMF 31670, 2 6 , southern part of Dhaulagiri Massive, Dhorpatan, 3000 m a.s.L; 



L1NYPHI1D SPECIES FROM PAKISTAN AND INDIA 



575 




Figs 47-52 

Gorboîhorax aff. ungibbus Tanasevitch, 1998. (47-48) Right palp, retrolateral and prolateral 
views, respectively. (49-50) Palpai tibia, dorsal and prolateral vievvs, respectively. (51) Distal 
suprategular apophysis and embolus. (52) Distal suprategular apophysis and embolie division. 



24.V.1973; leg J. Martens. - SMF 31671, 1 9 , vestern part of Dhaulagiri Massive, trail from 
Dhorpatan to Tarakot, 3100-3600 m a.s.l.; 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.l.; IX. 1969; leg J. Martens. Ail identified by J. 
Wunderlich in 1979. 

ETYMOLOGY: The spécifie 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., â holotype. (53-54) Right palp, retrolateral and ventral views, 
respectively. (55) Palpai tibia, dorsal view. (56) Distal suprategular apophysis. (57) Embolie 
division. 

DïAGNOSlS: The new species is characterized by the peculiar shape of its 
embolus. 

Description: Maie (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. Tml 0.44. Metatarsus IV without trichobothrium. Palp 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



577 



(Figs 53-57): Tibia abruptly narrowed distally, ending with a tapering transparent 
membranous process. Posterodorsal cymbial outgrovvth keel-shaped, with a shallovv 
saddle. Distal suprategular apophysis flat, spatulate, with a narrow membrane distally. 
Radical part of embolie 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 embolie division. The new species 
clearly differs from H.japonica (Oi, 1964) (see Wunderlich, 1983), the only congener 
that also occurs in the Himalayas, by the shorter and wider lobes of the embolie 
division. 

Variability. Détails of the both palps in both maies examined look identical. 
Distribution: Known from the type locality only. 

Indophantes tonglu sp. n. Figs 58-63 

Holotype: S , 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 6 , West Bengal, Darjeeling District, Tonglu, northern slope, 2700 m a.s.l., 
forest, sifting litter; 16.X.1978; leg. C. Besuchet & I. Lôbl [17]. 

ETYMOLOGY: The spécifie 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: Maie (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, 
Mtl-IV: 1-0-0-0. Metatarsus IV without trichobothrium. Tml 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, its lower one wide, bifurcated. Embolus large, its 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 Népal (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 




Figs 58-63 

Indophantes tonglu sp. n.. 6 holotype (58-59. 61-63) & 6 paratype (60). (58) Right palp. retro- 
lateral view. (59-60) Paracymbium. différent aspects. (61) Embolie division. (62) Embolus. (63) 
Lamella characteristica. 



Indophantes digitulus (Thaler. 1987) 

Material: 2 9 . PAKISTAN. Punjab. Murree. 1950 m a.s.l.. leaf litter under Quercus; 
23.IV.1984: leg. S. Vit [PAK-84/20]. -19. Hazara District. Kâghân Valley. NW of Mahandri. 
Kamalban Forest. 1800 m a.s.l.. fern litter: 3.VII.1985: leg. S. Vit [PAK-85/15]. -2 6. Nathia 
Gali. 2300 m a.s.l.. under stones: 17.IV.1984: leg. S.Vit [PAK-84/4]. -19. INDIA, Himachal 
Pradesh. Kulu Valley. Chijoea (S of Manali). 1900 m a.s.l.. in rotten stump of coniferous tree; 
12.X. 1988: leg. S. Vit [16]. - 1 9 . Kulu Valley, Vashisht Baths (N of Manali). bank of Beas 
River. 1900 m a.s.l.. fern litter: 13.X.1988: leg. S. Vit [20]. - 16.39, Chijoga (S-Manali). 1900 
m a.s.l.. wet ravine, from rodent burrovv; 14.X.1988: leg. S. Vit [22]. - 3 6 , 2 9 . Khajjiar. E of 
Dalhousie. 1950 m a.s.l.. 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 Népal (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. S , 2 9 , PAKISTAN, Punjab, Murree, 1950 m a.s.l., leaf litter under 
Quercus; 23.IV.1984; leg. S. Vit [PAK-84/20]. -19,3 subad. 9, same, leaf litter under 
Aesculus; 23.IV.1984; leg. S. Vit [PAK-84/22]. -19, Hazara District, Kâghân Valley, NW of 
Mahandri, Kamalban Forest, 2200 m a.s.l., leaf litter under Viburnum; 3.VII.1985; leg. S. Vit 
[PAK-85/18]. - 19,3 subad. 9 , Dunga Gali, 2300 m a.s.l., leaf litter in scree; 22.IV.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 6 , INDIA, New Dehli, coll. Heimer. - 1 6 , Uttar Pradesh, Kumaon, Bhim 
Tal, eastern slope, 1500 m a.s.l., 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, lying just at the border between the Palaearctic and the 
Oriental régions. 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 a.s.l., 
23.VII.1976, leg. G. Osella. - 4 9 (MSNV), 2 9 (ZMMU), Northern Areas, Skardu District, 
Skardu, Pakova, 2300 m a.s.l., 27 .VII. 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.l., 14.X.1978, leg. C. Besuchet & I. Lobl [14]. 

REMARKS: This species was hitherto known only from Népal (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 



DlAGNOSlS: The new genus is characterized by the highly modifiée! palpai 
tibia, the narrow, long and mammoth-tusk-like paracymbium, the simple embolie 
division with a long embolus, and the well-developed médian membrane. 

Description: Small-sized erigonines, total length 1.50-1.80 mm. Carapace 
unmodified, sulci absent, eyes normal. Chaetotaxy 1.1.1.1. Tml 0.30-0.36. Metatarsus 
IV without trichobothrium. Palpai tibia highly modified. Paracymbium narrow, very 
long and strong curved. Médian membrane well developed and protruded forward. 
Embolie division very simple: radical part elongated, flat, without outgrowths or 
processes; embolus thin, very long and slightly curved distally. Epigyne without cavity, 
its ventral surface (= "ventral plate" sensu Millidge (1984)) hairy. Réceptacles rela- 
tively large, complex. 

Taxonomic remarks: It is difficult to say anything about the closest relatives 
of this genus at présent. It is 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: â, INDIA, Tamil Nadu, Nilgiri Hills, Ootacamund, 2150 m a.s.l., sifting 
litter under bushes in ravine; 21. XI. 1972; leg. C. Besuchet & I. Lobl [41]. 

Paratypes: 2c5,6 9;2c?,2 9 (ZMMU), from same locality, collected together with 
the holotype. - 1 6 , 1 9 , Tamil Nadu, Nilgiri Hills, Coonoor, 1600 m a.s.l., forest, sifting litter; 
22.XI.1972; leg. C. Besuchet & I. Lobl [43]. - 3 9 , same, 6 km E of Coonoor, forest, sifting 
litter; 22.XI.1972; leg. C. Besuchet & I. Lobl [42]. -16, Kerala, NW of Anaimalai Hills, 
Nelliampathi Hills, Kaikatty, 900 m a.s.l., forest, sifting litter near brook; 30.XI.1972; leg. C. 
Besuchet & I. Lobl [58]. -19, Madras (= Chennai), Cardamom Hills, 6 km NE of Kumily, 
700 m a.s.l., forest, sifting litter; 3.XI.1972; leg. C. Besuchet & I. Lobl [7]. 

ETYMOLOGY: The species name, an adjective, refers to the peculiar shape of the 
maie palpai tibia. 

DlAGNOSlS: The new species can be easily recognized by the peculiar shape of 
the maie palpai tibia. 

DESCRIPTION: Maie (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. Tml 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. Médian mem- 
brane very long, protruded forward and covering distal part of embolus. Radical part 
of embolie division elongated and lacking of outgrowths. Base of embolus wide, bent 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



581 




Figs 64-67 

Paracymboides tibialis sp. n., 6 & 9 paratypes from Ootacamund, Tamil Nadu. (64-65) Right 
palp, retrolateral and ventral views, respectively. (66) Embolie 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). Tml 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 maie. 

TAXONOMIC REMARKS: Paracymboides tibialis sp. n. is similar to P. aduncus 
sp. n.: their embolie division is almost identical. Thèse species can be easily distin- 



582 



A. V. TANASEVITCH 




Figs 68-72 

Paracymboides aduncus sp. n., S holotype, 9 paratype from Cardamom Hills, Muttapatti. 
(68-69) Right palp, retrolateral and prolateral views, respectively. (70-71) Palpai tibia, dorsal and 
prolateral views, respectively. (72) Epigyne, ventral view. 

guished by the shape of the palpai 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: S , INDIA, Tamil Nadu, Palni Hills, 23 km W of Kodaikanal, Lake Berijam, 
2150 m a.s.l., Rhododendron forest, sifting litter; 14.X1.1972; leg. C. Besuchet & I. Lôbl [26]. 

Paratypes: 1 6 , 1 9 , INDIA, Kerala, Cardamom Hills, 13 km NE of Munnar, 1900 m 
a.s.l., forest, sifting litter; 26.XI.1972; leg. C. Besuchet & I. Lôbl [51]. - 1 9 , Cardamom Hills, 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



583 



near Munnar, Muttapatti, 1700 m a.s.l, forest, sifting litter under tree ferns; 24.XI.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 palpai tibia outgrowth. 

DlAGNOSIS: The new species can be easily recognizable by the peculiar shape 
of the maie palpai tibia. 

Description: Maie (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. Tml 0.31. Metatarsus IV without tricho- 
bothrium. Palp (Figs 68-71): Palpai tibia with a long and narrow outgrowth starting at 
retrolateral side and running orthogonally of main palpai axis to prolateral side. 
Paracymbium very long, narrow, strongly curved. Distal suprategular apophysis short, 
wide and rounded. Médian membrane very long, protruded forward and covering distal 
part of embolus. Radical part of embolie 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). Tml 0.32. Epigyne and vulvae as shown in Figs 72, 116. Body and 
leg coloration, as well as chaetotaxy, as in maie. 

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 â ,1 9;2 6 ,2 9 (ZMMU), India, Madras (= Chennai), Anaimalai Hills, 
18 km N of Valparai, 1250 m a.s.l., forest, sifting litter; 18. XI. 1972; leg. C. Besuchet & I. Lôbl 
[35]. 

REMARKS: This species was hitherto known only from Népal (Wunderlich, 
1973, 1983), and is here recorded from India for the first time. 

Distribution: Népal Himalayas and southern India. 

Pelecopsis indus sp. n. Figs 73-80 

Holotype: cT, INDIA, Uttar Pradesh, Garhwal, above Pauri, northern slope, 1900 m 
a.s.l., Quercus dry forest, sifting leaf litter and moss; 28.X.1979; leg. I. Lôbl [28]. 

Paratypes: 1 cT , 3 9 , from same locality, collected together with the holotype. - 1 <?, 
4 9, Uttar Pradesh, Kumaon, Rangarh, 2000 m a.s.l., forest in ravine, sifting leaf litter; 
9.X.1979; leg. I. Lôbl [6b]. -19, 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]. - 1 9 , 
Kumaon, Naini Tal, 2000-2100 m a.s.l., forest in ravine, sifting leaf litter and moss; 8.X.1979; 
leg. I. Lôbl [5b]. - 1 S , Himachal Pradesh, Kulu Valley, Naggar, 1850 m a.s.l., rotten stump of 
Cedrus, under bark; 16.X.1988; leg. S. Vit [25]. - 1 <J, Kulu Valley, Vashisht Baths (N of 
Manali), 1900 m a.s.l.; 13.X.1988; leg. S. Vit [18]. -19, PAKISTAN, Swat District, Kalam, 
2100 m a.s.l., Quercus forest, sifting leaf litter; 12. VI 983; leg. C. Besuchet & I. Lôbl [9b]. - 



584 



A. V. TANASEVITCH 




Figs 73-80 

Pelecopsis indus sp. n., S & 9 paratypes from Pauri, Uttar Pradesh. (73-74) Maie carapace, 
latéral and dorsal views, respectively (not to scale). (75-76) Right palp. retrolateral and prolateral 
views, respectively. (77) Palpai tibia, dorsal vievv. (78-80) Epigyne and vulva, ventral, dorsal 
(inclined 90°) and anterodorsal views, respectively. 



1 9 . S of Saidu Sharif, Murghazar, leaf litter under Platanus, 1300 m a.s.l.; 8.V.1983; leg. C. 
Besuchet & I. Lôbl [2b]. - 1 S, Ushu Valley, upper reaches of Kalam River, 2300 m a.s.l., 
Cedrus forest, sifting leaf litter under Corylus; 15.V.1983; leg. C. Besuchet & I. Lôbl [13b]. - 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



585 



1 9 , Malam Jabba, 2300 m a.s.l., Pinus forest, sifting under Pinus and Corylus; 9.V.1983; leg. 
C. Besuchet & [. Lobl 14c]. 

Comparatyvematerialexamined: Pelecopsis minor, SMF60195,3 6,2 9 paratypes, 
Dundgovi Aimak, 20 km S of Somon Delgertsogt, 1480 m a.s.l.; 9.VI.-1 3.VII.1967; leg. Z. 
Kaszab, det. J. Wunderlich. 

ETYMOLOGY: The spécifie name, a noun in apposition, refers to an inhabitant of 

India. 

Diagnosis: The new species is characterized by the smooth surface of its cara- 
pace (without pits), by the shape of the maie palpai tibia, as well as by the peculiar 
shape of the ventral plate of the epigyne. 

DESCRIPTION: Maie (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. Tml 0.39. Ail 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. 
Embolie 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. Tml 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 maie. 

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 présence a 
trichobothrium on metatarsus IV, by the smooth carapace surface without pits, as well 
as by some détails of the genitalia. 

RANGE: Himalayan. 

Piniphantes himalayensis (Tanasevitch, 1987) 

Material: 1 9 , PAKISTAN, Swat District, above Utrot, 2800 m a.s.l., Abies & Cedrus 
forest, sifting rotten coniferous litter; 13.V.1983; leg. C. Besuchet & I. Lobl [11c]. 

REMARKS: This species was hitherto known from Kashmir (Thaler, 1987) and 
from numerous localities in Népal (Tanasevitch, 1987). 

RANGE: Himalayan. 

Porrhomma pygmaeum (Blackwall, 1834) Fig. 117 

Material: 1 9 (MSNV), PAKISTAN, Northern Areas, Ghangche District, Ghangche, 
Kaplu Ghwari, 2480 m a.s.l.; 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 S (MSNV), PAKISTAN, Karakorum. Baltistan, Shalabot, 1700 m a.s.l.; 
15.VIII.1976; leg. G. Osella. - 1 S (MSNV), Northern Areas, Skardu District, Skardu, Pakova, 
2300 m a.s.l.; 27 .VII. 1976; leg. G. Osella. - 1 6 (MSNV), Skardu. Lake Satpara, 2700 m a.s.l.; 
1. VII. 1976; leg. G. Osella. -3 6 (MSNV), Ghangche District, Khaplu, Sciaiak Channel, 2400 
m a.s.l.; 10.VII.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 9 . PAKISTAN, Hazara District. Kâghân Valley, 1450 m a.s.l., Malkandi 
Forest, rotten coniferous stub; 29 .VI. 1985: leg. S. Vit [PAK-85/3]. - 4 S , 8 9 , INDIA, Himachal 
Pradesh, Kulu Valley, Naggar. 1850 m a.s.l.. litter; 16.X.1988; leg. S. Vit [24]. - 3 9, 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 6\ 1 9 (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 9, Garhwal, 10 km W of Chamba. 2200 m a.s.l.. sifting mosses under bushes; 
20.X. 1979; leg. I. Lôbl [17]. - 1 SA 9 (ZMMU), Garhwal, 4 km S ofBhatwari, 1400 m a.s.l., 
timberline. degraded forest. sifting moss and fern litter; 23.X.1979; leg. I. Lôbl [22]. - 1 S , 
Garhwal. above Pauri. northern slope, 1900 m a.s.l., Quercus dry forest, sifting leaf litter and 
moss; 28.X. 1979: leg. I. Lôbl [28]. - 1 S , 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: S , 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 nevv species was found. 

DlAGNOSlS: The new species is characterized by the peculiar shape of the split 
palpai tibia. 

DESCRIPTION: Maie. 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. Médian membrane well developed, 
protruded forward, distally extended. Radical part of embolie division liât, 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 




Figs 81-85 



Tapinocyboides bengalensis sp. n., 6 holotype. (81-83) Right palp. retrolateral, prolateral and 
dorso-prolateral views, respectively. (84) Palpai tibia, dorsal vievv. (85) Distal suprategular 
apophysis and médian 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 embolie division and no sulci on the maie 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.VIII.1976; leg. G. Osella. 



588 



A. V. TANASEVITCH 




Figs 86-88 

Tchatkalophantes baltistan sp. n.. S holotype. (86-88) Epigyne. ventral, dorsal (upside down) 
and latéral views. respectiveh . 

Etymology: The spécifie 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 biown, 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). FeI0.90 long. Chaetotaxy: Fel: 
0-1-0-0; TO: 2-1-1-0, HII: 2-0-1-0. III-IY: ?; Mtl: 1-0-0-0. II-IV: ? T ml 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 médian plate triangular. 

Maie unknown. 

TAXONOMIC REMARKS: The new species is similar to the Mongolian T. hyper - 
auritus (Loksa. 1965) and differs by a shallower dépression in the distal part of the 
proscape. as well as by the shape of the posterior médian plate: in T. baltistan sp. n. it 
wider than high vs higher than wide in T. hyper auritus. 

Distribution: Known from the type locality only. 

Tenuiphantes tennis (Blackwall. 1852) 

Material: 2 9 (MSNY ). PAKISTAN. Karakorum. Baltistan. Shalabot. 1700 m a.s.l.: 
15.VHL1976; leg. G. Osella. - 1 6 (MSNV). Northern Areas. Gilgit District. Bagrot Valley, trap 



UNYPHIID SPECIES FROM PAKISTAN A\D INEHA 



589 




Figs 89-96 

Tiso incisus sp. n., 6 & 9 paratypes f r0 m Jutogh. Himachal Pradesh. (89) Right palp. retro- 
lateral vievv. (90) Paracymbium. (91-92) Palpai tibia, dorsal and prolateral views. respectively. 
(93) Distal suprategular apophysis. (94) Embolie division. (95) Epigyne. ventral view. i96> 
Vulva, dorsal vievv. 



#5: 25.X. -2. XI .2008: leg. L. Latella & R. Ahmed. - 1 5 (MSNV), Bagrot Valley: 17A1.2008: 
leg. L. Latella. 

Remarks: The species is here recorded from the Himala>as and Pakistan for 
the first time. 



Range: European-Ancient Mediterranean. 



590 



A. V. TANASEVTTCH 



Tiso incisus sp. n. Figs 89-96 

Holotype: 6. INDIA. Himachal Pradesh. Jutogh. 10 km W of Shimla. 2000 m a.s.l.. 
leaf litter near foot of rock: 29.X.1988: les. S. Vit [37]. 

PARATYPES: 13 6.19 9;5 d,3 9 (ZMMU), from same locality. collected together with 
the holotvpe. -12. Himachal Pradesh. Kulu Valley, S of Manali. Chijoga. 1900 m a.s.l., hollow 
in Quercus; 14.X.1988: leg. S. Vit [21]. - 1 6 , 2 9. Uttar Pradesh. Garhwal. 10 km E of 
Dhanolti. 2450 m a.s.l.. northern slope. Quercus forest. sifting litter: 21. X. 1979; leg. I. Lôbl 
[21b].- 1 9 . PAKISTAN. Swat District. S of Saidu-Sharif. Marghuzar. 1300 m a.s.l.. sifting leaf 
litter under Platanus; 8.V.1983: leg. C. Besuchet & I. Lôbl [2b]. - 1 S, 3 9 , Malam Jabba. 
2300 m a.s.l.. Pinus forest. sifting litter under Pinus and Conius: 9.V.1983: leg. C. Besuchet & 
I. Lôbl [4c]. - 1 6 . same. 2500-1600 m a.s.l.. under stones; 18.V.1983: leg. C. Besuchet & I. 
Lôbl [17a].- 1 d,5 9 , same, 2500-2600 m a.s.l.. Abies forest. sifting litter and moss: 18.V.1983; 
leg. C. Besuchet & I. Lôbl [17b]. - 1 6.3 9 , Kalam. 2100 m a.s.l., Quercus forest. sifting leaf 
litter: 12.V.1983: leg. C. Besuchet & I. Lôbl [9b]. - 7 9 , Ushu Valley, upper reaches of Kalam 
River. 2300 m a.s.l.. Cedrus forest. sifting leaf litter under Corylus; 15.V.1983; leg. C. Besuchet 
& I. Lôbl [13b]. - 1 S , 1 5 . above Miandam. 2400-2500 m a.s.l.. Abies forest. sifting litter and 
moss: 17.V.1983: leg. C. Besuchet & I. Lôbl [15b]. - l S, 1 9, same. 2400-2500 m a.s.l.. 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. 

DlAGXOSlS: The new species is characterized by the peculiar shape of the palpai 
tibia and the embolie division, as well as by the shape of the ventral plate of the 
epigyne. 

Description: Maie. 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. Tml 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 différent 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 embolie 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). Tml 0.43. Abdomen 0.80 long. 0.55 wide. Epigyne 
and vulva (Figs 95-96): Ventral plate of epigyne long and narrow. Réceptacles sphe - 
rical. far apart from each other. Body and leg coloration, as well as chaetotaxy, as in 
maie. 

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 embolie division: in T. incisus sp. n. the loop of the embolus is almost orthogonal 
to the main axis of the palp. whereas in T. aestivus the loop lies almost in the same 
plane; there are also other small différences in the shape of the palpai tibia and the 
embolie division. The female differs by the shape of the ventral plate of the epigyne: 
in T. aestivus the ventral plate is triangular. in T. incisus sp. n. it is as a narrow trans- 
versal stripe. 

Range: Himalayan. 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



591 




Figs 97-100 

Tiso (?) indianus sp. n., 6 paratype. (97-98) Right palp, retrolateral and prolateral views, 
respectively. (99) Palpai tibia, dorsal view. (100) Palpai tibia and paracymbium, latéral view. 



Tiso (?) indianus sp. n. Figs 97-100, 118 

Holotype: 6 , INDIA, West Bengal, Darjeeling District, Tiger Hill, 2500-2600 m a.s.l., 
near top, forest, sifting litter; 18.X.1978; leg. C. Besuchet & I. Lôbl [19]. 

Paratype: 1 S , from same locality, collected together with the holotype. 

ETYMOLOGY: The spécifie name is derived from the name of the country of 

origin; adjective. 

DlAGNOSiS: The new species is characterized by the peculiar shape of the 
embolus and the palpai tibia. 

Description: Maie (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+039+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. Tml 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 




FlGS 101-104 

Maie carapace of Walckenaeha martensi Wunderlich, 1972, carapace, latéral view. (101-103) 
Spécimens from Tonglu, West Bengal. (104) Spécimen from Thare Pati, Népal. 



shaped distally. Radical part of embolie 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 représentatives of the genus Paracymboides gen. n., but the structure of the 
embolie division is quite différent 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 Tiso Simon, 1884 is thus only provisional. 

Distribution: Known from the type locality only. 

Walckenaeria martensi Wunderlich, 1972 Figs 101-104 

Walckenaeha nepalensis Wunderlich, 1972 syn. n. 

Type material examined: Walckenaeria martensi, SMF 25298/1, S holotype, NEPAL, 
Khumbu. Everest Région at confluent of Imja- and Phunki-Drangka rivers, Betula forest, 3200- 
3250 m a.s.l.; 30.IX.-2.X.1970; leg. J. Martens. - SMF 25299/7, 1 6, 6 9 paratypes, same 
locality, collected together vvith holotype. - SMF 25300, 2 9 paratypes, NEPAL, Khumbu, Pare, 
Nangba-Tsangpo Valley, subalpine forest with Betula utilis and Rhododendron campanulatum, 
3350 m a.s.l.; 14.16.X.1970; leg. J. Martens. - SMF 25302/1, 1 subad. 9 paratype, Gorapani, 
Rhododendron forest along stream, 2850-2900 m a.s.l.; 10.-14.XII. 1969: leg. J. Martens. W. 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



593 




Figs 105-108 

Walckenaeria saetigera sp. n., 6 holotype (105-106), S paratype from Weiloi, Meghalaya (107) 
and £ paratype from Shillong, Meghalaya (108). (105-107) carapace, latéral (105, 107) and dor- 
sal (106) views, respectively. (108) Epigyne, ventral vievv (not to scale). 



nepalensis, SMF 25303/1, d holotype, NEPAL, Kathmandu-Tal, Mt Phulchoki, Quercus semi- 
carpifolia forest, 2600-2700 m a.s.L; 25.-30.I.1970; leg. J. Martens. - SMF 25304/4, 4 9 
paratypes, same locality, collected together with holotype. 

Other material: 2 â , INDIA, West Bengal, Darjeeling District, Tonglu, 3100 m a. s. 1., 
near top, under stones;16.X.1978; leg. C. Besuchet & I. Lôbl [16a]. - 1 6 , 3 9 , same, sifting in 
dwarf forest under brushes on pasture; 16.X.1978; leg. C. Besuchet & I. Lôbl [16b]. New records 
of W. martensv.l 9 , NEPAL, Bagmati Province, above Gui Bhanjyang, 2600 m a.s.l., northern 
slope, old Quercus forest, sifting litter and rotten wood under trees; 6.IV.1981; leg. I. Lôbl & A. 
Smetana [9]. - 1 S , 1 9 , lower Thare Pati, 3300 m a.s.L, sifting litter under fern and Acer near 
stream; 10.IV.1981; leg. I. Lôbl & A. Smetana [18b]. - 1 9 , lower Thare Pati, 3500 m a.s.L, Acer 
forest, sifting litter; 12.IV.1981; leg. I. Lôbl & A. Smetana [21]. - 3 9 , Malemchi, 2800 m a.s.L, 
sifting litter; 14.IV.1981; leg. I. Lôbl & A. Smetana [24]. -39, above Shermathang, 2900 m 
a.s.L, in faeces; 26.IV.1981; leg. I. Lôbl & A. Smetana [47b]. - 1 9, 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 S , 1 9 , Sankhuwasawa District, Kosi Province, 
NE of Kuwapani, 2350 m a.s.L, sifting litter and humus near spring; 5. IV. 1984; leg. I. Lôbl & A. 
Smetana [5]. - 1 9 , same, 2250 m a.s.L, sifting litter near rotten trunk, 6.IV.1984; leg. I. Lôbl & 
A. Smetana [6]. - 1 9 , NE of Mangmaya, 2300 m a.s.L, northern slope, dry forest, sifting rotten 
wood and litter; 6.IV.1984; leg. I. Lôbl & A. Smetana [7]. - 2 9 , southern part of Mangsingma, 
2200 m a.s.L, and rotten wood in ravine; 11.IV.1984; leg. I. Lôbl & A. Smetana [13]. -19, 
Induwa Kola Valley, 2000 m a.s.L, sifting litter and rotten wood; 14.IV.1984; leg. I. Lôbl & A. 
Smetana [18]. -19, same, 2100 m a.s.L, sifting litter; 17.IV.1984; leg. I. Lôbl & A. Smetana 
[27]. 

REMARKS: Ail three maies of W. martensi from Tonglu, West Bengal have palps 
identical to those of the type spécimens, but the shapes of the cephalic élévations are 
slightly différent from each other (see Figs 101-104) and similar to that of the holotype 



594 



A. V. TANASEVITCH 




FlGS 109-113 

Walckenaeria saetigera sp. n., S holotype. (109-110) Right palp, retrolateral and prolateral 
views, respectively. (111) Palpai tibia, dorsal view. (112-113) Embolie division and distal 
suprategular apophysis, différent aspects. 



of W. nepalensis, described from a single maie and several females from Népal 
(Wunderlich, 1972). A detailed comparison of the maie palps of ail 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 spécimens 
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., 9 paratype from Cardamom Hills, Muttapatti, India (114); 
Paracymboides tibialis sp. n., 9 paratype from Ootacamund, Tamil Nadu, India (115); P. adun- 
cus sp. n., 9 paratype from Cardamom Hills, Muttapatti, India (116); Porrhomma pygmaeum 
(Blackwall, 1834), 9 from Kaplu Ghwari, Pakistan (117); Tiso (?) indianus sp. n., S paratype 
from Tiger Hill, India (118); Walckenaeria saetigera sp. n., 9 paratype from Shillong, 
Meghalaya, India (119). (114-117) Vulva. (118) Embolie division and distal suprategular apo - 
physis. (119) Génital région, ventral view. Not to scale. 

Walckenaeria saetigera sp. n. Figs 105-113, 119 

Holotype: S, INDIA, Meghalaya, above Shillong, Khasi Hills, northern slope, 
1850-1950 m a.s.l., primary forest near Shillong Peak, sifting litter; 25 .X. 1978; leg. C. Besuchet 
&I.Lôbl[27]. 



596 



A. V. TANASEVITCH 



Paratypes: 2 9, from same locality, collected together with holotype. - 1 9, 
Meghalaya, above Cherrapunjee, Khasi Hills, 1200 m a.s.l., forest, near foot of rock, sifting 
litter; 26.X.1978; leg. C. Besuchet & I. Lôbl [28b]. - 1 d, Khasi Hills, Weiloi, 1700 m a.s.l, 
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 maie palpai tibia of this species. 

DiAGNOSiS: The new species is characterized by the peculiar shape of the maie 
carapace and the shape of the palpai tibia. The epigyne is also well distinct from that 
of congeners. 

Description: Maie (holotype). Total length 2.30. Carapace 1.08 long, 0.75 
vvide, 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. Tml 0.53. Ail 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 embolie 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 1.08 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). Tml 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. Réceptacles oval, well visible on both sides of ventral plate. 
Body and leg coloration, as well as chaetotaxy, as in maie. 

Variability: The shapes of the maie carapaces in the holotype and the 
paratype are slightly différent (Fig. 105 cf. Fig. 107). 

Taxonomic REMARKS: The epigyne bears some resemblance to that of the 
représentatives of the subgenus Wideria Simon, 1864, sensu Wunderlich (1972), but 
the embolie division shows similarities to that of the subgenus Prosopotheca Simon, 
1884. 

Distribution: Known from Meghalaya, northeastern India. 

ACKNOWLEDGEMENTS 

I am most grateful to ail persons whose collections I have used for the présent 
study. I wish to thank Peter Jàger and Julia Altmann (SMF) for providing types and 
comparative material. Spécial 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 (MHNG) 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 # 09-04-0 1365-a. 



LINYPHIID SPECIES FROM PAKISTAN AND INDIA 



597 



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REVUE SUISSE DE ZOOLOGIE 



Tome 118 — Fascicule 3 



Pages 

G att, Paul. Microphorella cassari sp. n., a new species of Microphorella 

Becker (Diptera: Dolichopodidae) from Tunisia 401-412 

Tran, Dao Thi 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 413-421 

Lienhard, Charles & Ashmole, N. Philip. The Psocoptera (Insecta: 
Psocodea) of St Helena and Ascension Island (South Atlantic) with a 
new record from South Africa 423-449 

Lienhard, Charles & Baz, Arturo. Redescription of the genus Marce - 
nendius Navâs (Psocodea: 'Psocoptera': Amphientomidae) with a 
key to western Palaearctic amphientomids 451-466 

Gonzâlez-Solis, David & Mariaux, Jean. An annotated list of the para- 
sitic nematodes (Nematoda) of freshwater fishes from Paraguay 
deposited in the Muséum of Natural History of Geneva 467-484 

Weber, Jean-Marc. Food habits of escaped Eurasian otters {Lutra lutra) 

in a suburban environment in Switzerland 485-489 

Hlavâc, Peter. Claude Besuchet, an eminent swiss coleopterists, 80 years 

old 491-512 

Senglet, Antoine. New species in the Zelotes tenuis-group and new of 

little known species in other Zelotes groups (Gnaphosidae, Araneae) 513-559 

Tanasevitch, Andrei V. Linyphiid spiders (Araneae, Linyphiidae) from 

Pakistan and India 561-598 



REVUE SUISSE DE ZOOLOGIE 
Volume 1 1 8 — Number 3 

Pages 

Gatt, Paul. Microphorella cassari sp. n., a new species of Microphorella 

Becker (Diptera: Dolichopodidae) from Tunisia 401-412 

Tran, Dao Thi 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 413-421 

Lienhard, Charles & Ashmole, N. Philip. The Psocoptera (Insecta: 
Psocodea) of St Helena and Ascension Island (South Atlantic) with a 
new record from South Africa 423-449 

Lienhard, Charles & Baz, Arturo. Redescription of the genus Marce- 
nendius Navâs (Psocodea: 'Psocoptera': Amphientomidae) with a 
key to western Palaearctic amphientomids 451-466 

Gonzâlez-Solis, David & Mariaux, Jean. An annotated list of the para- 
sitic nematodes (Nematoda) of freshwater fishes from Paraguay 
deposited in the Muséum of Natural History of Geneva 467-484 

Weber, Jean-Marc. Food habits of escaped Eurasian otters {Lutra lutra) 

in a suburban environment in Switzerland 485-489 

Hlavâc, Peter. Claude Besuchet, an eminent swiss coleopterists, 80 years 

old 491-512 

Senglet, Antoine. New species in the Zelotes tenuis-group and new of 

little known species in other Zelotes groups (Gnaphosidae, Araneae) 513-559 

Tanasevitch, Andrei V. Linyphiid spiders (Araneae, Linyphiidae) from 

Pakistan and India 561-598 



Indexed in Current Contents, Science Citation Index 



PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENÈVE 
CATALOGUE DES INVERTEBRES DE LA SUISSE. N os 1-17 (1908-1926) série Fr. 285.— 



(prix des fascicules sur demande» 



LE RHINOLOPHE (Bulletin du centre d étude des chauves-souris) par fascicule Fr. 35. — 

THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND 
DISTRIBUTION. WITH KEYS FOR DETERMINATION 

J. Nosek, 345 p., 1973 Fr. 30.— 

CLASSIFICATION OF THE DIPLOPODA 

R. L. Hoffman, 237 p., 1979 Fr. 30.— 

LES OISEAUX NICHEURS DU CANTON DE GENÈVE 
P. Géroldet, C. Glex & M. Maire 

351 p.. nombreuses cartes et figures. 1983 Fr. 45. — 

CATALOGUE COMMENTÉ DES TYPES D'ECHINODERMES ACTUELS 
CONSERVÉS DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI D UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ 
À LA CONNAISSANCE DES ECHINODERMES ACTUELS 

M. Jangoux, 67 p., 1985 Fr. 15.— 

RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE 
(COTENTIN-BAIE DE SEINE. FRANCE i 

Y. Finet, J. WLest & K. Mareda, 62 p.. 1991 Fr. 10.— 

GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: 
SHELLS AND RADULAS 

Y. Finet. J. WCest & K. Mareda. 1992 Fr. 30.— 

O. SCHMIDT SPONGE CATALOGUE 

R. Desqueyroux-Faundez & S.M. Stone, 190 p.. 1992 Fr. 40.— 

ATLAS DE RÉPARTITION DES AMPHIBIENS 
ET REPTILES DU CANTON DE GENÈVE 

A. Keller, V. Aellen & V. Mahnert. 48 p.. 1993 Fr. 15.— 

THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: 
A DOCUMENTED FAUNAL LIST 

Y. Finet, 180 p., 1995 Fr. 30.— 

NOTICE SUR LES COLLECTIONS MALACOLOGIQUES 
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 

J.-C. Cailliez, 49 p., 1995 Fr. 22.— 

PROCEEDINGS OF THE XlIIth INTERNATIONAL CONGRES S 

OF ARACHNOLOGY. Geneva 1995 (ed. V. Mahnert), 720 p. (2 vol.). 1996 Fr. 160.— 

CATALOGUE OF THE SCAPHIDILNAE (COLEOPTERA: STAPHYLINTDAE i 

(Instrumenta Biodhersitatis I). L LôBL, xii + 190 p.. 1997 Fr. 50. — 

CATALOGUE SYNONYMIQUE ET GEOGRAPHIQUE DES SYRPHIDAE (DIPTERA I 
DE LA REGION AFROTROPICALE 

(Instrumenta Biodh ersitatis II), H. G. Dirickx, x +187 p.. 1998 Fr. 50. — 

A REVISION OF THE CORYLOPHIDAE (COLEOPTERA i OF THE 
WEST PALAEARCTIC REGION 

(Instrumenta Biodhersitatis III). S. Bowestead, 203 p., 1999 Fr. 60. — 

THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE 
SOKOTRA ARCHIPELAGO 

(Instrumenta Biodh ersitatis IV). B. Schatti & A. Desvoignes. 

178 p., 1999 Fr. 70.— 

PSOCOPTERA (INSECTAi: WORLD CATALOGUE AND BIBLIOGRAPHY 
(Instrumenta Biodh ersitatis V). C. Lienhard & C. N. Smithers. 

xli + 745 p., 2002 Fr. 180. — 

REVISION DER PALÂARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA 
THOMSON. 1859 (COLEOPTERA. STAPHYLINTDAE) ( 1. Teil) 
(Instrumenta Biodh ersitatis VI). G. Sabella. Ch. Buckle. V. Brachat 

& C. Besuchet. vi + 283 p.. 2004 Fr. 100.— 

PHYLOGENY. TAXONOMY. AND BIOLOGY OF TEPHRITOID FLIES 
(DIPTERA. TEPHRITOIDEA I 

Proceedings of the "3rd Tephritoid Taxonomist"s Meeting. Geneva. 19.-24. July 2004" 
(Instrumenta Biodh ersitatis VII). B. Merz. vi + 274 p.. 2006 Fr. 100. — 



Volume 1 18 - Number 3 - 201 1 



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