QL

453.4

^NT Arachnologische

E

Arachnology 2012

O 2.9.-7.9.201 2 Ljubljana, Slovenia
www.european-arachnology.org

Heft 45

Karlsruhe, Juni 2013

ISSN 1018-4171

www.AraGes.de/aramit

r

Herausgeber:

Arachnologische Gesellschaft e.V.

URL: http://www.AraGes.de

Arachnologische

Schriftleitung:

Theo Blick, Senckenberg Gesellschaft für Naturforschung,

Terrestrische Zoologie, Projekt Hessische Naturwaldreservate, Senckenberganlage 25,

D-60325 Frankfurt/M., E-Mail; theo.blick@senckenberg.de, aramit@theoblick.de
Dr. Sascha Buchholz, Technische Universität Berlin,

Institut für Ökologie, Rothenburgstr. 12, D-12165 Berlin,

E-Mail: sascha.buchholz@tu-berlin.de

Gast- Editor für Artikel 2-4:

Dr. Matjaz Kuntner, University of Ljubljana,

Institute of Biology, Scientific Research Centre of the Slovenian Academy of Sciences and Arts, Novi trg 2, P. O. Box 306, SI-1001
Ljubljana, Slovenia, E-Mail: kuntner@zrc-sazu.si

Redaktion:

Theo Blick, Frankfurt Dr. Sascha Buchholz, Berlin

Dr. Jason Dunlop, Berlin Dr. Ambros Hänggi, Basel

Dr. Hubert Höfer &. Stefan Scharf, Karlsruhe (Satz und Repro, E-Mail: hubert.hoefer@smnk.de)

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Dr. Peter Jäger, Frankfurt/M. (D)

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Dr. Volker Mahnert, Douvaine (F)

Prof. Dr. Jochen Martens, Mainz (D)

Dr. Dieter Martin, Waren (D)

Dr. Uwe Riecken, Bonn (D)

Prof. Dr. Wojciech Star^ga, Warszawa (PL)

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Umschlagzeichnung: P. Jäger, K. Rehbinder
Arachnologische Mitteilungen 45: 1-53

APR 29 2015

Karlsruhe, Juni 2013

Arachnologische Mitteilungen 45: 1-3

Karlsruhe, Juni 2013

Preface to the section of the Proceedings of the 27th European Congress of
Arachnology, Ljubljana, 201 2 Sept. 2-7 (pp. 4-20)

doi: 1 0.5431 /aramit4501

Almost four years ago, du-
ring the 2009 congress in
Greece, I was called away
from the Russian party
only to be cornered by the
esteemed colleagues of the
ESA council and be asked
the unnerving question, to
which, of course, a negative
response was not an opti-
on: “Matjaz, how would
you feel about organizing
a future European con-
gress”? Three years later, in
September 2012, not only
European arachnologists,
but also many overseas col-
leagues came to Ljubljana with huge expecta-
tions from me and my crowd of enthusiastic,
yet inexperienced, sidekicks, of which only one
was a professional researcher, and the rest were
graduate or undergraduate students. 2012 was a
recession year, and with hundreds of our letters
asking for sponsorship of this “unique and pres-
tigious scientific event” ignored (though with a
handful of exceptions, see below), we had to pull
the event off on a shoestring. We did, however,
keep the fees low for numerous colleagues to be
able to attend, and in addition we sponsored an
unprecedented number of students that would
otherwise be unable to attend the meeting; no
fewer than 30 lucky international students were
thus awarded grants that in addition to waived
fees also covered their accommodation.

And so we (EZ lab) hosted, at the Scientific
Research Centre of the Slovenian Academy of
Sciences and Arts (ZRC SAZU) in Ljubljana,
between 2""^ and September 2012, 156 col-
leagues from 39 countries. The participants re-
presented most European countries, and those
from further afield came from Brazil, Uruguay,
Mexico, USA, Algeria, India, Pakistan, Japan,
China, Taiwan, Singapore and Australia. As
people told us, the congress was a lot of fun.

The event would never have been possib-
le without the amazing help from my EZ Lab

EGA

, SLOVENIA

2012

organizing team. Thank you Matjaz Gregoric,
Ren-Chung Cheng, Nina Vidergar, Kiemen
Candek, Urska Pristovsek and Miha Pipan, who
made the impossible things work, with further
help coming from Olga Kardos, Martin Turjak,
Tea Knapic, Klavdija Suen and selected student
grantees. I am also indebted to the wonderful
scientific committee, Drs. Simona Kralj-Fiser,
Ingi Agnarsson, Christian Kropf, Daiqin Li
and Miquel Arnedo. These colleagues helped
me select and invite four outstanding plenary
speakers from around the World. We were pri-
vileged to welcome and listen to the lectures of
Jason Bond (Auburn University, USA) on inte-
grative approaches to delimiting species and ta-
xonomy, Fiona Cross (University of Canterbury,
New Zealand) on olfaction based behaviour of a
mosquito-eating, and thus blood-sucking, jum-
ping spider from Africa, Eileen Hebets (Univer-
sity of Nebraska, USA) on interacting traits and
secret senses in arachnids, proposing them as
model organisms in studying behavioural evo-
lution and Fritz Vollrath (University of Oxford,
UK), who spoke of the secrets of silk. These lec-
tures were all fascinating and of high scientific
value. Most of them can be viewed online at
http://videolectures.net/ eca2012_ljubljana/.

We grouped the presentations into logical
sessions, some of which had to be parallel, with

2

M. Kuntner

non-overlapping presentations of the students
competing for awards. With the help of Wolf-
gang Nentwig we also organized a symposium
on DNA barcoding, where longer introductory
talks by Miquel Arnedo, Ingi Agnarsson and
Jeremy Miller introduced the concepts and re-
cent practices in discovering, delimiting and in-
ventorying species using DNA barcodes, while
more specific presentations introduced recent
European barcoding initiatives. There were also
two poster sessions, and many social events.
We congratulate again those students who won
awards, kindly provided by our sponsors, alt-
hough the selection amongst many high quality
student presentations was difficult. Within the
broad category “Systematics and Evolution”,
the awarded oral presentations were those of
Henrik Krehenwinkel (1), Elisabeth Lipke (2)
and Marija Majer (3), the awarded posters were
those of Jana Plfskova (1), Susanne Meese (2)
and Cheng-Ya Chang (3). Within the catego-
ry “Ecology and Behavior” the awards for oral
presentations went to Katrin Kunz (1), Shazia
Quasin (2) and Reiner Neumann (3), while
those for posters went to Urska Pristovsek (1),
Hsien-Chun Liao (2) and Klaas Welke (3).

The main congress sponsors in addition to
the European Society of Arachnology were the
Scientific Research Centre of the Slovenian

Academy of Sciences and Arts (http://www.zrc-
sazu.si/) and its EZ Lab (http://ezlab.zrc-sazu.
si/), the Slovenian Academy of Sciences and
Arts (http://www.sazu.si/en/o-sazu.html) and
the Slovenian Research Agency (http://www.
arrs.gov.si/en/dobrodoslica.asp). Further gene-
rous sponsorships came from Pensoft (http://
www.pensoft.net/). Bioform (https://www.bio-
form.de/), the British Arachnological Society
(http://britishspiders.org.uk/), the American
Arachnological Society (http://americanarach-
nology.org/) and the Arachnologische Gesell-
schaft (http://arages.de/).

Finally, although the congress saw nume-
rous talks and posters (the Abstract book can
be downloaded at http://ezlab.zrc-sazu.si/
eca2012#abstracts), the future of the proceedings
from the European Congress of Arachnology
may be bleak, as no more than five manuscripts
in total were submitted from the Ljubljana con-
gress, of which three withstood peer review and
were evaluated as appropriate for publication.
Perhaps it’s time for the society to reconsider
the future of publishing presentations from the
congresses in manuscript form, but rather move
towards publishing and indexing only presenta-
tion abstracts.

Matjaz Kuntner, Ljubljana

Preface

3

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Arachnologische Mitteilungen 45:4-7

Karlsruhejuni 2013

On the identity of the poorly known spider species Zelotes strandi (Araneae:
Gnaphosidae)

Christo Deltshev

doi: 1 0.543 l/aramit4502

Abstract. The taxonomic status and distribution of the poorly known gnaphosid species Zelotes strandi (Nosek,
1 905) is clarified, discussed and mapped on the basis of both the holotype (a single female from Turkey) and newly
collected material (males and females from Bulgaria). This species was originally described from the island of Prink-
ipo (= Büyükada, District of Istanbul, Turkey) and the holotype is currently housed in the Naturhistorisches Museum,
Vienna. Both sexes are here diagnosed and illustrated, whereby the male is described for the first time.

Keywords: Bulgaria, first description of male, new records, taxonomy, Turkey

Nosek (1905) described 23 new spider species from
the families Dysderidae,Theridiidae, Lycosidae, Age-
lenidae, Gnaphosidae and Thomisidae collected by
the well-known Austrian naturalist Arnold Penther
in different regions of Turkey, including the Island
Prinkipo (= Büyükada) in the Marmara Sea, near Is-
tanbul. From the material collected in the territory of
this island, Nosek described the species T^elotes stran-
di (Nosek, 1905) (originally in Prosthesima) on the
basis of a single female specimen. Since its original
description, there have been no further records con-
cerning the taxonomy and distribution of the species
(Chatzaki 2010, Senglet 2011, 2012, Platnick 2013).
Having had the opportunity to examine the collec-
tion of A. Nosek kept in the Vienna Museum - and
comparing it with material collected from different
regions of Bulgaria - it became possible to provide a
description of the previously unknown male togeth-
er with a redescription of the female based on new
drawings and photos.

Material and methods

The type material (a single female) was studied and
photographed, without removing the epigyne. The
spider material from Bulgaria was collected by hand
and by pitfall traps from various localities (Fig. 1).
The map of the Balkan Peninsula follows Abadjiev
(2001). Measurements of the legs were taken from
the dorsal side. Total body length includes chelice-

This contribution was presented at the 27th European Congress of
Arachnology, Ljubljana, 2012 Sept. 2-7.

Christo DELTSHEV, National Museum of Natural History, Bulgarian Aca-
demy of Sciences, Tsar Osvoboditel Blvd. 1 , 1 000 Sofia, Bulgaria, e-mail:
deltshev@gmail.com

submitted 4.1.201 3, accepted 27.3.2013, online 13.5.2013

rae. Photographs of the holotype were made using a
Nikon camera connected to a Leica MZ 12.5 stereo-
scope; those of the newly described male with a
Lumix camera connected to a Wild MD stereo-
scope. The drawings were made with a camera lucida
mounted on a Leica MZ 12.5 stereoscope.

The following abbreviations are used in the text
and figures: AME - anterior median eyes, PLE -
posterior lateral eyes, PME - posterior median eyes.

Taxonomy

Zelotes strandi (Nosek, 1905) (Figs 2-8, 10, 12, 14)
Prosthesima strandi Nosek, 1905: 126, f. 7

Material examined

TURKEY, Princess Islands, Prinkipo (Büyükada), Marmara
Sea, #249 coll. A. Penther, 9 holotype, 5.5.1902 (A. Pen-
ther leg.). BULGARIA: North Black Sea: Kavarna town,
N43.458498°, £28.361197°, alt. 122 m, 19, 14.08.1996;
Balchik town (dunes), N43. 406734°, £28.125257°, alt.
Ill m. Id 299, 17.6.1995; Albena resort, N43. 395426°,
£28.088753°, alt. 234 m. Id, 23.06.1996; id l9, 11.10.1997
(V. Popov leg.). South Black Sea: Arkutino (dunes).

The poorly known Zelotes strand!

5

N42.330895°, £27.734050°, 1 m alt., l6, 12.06.1996;
Ropotamo river, N42. 324380°, £27.730782°. 21 m alt.,
l9, 8.6.1996 (V. Popov leg.). Sushtinska Sredna Gora
Mountains: Panagyuriste town, N42. 508435°, £24.188566°,
533 m, 266 4$9, 9.10.1998; Panagyuriste town, Kolonijte,
N42.583370°, £24.190925°, alt. 1133 m, l6, 8.10.1997;
Panagyuriste town, Milej, N42. 541801°, £24.196221°,
alt. 622 m, 1$, 5.5.1997; Panagyuriste town, Nikulden,
N42.516859°, £24.181875°, alt. 606 m, l9, 21.5.1998; 29?,
08.09.1998; 566, 10.10.1998; 266 19, 14.5.2000; Panagyuri-
ste town, Sredoryak,N42. 551685°, £24.188457°, alt 696 m,
l9, 6.5.1998; Panagyuriste town, Sveti Ivan, N42. 515571°,
£24.183635°, alt. 572 m, 399, 2.6.1998; Strelcha town,
Barikadite, N42.574502°, £24.456701°, alt. 1284 m, 299,
2.6.1996 (S. Bazarov leg.).

Comparative material

Zelotes cingarus (O. P. -Cambridge, 1874). Bulga-
ria, Slavianka Mt, Kalimantsi vill., N 41.459735°,
23.489964°, alt. 462 m, l6 399, 09.11.2002 (S. La-
zarov leg.).

Depository

The holotype is housed in the Naturhistorisches Mu-
seum, Vienna. One pair (male and female) from Bul-
garia, Panagyuriste town have been deposited at each
of the following museums: Museum für Naturkunde,
Humboldt-Universität, Berlin and Senckenberg
Museum, Frankfurt am Main. The remaining 12
males and 18 females from Bulgaria are held in the
collections of the National Museum of Natural His-
tory, Sofia.

Diagnosis and affinities

Somatic characters corresponding well to those of
the genus. Male palp (Figs 8, 10) distinguished by
the long tapering tibial apophysis, coiled embolar
base, stout terminal apophysis with a bifid end and
posterior sclerite with a characteristic shape; bifid
distally. Female epigyne (Figs 12, 14) distinguished
by the almost straight and parallel epyginal late-
ral margins, posteriorly forming U-shaped pockets.
Anterior margins much wider than lateral ones. Z.
strandi resembles Zelotes cingarus (O. P.-Cambridge,
1874) (Figs 9, 11, 13, 15), but differs in the following
characters. In males of Z. strandi (Figs 8, 10) the
tibial apophysis is longer; the terminal apophysis is
longer and the bifid end is better expressed; the pos-
terior sclerite of the terminal apophysis is bifid dis-
tally. In females of Z. strandi (Figs 12, 14), the lateral
epigynal margins are less spaced, the spermathecae
are smaller and the epigynal ducts are less coiled.

The original figure of the epigyne of Z. strandi
by Nosek (1905) is very schematic and these features

Figs 2-7: Zelotes strandi: 2 - epigyne (type); 3 - epigyne (type),
figure of Nosek (1905); 4 - epigyne (Bulgaria, Panagyuriste); 5
- vulva (Bulgaria, Panagyuriste); 6 - male palp, ventral view; 7 -
male palp, retrolateral view

are not represented. The taxonomic value of the pic-
ture is thus low. It is effectively impossible to identify
the species using the original illustration and this is

6

C Deltshev

E

Figs 8-1 1 : Zelotes strandi: 8 - male palp, ventral view; 1 0 - male
palp, retrolateral view. Zelotes cingarus (Bulgaria, Slavianka Mt):
9 - male palp, ventral view; 1 1 - male palp, retrolateral view.
E - embolus; M - median apophysis; PS - posterior sclerit;TA -
terminal apophysis. Scale bar: 0.3 mm

probably the reason why Z. strandi was not recog-
nised again in previous studies.

Description

Male: Total length 5.3; cephalo thorax length 2.52;
cephalothorax width 1.94 mm. Leg lengths:

leg

I

II

III

IV

Femur

1.62

1.44

1.26

2.16

Patella

1.01

0.97

0.72

1.10

Tibia

1.26

1.01

0.90

1.44

Metatarsus

0.97

0.90

1.08

1.80

Tarsus

0.83

0.79

0.72

0.90

Total

5.69

5.11

4.68

7.31

Female: Total length 9.0; cephalothorax length 4.68;
cephalothorax width 3.06 mm. Leg lengths:

leg

I

II

III

IV

Femur

2.88

2.27

2.16

2,80

Patella

1.62

1.26

1:08

1.62

Tibia

1.98

1.55

1.26

1.98

Metatarsus

1.62

1.40

1.26

2.52

Tarsus

1.26

1.08

1.08

1.26

Total

9.36

7.56

6.84

10.18

Posterior eye row with eyes of equal size and
nearly equidistant. Chelicerae red-brown, armed
with 3 teeth on outer margin of cheliceral furrow and
2 denticles on inner margin. Carapace, sternum and
abdomen brown. Legs, spination: Fe: I-II d 2; II-IV
d 6. Ti: I-II spineless; II-IV spinose. Me: I-II v 2;
III-IV spinose.

Male palp (Figs 6-8, 10): Tibial apophysis long,
slender and tapering. Embolic complex with an S-
shaped base and a small, sclerotized embolus. Ter-
minal apophysis stout with a bifid end. Posterior

12 13

Figs 1 2-1 5: Zelotes strandi: 1 2 - epigyne, 1 4 - vulva. Zelotes cin-
garus (Bulgaria, Slavianka Mt): 13 - epigyne, 15 - vulva. Scale
bar: 0.3 mm

The poorly known Zelotes strand!

1

sclerite bifid distally. Median apophysis concave in
the middle.

Epigyne (Figs 2-4, 12): Long and relatively
narrow. Lateral epigynal margins anteriorly almost
straight and parallel, posteriorly forming U-shaped
pockets, anterior margins much wider than
lateral ones.

Vulva (Figs 5, 14): Copulatory ducts long, leading
to large, globular, slightly separated spermathecae.

Ecology

This species reaches altitudes of up to 1,200 m, but
prefers lowlands. At higher altitudes it is found un-
der stones, mainly at the edge of beach forests and
meadows, in lowlands in dry grasslands, close to oak
forests and dunes near the sea shore.

Phenology

Mature males were captured from April to Septem-
ber, females from May to October.

Distribution

Z. strandi seems to be widespread over the eastern
part of the Balkan Peninsula. Until now, it has been
captured in Bulgaria in several places, and in Turkey
on the Prinkipo (Biiyiikada) Island (Fig. 1).

Acknowledgements

I am especially grateful to my colleagues S. Lazarov, I.
Krusteva and V. Popov, for making their collected material
available, to C. Hörweg for access to the collections of the

Natural History Museum Vienna and to Dr S. Abadjiev
for computerizing the map. The research in the collections
of Natural History Museum in Vienna was supported
by a grant from the “European Commission’s Research
Infrastructure Action via the SYNTHESYS Project” -
V ienna-TAF-24.

References

Abadjiev S 2001 An atlas of the distribution of the butterflies
in Bulgaria (Lepidoptera: Hesperioidea ScPapilionoides).
Pensoft Publishers, Sofia-Moscow. 335 pp.

Chatzaki M 2010 New data on the least known zelotines
(Araneae, Gnaphosidae) of Greece and adjacent regions.

- Zootaxa 2564: 43-61

Nosek A 1905 Araneiden, Opilionen und Chernetiden. In:
Penther A. &E. Zederbauer: Ergebnisse einer naturwis-
senschaftlichen Reise zum Erdschias-Dagh (Kleinasien).

- Annalen des Naturhistorischen Hofmuseums Wien
20: 114-154

Platnick NI 2013 The world spider catalog, version 13.5
American Museum of Natural History. - Internet:
http://research.amnh.org/iz/spiders/catalog/ (accessed
January 2013)

Senglet A 2011 New species in the Zelotes tenuis-groMp
and new or little known species in other Zelotes groups
(Gnaphosidae, Araneae). - Revue suisse de Zoologie
118:513-559

Senglet A 2012 Civizelotes new genus, and other new or
little known Zelotinae (Araneae, Gnaphosidae). - Revue
suisse de Zoologie 119: 501-528

Arachnologische Mitteilungen 45: 8-14

Karlsruhe, Juni 2013

Ground-living spiders in wooded habitats under human influence on an island in
Finland

Seppo Koponen

doi: 10.5431 /aramit4503

Abstract. Spiders were collected by pitfall traps in the south-western archipelago of Finland. Wooded study habitats
on a small-sized (1.2 km^) island were: 1) natural open ash grove, 2) dense mixed grove (old overgrown wooded
meadow), 3) wooded aspen pasture and 4) wooded meadow, both restored ten years earlier, 5) natural wooded
meadow. Highest species and family numbers were found at the natural sites (1 and 5) and the lowest in the dense
grove (site 2). Linyphiidae dominated, both at species and individual level, in the groves. Lycosidae were abundant
on the wooded meadows and Gnaphosidae on the wooded pasture. The highest faunal similarities were between
the groves (70 % species in common) and between the wooded meadows (64 %). The lowest similarity was found
between the dense grove (17 %) and the ash grove (23 %) with the aspen pasture. Ten years after clearing, sites 3 and
4 had diverse spider faunas. The fauna at site 4 resembled that on the corresponding natural site (5), thus showing
restoration success. Altogether 84 species of spiders were caught. The proportion of Gnaphosidae (16 species found)
is high. Most species found in the study are common in south-western Finland and many occur across the whole
country. Pardosa lugubris was most dominant at three sites, P pullata and Diplostyla concolor both at one site. Two
species, Enoplognatha thoracica and Micaria fulgens, are included in the Finnish Red Data Book.

Keywords: Araneae, archipelago, groves, habitat restoration, wooded meadows, wooded pastures

The island group Jungfruskär comprises three island
and a few small islets. It is situated in the south-
western archipelago of Finland, between the Turku
archipelago and the Aland archipelago in an open
sound called Skiftet (Fig. 1). Jungfruskär is located in
the north-westernmost corner of the South-western
Archipelago National Park. Due to its exceptionally
diverse flora and fauna it is known as “the flagship of
the Park” (Lindgren 2000).

The islands have been used as grazing sites for
sheep, goats and cows for a few hundred years.
Around 1850 some permanent inhabitants were
settled on Storlandet (the study island), and after
that felling of trees, cutting of upper parts of de-
ciduous trees (pollarding) for fodder, and grazing
itself became more active. This human activity pro-
duced special semi-natural habitats, like wooded
meadows and wooded pastures. Wooded mead-
ows are situated on the most fertile soils. They are
rather open: usually 15-25 % coverage by copses
of deciduous trees and shrubs, the rest being open
meadow. Grass vegetation is rather short (due to
hay harvesting and grazing) and the sites are char-
acterized by many flowering plants, like the orchid

This contribution was presented at the 27th European Congress of
Arachnology, Ljubljana, 2012 Sept. 2-7.

Seppo KOPONEN, Zoological Museum, University ofTurku,

FI-2001 4Turku, Finland, e-mail: sepkopo@utu.fi

Dactylorhiza sambucina. Wooded pastures are
found on fertile, but more stony, sites. In addition,
also forests, dry meadows and shores were used as
pastures.

During the 1960s human activity diminished,
and many wooded meadows and other pastures
began to overgrow. Clearing and opening of shrub
lands started by volunteers in the 1980s and later also
by the staff of the National Park (Lindgren 2000).
This restoration activity has been very effective.

The aim of this paper is to document the compo-
sition and differences of ground-living spider com-
munities in habitats which have been under differ-
ent level of human activity on Storlandet, the main
island of Jungfruskär.

Study area, material and methods

Jungfruskär is isolated from larger, forest growing
land masses (islands). Larger islands to the south-
east, south-west and west lay ca. 20 km from Jung-
fruskär, and those to the east, north and north-west
ca. 10 km apart. The distances to the Finnish, Swed-
ish and Estonian mainland are 60, 120 and 170 km
respectively. The land area of the Jungfruskär island
group is ca. 2 km^ and that of the main island Stor-
landet 1.2 km^. Its geographic location is 60° 8.35’N,
21° 4.69’E.

The land elevation here, caused by the Ice Age, is
about half a meter per century. Therefore the highest
hills in the island are about 3000-4000 years old, and

submitted 17.12.2012, accepted 19.3.2013, online 13.5.2013

Ground-living spiders on an island, Finland

9

the present study sites have been above sea level for
about 400-1000 years.

Five wooded sites were studied on Storlandet.

The elevation of the sites varied from 2 to 5 m a.s.L

1) Open ash grove (natural). Trees: Fraxinus ex-
celsior and Alnus glutinosa\ abundant field layer
plants: Filipendula ulmaria, Urtica dioica, Anthr-
iscus sylvestris and Allium scorodoprasum.

2) Mixed dense grove (old overgrown wooded
meadow), Fig. 2. Trees and shrubs: Betula pubes-
cens, Alnus glutinosa, Prunus padus, Viburnum opu-
lus and Kibes alpinum; field layer: Milium effusum,
Geum rivale, Flipendula ulmaria and Silene dioica.

3) Wooded aspen pasture (junipers and coppice
removed in 1997-98). Trees and shrubs: Populus
tremula and Kibes alpinum; field layer: Fragaria
vesca, Kubus saxatilis, Primula veris and different
grasses. Also ground-laying rotten trunks of as-
pen. Before clearing it resembled site 2.

4) Wooded meadow (cleared in 1997-98). Trees: Al-
nus glutinosa, Morbus aucuparia and Prunus padus;
field layer: Deschampsia cespitosa, Filipendula ul-

maria, Kubus idaeus, Geranium sylvaticum and Si-
lene dioica. Before clearing it resembled site 2.

5) Wooded meadow (natural) on calcareous mo-
raine, Fig. 3. Trees: Alnus glutinosa and Betula
pubesce?2s; field layer: Convallaria majalis, Agrostis
capillaris, Anthoxanthum odoratum, Khinanthus
minor and Galium bore ale.

Ground-living spiders were collected using pitfall
traps from 22 May to 11 September, 2007. Ten traps
(diameter 6 cm, with propylene glycol and detergent,
and covers) were placed at each site. Material consists
of 1,299 adult, identifiable specimens, and is deposit-
ed in the Zoological Museum, University of Turku.

Results

Altogether, 84 species of spiders belonging to 12
families were caught by pitfall trapping (Tab. l).The
most species-rich families were Linyphiidae (41 spe-
cies), Gnaphosidae (16), Lycosidae (9) and Thom-
isidae (7). The structure of the spider communities
at the study sites is shown in Table 2. The highest

10

5. Koponen

Tab. 1 : Spider species collected in study habitats (sites 1 -5) using pitfall traps, 2007.

site

1

2

3

4

5

Araneidae (1)

Zygiella atrica (C. L. Koch, 1845)

1

Clubionidae (1)

Clubiona lutescensV^t^Xxvag^ 1851

2

Corinnidae (1)

Phrurolithus festivus (C. L. Koch, 1835)

2

1

Gnaphosidae (16)

Callilepis noctmma (Linnaeus, 1758)

2

Drassodes pubescens (Thorell, 1856)

4

1

Drassyllus praeficus (L. Koch, 1866)

7

Drassyllus pusillus (C. L. Koch, 1833)

2

8

Gnaphosa bicolor (Hahn, 1833)

8

Haplodrassus sigftifer (C. L. Koch, 1839)

6

2

Haplodrassus sihestris (Blackwall, 1833)

2

9

3

1

Micaria aenea Thorell, 1871

22

Micaria fidgens (Walckenaer, 1802)

1

Micaria pulicaria (Sundevall, 1831)

5

2

Micaria siibopaca Westring, 1861

1

Zelotes clivicola (L. Koch, 1870)

1

Zelotes latreillei (Simon, 1878)

1

16

1

2

Zelotes longipes (L. Koch, 1866)

1

1

Zelotes petrensis (C. L. Koch, 1839)

11

Zelotes subterraneus (C. L. Koch, 1833)

4

Linyphiidae (41)

Agnyphantes expunctus (O. P.-Cambridge, 1875)

1

Agyneta afinis (Kulczyhski, 1898)

3

2

Agyneta conigera (O. P.-Cambridge, 1863)

1

Agyneta Jackson, 1912

2

15

1

Agyneta subtilis (O. P.-Cambridge, 1863)

2

1

Allomengea scopigera (Grube, 1859)

1

Anguliphantes angulipalpis (Westring, 1851)

2

1

Bathyph antes gracilis (Blackwall, 1841)

1

11

Bathyphantes nigrinus (Westring, 1851)

1

Bathyphantes parvulus (Westring, 1851)

3

6

Centromerus arcanus (O. P.-Cambridge, 1873)

1

Ceratinella brevis (Wider, 1834)

2

1

3

2

1.

Ceratinella scabrosa (O. P.-Cambridge, 1871)

16

*Dicymbium nigrum (Blackwall, 1834)

33

4

6

5

Diplocephalus picinus (Blackwall, 1841)

16

5

1

Diplostyla concolor (Wider, 1834)

3

67

1

3

Erigone atra Blackwall, 1833

2

3

Erigonella hiemalis (Blackwall, 1841)

3

Gonatium rubens (Blackwall, 1833)

1

Gongylidiellum latebricola (O. P.-Cambridge, 1871)

1

Gongylidium rufipes (Linnaeus, 1758)

4

Macrargus carpenteri (O. P.-Cambridge, 1894)

1

Macrargus rufus (Wider, 1834)

4

Maro minutus O. P.-Cambridge, 1906

1

Maso sundevalli (Westring, 1851)

1

Ground-living spiders on an island, Finland

11

site

1

2

3

4

5

Micrargus herbigradus (Blackwall, 1854)

2

Microneta viaria (Blackwall, 1841)

3

Pana?nomops mengei Simon, 1926

1

Porrhomma pallidum]dicVsor\, 1913

1

2

Tapinocyba pallens (O. P.-Cambridge, 1872)

4

9

Tapinocyboides pygmaea (Menge, 1869)

2

Tapmopa longidens (Wider, 1834)

2

Tenuiphantes mengei (Kulczynski, 1887)

1

Tenuiphantes tenebricola (Wider, 1834)

17

27

Tiso vagans (Blackwall, 1834)

16

39

11

Walckenaeria antica (Wider, 1834)

2

Walckenaeria atrotibialis (O. R-Cambridge, 1878)

6

21

3

3

Walckenaeria cucullata (C. L. Koch, 1836)

3

2

1

Walckenaeria dysderoides (Wider, 1834)

1

Walckenaeria unicornis O. P.-Cambridge, 1861

1

Walckenaeria vigilax O. P.-Cambridge, 1861

1

1

1

Lycosidae (9)

Alopecosa puherulenta (Clerck, 1757)

4

62

2

Alopecosa taeniata (C. L. Koch, 1835)

1

1

Pardosa fulvipes (Collett, 1876)

8

9

Pardosa lugubris (Walckenaer, 1802)

88

4

72

81

12

Pardosa palustris (Linnaeus, 1758)

1

7

Pardosa prativaga (L. Koch, 1870)

1

1

Pardosa pullata (Clerck, 1757)

1

40

Trochosa spmipalpis (R O. P.-Cambridge, 1895)

6

7

Trochosa t err kola Thorell, 1856

17

2

48

47

24

Salticidae (1)

Neon reticulatus (Blackwall, 1853)

1

Segestriidae (1)

Segestria senoculata (Linnaeus, 1758)

1

Tetragnathidae (2)

Pachygnatha degeeri Sundevall, 1830

47

11

25

Pachygnatha listeri Sundevall, 1830

19

11

2

Theridiidae (3)

Enoplognatha thoracica (Hahn, 1833)

1

1

Euryopis flavomaculata (C. L. Koch, 1836)

8

Robertas lividus (Blackwall, 1836)

1

Thomisidae (7)

Ozyptila atomar ia (Panzer, 1801)

3

Ozyptila praticola (C. L. Koch, 1837)

32

8

1

3

Ozyptila trux (Blackwall, 1846)

3

2

Xysticus bifasciatus C. L. Koch, 1837

1

Xysticus cristatus (Clerck, 1757)

4

Xysticus erraticus (Blackwall, 1834)

1

Xysticus obscurus Collett, 1877

5

Zoridae (1)

Zora spinimana (Sundevall, 1833)

3

1

Total specimens

322

194

351

224

208

Total species

38

23

30

25

38

* editor's comment: only Dicymbium nigrum brevisetosum Locket, 1962 is known from Scandinavia

12

5. Koponen

species and family numbers were found in the most
natural sites (1 and 5), and smallest numbers in
the dense, shady mixed grove (2). Clear differences
were found in the family composition between sites.
Linyphiidae clearly dominated, both at species and
individual level, in the two studied groves. Lycosidae
were most abundant on the two wooded meadows
and Gnaphosidae on the wooded pasture (Tab. 2).

The six most abundant spider species at each site
are listed in Tab. 3. Among the ten most abundant
species in the groves (sites 1 and 2), the following six
were shared in common: Pardosa lugubris, Dicymbium
nigrum, Ozyptila praticola, Pachygnatha list er i, Ten-
uiphantes tenebricola and Diplocephalus picinus. Three
species, Pardosa lugubris, Pachygnatha degeeri and
Trochosa terricola,wQct among the top-ten at all three
more open sites (3-5: wooded pasture and wooded
meadows). In addition, Haplodrassus silvestris, Tiso
vagans and Trochosa spinipalpis were abundant at two
sites, i.e. in wooded meadows (sites 4 and 5). Deal-
ing with all five habitats, a Pardosa species was the
most dominant at four sites {P lugubris at three and
P pullata once), and Diplostyla concolor in the dense
grove. Species with high numbers at many sites also

Fig. 2: Mixed dense grove (site 2). - Photo: V. Rinne 2007

include Pachygnatha degeeri and Trochosa terricola.
Interestingly, only one species {Pardosa lugubris) was
common in the top-ten lists of the mixed grove and
wooded ash pasture. These are rather closely situated
sites, and a decade prior to trapping (i.e. before res-
toration work) the wooded pasture was rather similar
to the mixed grove. Only three of the total 84 species,
Ceratinella brevis, Pardosa lugubris and Trochosa ter-
ricola, were caught at all five study localities.

Faunal similarity between the study sites is shown
in Tab. 4. The highest similarity is between the two
groves (Sorensen index: 0.52) and two wooded
meadows (0.51), the smallest similarity to others is
seen in the wooded aspen pasture: with mixed grove
(0.15) and with ash grove (0.21). Interestingly, the
mixed grove has low species similarity with wooded
aspen pasture and wooded cleared meadow. Before
restoration, i.e. ten years prior to trapping, the nature
of these three sites resembled each other.

Discussion

The proportion of Gnaphosidae among the study
material is high, especially in terms of species num-
bers. Altogether, 16 gnaphosid species were found, i.e.
19 % of the total species. For comparison, only 7 %
of the Finnish spider fauna belongs to Gnaphosidae
(Koponen 2008a). Reasons for the large number of
Gnaphosidae are open and warm habitats as well as
the situation of Jungfruskär in the southernmost part
of Finland. High numbers of Gnaphosidae species
and specimens have also been found previously on
dry and open island habitats in the south-western
Finnish archipelago (e.g. Koponen 2000, 2008b).

Ten years after restoration work, i.e. the clear-
ing of sites 3 and 4, their spider fauna was rich and
clearly different (both at species and family level)
from the fauna found in their original habitat type
(site 2). Very probably the openness (more light and
warmth in summertime) is the key factor for a new
faunal composition of the restored sites 3 and 4. The
fauna of the cleared wooded meadow (site 4) clearly
resembled that of the more natural wooded meadow
(site 5). At least here, the restoration work has thus
been successful. Although some studies on differ-
ences in spider communities during woodland res-
toration have been done (e.g. Ryndock et al. 2012),
there seems to be no comparable data known from
adjacent countries.

One of the species discovered, Enoplognatha tho-
racica, has been listed as a vulnerable species (VTJ)

Ground-living spiders on an island, Finland

13

Fig. 3: Wooded meadow (site 5).
- Photo: V. Rinne 2007

in the recent Finnish Red Data Book (Pajunen et
ah 2010). Jungfruskär (where the species was found
on both wooded meadows) represents its third local-
ity in Finland. In addition, Micaria fulgens, found in
wooded aspen pasture, is listed as NT (nearly threat-
ened) in the Red Data Book.

In general, the spider fauna found on Jungfruskär
is typical for the south-western archipelago of Fin-
land (cf Koponen 2008b). Species with a general
southerly distribution dominated. Thus the record

of Xysticus obscuruSy known in Finland as a northern
or boreal species, is interesting. Koponen (2008b)
studied ground-living spiders in groves on two is-
lands ca. 35 km east of Jungfruskär. Among the most
abundant species in groves both on Jungfruskär, and
on these eastern islands, were e.g. Diplostyla concolor,
Pardosa lugubris and Tenuiphantes tenebrkola. Some
species, like the linyphiids Dicyrnbium nigrum and
Tiso vagans, were found in high numbers on Storlan-
det in Jungfruskär compared to earlier studies done

Tab. 2: The structure of spider communities. Sites 1) natural open ash grove, 2) mixed dense grove, 3) wooded aspen pasture,
4) wooded meadow, 5) natural wooded meadow.

Site

1

2

3

4

5

Species

38

23

30

25

38

Families

9

5

8

6

8

Specimens

322

194

351

224

208

Species ( %) of total

Linyphiidae

63.2

73.9

23.3

40.0

39.5

Lycosidae

13.2

8.7

16.7

20.0

21.0

Gnaphosidae

7.9

0

43.3

24.0

13.2

Thomisidae

2.6

8.7

3.3

5.0

15.8

Individuals ( %) of total

Linyphiidae

43.8

85.1

3.4

28.1

22.6

Lycosidae

36.6

3.1

52.4

60.7

49.0

Gnaphosidae

2.2

0

26.8

4.5

6.3

Thomisidae

9.9

5.7

0.3

0.5

8.7

14

S. Koponen

Tab. 3: Six most abundant spider species in the 5 sites.

Study site

%

1)

Open ash grove

Pardosa lugubris

27.3

Dicymbium nigrum

10.2

Ozyptila praticola

9.9

Pachygnatha listeri

5.9

Tenuiphantes tenebrkola

5.3

Trochosa terricola

5.3

2)

Mixed dense grove

Diplostyla concolor

34.5

Tenuiphantes tenebrkola

13.9

Walckenaeria atrotibialis

10.8

Agyneta ramosa

7.7

Pachygnatha listeri

5.7

Tapinocyba palleiu

4.6

3)

Wooded aspen pasture

Pardosa lugubris

20.5

Alopecosa pulverulenta

17.7

Trochosa terricola

13.7

Pachygnatha degeeri

13.4

Micaria aenea

6.3

Zelotes latreillei

4.6

4)

Wooded meadow

Pardosa lugubris

36.2

Trochosa terricola

21.9

Tiso vagans

17.4

Pachygnatha degeeri

4.9

Bathyphaiztes parvulus

2.7

Dicymbium nigrum

2.7

5)

natural woded meadow

Pardosa pullata

19.2

Pachygnatha degeeri

12.0

Trochosa terricola

1.5

Pardosa lugubris

5.8

Bathypha?ites gracilis

5.3

Tiso vagans

5.3

in the archipelago (Lehtinen et ah 1979, Clayhills et
ah 2000, Koponen 2000, 2008b). The present mate-
rial consists of 84 species, but the total number of
spiders recorded from the small (1.2 km^) island of
Storlandet in the rather isolated Jungfruskär island
group is as high as 167 (Clayhills et al. 2008). It is
more than one fourth of the known Finnish spider
fauna (Koponen 2008a).

Tab. 4: Similarity between study sites (Sorensen index).

Sites

1

2

3

4

5

1

-

0.52

0.21

0.38

0.45

2

-

0.15

0.33

0.33

3

-

0.36

0.35

4

-

0.51

5

-

Acknowledgements

I wish to thank Veikko Rinne and Anssi Teräs (University
of Turku) for help in the field and for sorting the material.
I am very grateful to Leif Lindgren (Finnish Forest and
Park Service) for important information about the nature
and history of Jungsfruskär and the archipelago in general;
he also helped with logistics.

References

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Arachnologische Mitteilungen 45: 15-20

Karlsruhe, Juni 2013

Male palp organ morphology of three species of ground spiders (Araneae,
Gnaphosidae)

Boris Zakharov & Vladimir Ovtcharenko

doi; 10.5431 /a ramit4504

Abstract. A detailed morphological account of the male copulatory organs of three species of ground spiders,
Sergioluscapulotus, Herpyllus propinquus and Callilepis pluto (Araneae, Gnaphosidae), is presented. The large sclerites
(subtegulum, tegulum and embolus) appear to be homologous in all spiders. Sergiolus and Zelando have a plesio-
morphic palp organization. The increased complexity in the male bulb organization creates a locking mechanism
that fixes the male palp position during intercourse in Callilepis, as well as in Encoptarthria, Trachyzelotes and Zelotes.
The palp of Herpyllus, together with Anzacia, Drassodes and Intruda, demonstrates progressive modification of the
male bulb.

Keywords: Callilepis pluto, ground-plan, Herpyllus propinquus, Sergiolus capulatus, tripartite male bulb

It is widely accepted that the male palp specifically
fits into the female epigynum of the same species.
The importance of male and female genitalia in spe-
cies identification has long been recognized, since
it was first used for this purpose (Westring 1861,
Menge 1866, Wagner 1886, 1888, Engelhardt 1910,
Comstock 1910, 1912).

Studies of male palp morphology show that the
plesiomorphic state for this organ is characterized by
a tripartite organization (Haupt 1983, Kraus 1978,
1984, Szombathy 1915). This type of palp consists
of three basic sclerites - a subtegulum, tegulum and
embolus - connected by three inflatable membranes:
the basal, medial, and embolar hematodochae. This
type of male palp organization was termed “hydrau-
lic”, in contrast to the other type of the male palp he
termed “glandular” (Kraus 1978, 1984). The latter is
characterized by the progressive fusion of all three
sclerites into one capsule, accompanied by complete
reduction of the membranes and two bulb muscles.
This progressive reduction of sclerites, membranes
and muscles evolved several times (Kraus 1978, 1984,
Huber 1994).

Gnaphosid spiders can be divided into three ma-
jor groups according to their male palp organization

This contribution was presented at the 27th European Congress of
Arachnology, Ljubljana, 2012 Sept. 2-7.

Boris ZAKHAROV, Department of Natural Sciences, LaGuardia Commu-
nity College of the City University of New York, New York, USA, e-mail:
bzakharov@lagcc.cuny.edu

Vladimir OVTCHARENKO, Department of Natural Sciences, Hostos
Community College of the City University of New York, New York, USA,
e-mail: vio@hostos.cuny.edu, ovtshare@amnh.org

submitted 19.11.2012, accepted 8.4.201 3, online 13.5.2013

(Zakharov &c Ovtcharenko 2011). The closest condi-
tion to the ancestral type of male palp was observed
in Zelanda erebus (L. Koch, 1873); a peculiar species
from New Zealand. Drassodes lapidosus (Walckenaer,
1802), Intruda signata (Hogg, 1900), and Anzacia
gemmea (Dalmas, 1917) demonstrate significant
“simplification” in the male palp construction. The
embolic division of these spiders tends to fuse with
the tegulum and, thus, transforms the palp into an
essentially bipartite structure. By contrast, spiders of
the genus Encoptarthria Main, 1954 demonstrate an
increase in the complexity of male palp organization
(Ovtcharenko & Zakharov 2007). Their palp con-
tains an additional sclerite positioned between the
tegulum and embolus, which probably functions as a
flexible bridge and facilitates movement between the
tegulum and embolus. This present study continues
the morphological study of the expanded male palps
of the ground spiders (cf Zakharov & Ovtcharenko
2011).

Materials and methods

Genital bulbs of the following species - which rep-
resent two groups of gnaphosid spiders (the Laro-
nius and Herpyllus groups) (Murphy 2007) - were
studied: Callilepis pluto Banks, 1896, Herpyllus pro-
pinquus (Keyserling, 1887) and Sergiolus capulatus
(Walckenaer, 1837). All three species were collected
as follows: USA, Black Rock Forest, Cornwall, NY,
4r24’29"N 74°01T8"W,June 1999, leg. A. Tanase-
vitch and V. Ovtcharenko, coll V. Ovtcharenko.

The male palps of ground spiders for this study
were prepared using a standard procedure (Comstock
1910, Sierwald 1990). The left palps were detached

16

B. Zakharov & V. Ovtcharenko

and submerged overnight in a weak watery solu-
tion of potassium hydroxide (KOH). It makes the
bulb expend to various degrees. The bulb was then
transferred into distilled water, where it continued to
inflate. All prepared palps were preserved in 75 %
alcohol. Drawings were made with the aid of a dis-
secting microscope (Nikon SMZ-U). Drawings were
scanned and corrected with the computer program,
Adobe Photoshop Lightroom 4.

Recent study on recognising homology status
demonstrates that homology based on topology is
the best criterion for male palp structures. A special
similarity is close to that of topology and each cri-
terion is better and contains fewer violations than
homology based strictly on function (Agnarsson &
Coddington 2008). This study supports the tradi-
tional view, and topology still remains the most relia-
ble criterion of homology. For this reason, in order to
reach a decision on the homologous status of a par-
ticular structure of the bulb, the following classical
and widely applied criteria were used: 1) position of
the structure; 2) morphological similarity with other
known structures; 3) correspondence of the structure
with other characteristics (Remane 1956, Patterson
1982, Coddington 1990, Sierwald 1990).

As noted above, it is generally accepted that the
tripartite genital bulb in male spiders is a plesiomor-
phic characteristic (Platnick & Gertsch 1976, Kraus
1978, Haupt 1983, Sierwald 1990). This present
study supports the conclusion that the large sclerites
(subtegulum and tegulum) are homologous across all
spiders (see also Kraus 1978, Coddington 1990, Si-
erwald 1990). These sclerites are organized around
a tube. This tube has an enlarged, closed end (fun-
dus), a long coiled tube (sperm duct), and a narrow
tube with an opening at the end (ejaculatory duct)
(Comstock 1910, 1912). This tube serves as a tempo-
rary sperm reservoir. Before mating, males fill their
palps with sperm, which is stored here until mating
occurs.

The terms proximal and distal here refer to the
position of a structure in relation to the trajectory of
the sperm duct. The structures that occupy a position
close to the fundus are considered proximally locat-
ed. On the other hand, the structures that are close to
the ejaculatory duct are referred to here as distal.

The terms median apophysis and conductor are
used to name tegular apophyses that are supposed to
be homologous within all gnaphosoids. An inflatable
membranous projection on the upper surface of the

first half of the tegulum - which is merely an out-
growth of the membranous walls of the tegulum and
is closely related to the tip of the embolus - is labeled
the conductor. The median apophysis is a heavy scle-
rotized structure that occupies a position more distal
on the tegulum than the conductor. It connects to the
tegulum via an inflatable membrane, and does not
directly associate with the embolus. The embolic di-
vision of the bulb is identified by the constriction of
the sperm duct and its transformation into the nar-
row ejaculatory duct. According to this definition of
the embolic division, the distal sclerotized tube is as-
signed to the embolic bulb division (Sierwald 1990).

Results

The retrolateral tibial apophysis in Sergiolus capula-
tus (Walckenaer, 1837) is a simple, massive, hooked
structure (Fig. 1). The basal and median hematodo-
chae are well developed. The subtegulum and tegu-
lum are open spirals with a single loop. The median
apophysis and conductor are absent. The embolus is
short, curved clock-wise, and grooved. The area of
the embolus, close to its tip, has a membrane (Fig. 1,
TM) which is associated with the embolus. The
function of this membrane is unknown; it probably
plays a supporting role during copulation. This mem-
brane does not connect to the tegulum. Instead it is a
membranous outgrowth of the base of the embolus.
Thus, taking into account the topological criterion of
homology (Agnarsson & Coddington 2008), it can-
not be regarded as a conductor, and we refer to it
as a “terminal membrane”. Proximally the embolus is
attached to the distal tubular membrane, which con-
nects it to the distal part of the tegulum.

In general, the bulb of the Sergiolus is very simple,
tripartite, has three well-separated major sclerites
(subtegulum, tegulum, and embolus), and lacks the
median apophysis and conductor.

The retrolateral tibial apophysis of Herpyllus pro-
pinquus (Keyserling, 1887) is short, broad, slightly
curved, and sharp at its tip (Fig. 2). The basal and me-
dial hematodochae are well developed. The subtegu-
lum and tegulum are heavily sclerotized rings. The
median apophysis is a simple hook. The conductor
is a comparatively simple, inflatable membrane. At
its tip this membrane is divided into two lobes. The
embolus is comparatively short. The proximal part of
the embolus is broad and fuses with the tegulum. Its
distal part is short, slender, hook-like, and rests in the
groove between the top lobes of the conductor.

Male palp organ morphology of ground spiders

17

Fig. 1 a-b: Sergiolus capulatus, left
palp; a. Ventral view; b. Lateral
view. BH - basal hematodocha;
Cy - cymbium; DTM - distal tu-
bular membrane; E - embolus;
ED - ejaculatory duct; MH - me-
dian hematodocha; RTA - retrola-
teral tibial apophysis; SD - sperm
duct; St - subtegulum; T - tegu-
lum;TIVl - terminal membrane.

In conclusion, the bulb of Herpyllus is among the
most modified male reproductive organs seen in the
ground spiders. The fusion of the embolus with the
tegulum transforms the male bulb of these spiders
into a bipartite structure.

The male of Callilepis pluto Banks, 1896 does
not have a retrolateral tibial apophysis (Fig. 3). The
cymbium of these spiders is very characteristic. It is
flattened; its length is two times longer than its width
and somewhat spoon-shaped. The bulbus is well-de-
veloped. A basal hematodocha connects the subtegu-
lum to the petiolus and the alveolus of the cymbium.
The median hematodocha connects the tegulum and
subtegulum. There is a distal tubular membrane be-
tween the tegulum and embolus that binds them

through the distal tegular projection on one side and
the broad base of the embolus on the other side.

The median apophysis is attached to the tegulum
via a flexible membrane, approximately two-thirds of
the way along the ventral part of the tegular ring. The
insertion of the median apophysis on the tegulum is
closer to the embolic division than the insertion of
the conductor. The shape of the median apophysis is
very unusual. It is divided into two large, massive, and
heavily sclerotized hooks. The conductor is a small,
weakly-developed outgrowth of the membrane that
covers the tegulum.

The embolic part of the bulb is connected to the
tegulum by the distal tubular membrane. This flex-
ible membrane permits the embolus to rotate around

Fig. 2a-b: Herpyllus propinquus,
left palp; a. Prolateral view; b.
Retrolateral view. BH - basal he-
matodocha; Co - conductor; Cy -
cymbium; E - embolus; MA - me-
dian apophysis; SD - sperm duct;
St - subtegulum;! - tegulum.

18

B. Zakharov & V. Ovtcharenko

Fig. 3a-c: Callilepis pluto; left palp; a. Antero-retrolateral view; b. Ventral view; c. Prolateral view. BH - basal hemato-
docha; Cy - cymbium; E - embolus; ED - ejaculatory duct; IS - intercalary sclerite;MA - median apophysis; SD - sperm
duct; St - subtegulum;T - tegulum.

the distal tegular projection as if it were an axis. The
embolus is very long, slender, slightly flattened and
semi-circularly curved. It has a broad basal part and
bulged outgrowths on its distal part (embolar distal
projection). The most peculiar structure found in the
bulb of this species is a long, narrow sclerite between
the tegulum and the embolus (Fig. 3, IS). Its posi-
tion, special relations with other sclerites of the bulb
and function are similar to the intercalary sclerite
found in Zelotes (Platnick & Shadab 1983). It allows
us to term this structure an “intercalary sclerite” here
too. Probably, this additional sclerite increases the
mobility of the embolus during copulation, as was
described for Zelotes (Senglet 2004, 2011, 2012).

Discussion

Analysis of the present material allows us to further
develop an understanding of the organization of the
male palp in gnaphosid spiders. This study supports
the previous observation that there are three basic
ground-plans in gnaphosid male bulb morphology
(Zakharov & Ovtcharenko 2011). The tripartite gen-
ital bulb in male spiders is a plesiomorphic charac-
teristic (Platnick & Gertsch 1976, Kraus 1978, 1984,
Haupt 1983, Coddington 1990, Sierwald 1990) and
includes three basic sclerites: a subtegulum, tegu-
lum and embolus that are bound together by inflat-
able membranes. These large sclerites (subtegulum.

tegulum and embolus) of all Entelegynae are ho-
mologous. The basic hematodocha — the membrane
that attaches the subtegulum to the alveolus of the
cymbium and the median hematodocha that binds
the subtegulum and tegulum - are also homologous
across all Entelegynae. The use of the term “termi-
nal hematodocha” (Comstock 1910, 1912) should be
avoided because its description and position in the
bulb has not been clearly identified. Instead, the term
“distal tubular membrane” is used here for the mem-
brane that connects the distal part of the tegulum
to the proximal end of the distal sclerotized tube or
embolus. The term “terminal membrane” is proposed
for the membranous outgrowth of the embolus.

Among ground spiders, the bulbs of Sergiolus and
Zelanda are closest in morphology to that of the an-
cestral male palp. All major sclerites and membranes
are present in the male bulbs of these spiders. Ad-
ditionally, some derived palp forms have increased
bulb complexity. The ground spiders of the genus
Callilepis^ as well as Encoptarthria, Zelotes and Tra-
chyzelotes (Miller 1967, Platnick & Shadab 1983,
Senglet 2004, 2011, 2012, Zakharov & Ovtcharenko
2011), have additional sclerites in the embolic divi-
sion. They have a distal sclerotized tube (in Enco-
ptarthrid) or an intercalary sclerite (in Callilepis and
Zelotes) between the tegulum and embolus that are
flexibly connected to each other. This additional scle-

Mole polp organ morphology of ground spiders

19

rite increases the mobility of the embolic part of the
bulb. Furthermore, on the embolic part of the bulb,
these spiders have subterminal and terminal apophy-
ses. The presence of a number of additional sclerites
in the embolar part suggests that increased complex-
ity in the male bulb organization creates a locking
mechanism during intercourse in these spiders (Sier-
wald & Coddington 1988, Huber 1994).

Herpyllus, together with the genera Anzacia,
Drassodes and Intruda (Zakharov & Ovtcharenko
2011), represents progressive reduction of the number
of sclerites in the male bulb. The proximal part of the
embolus and the distal end of the tegulum in these
spiders are fused. Thus, the embolus in these spiders
is firmly attached to the tegulum and the distal tubu-
lar membrane completely disappears. The other ma-
jor sclerites of the bulb in spiders of these genera are
also significantly reduced.

Male bulb evolution in ground spiders was not
linear, and analogous structures may appear inde-
pendently in different groups. Such parallel evolu-
tion of the male palp - characterized by a secondary
simplification through fusion of the apical and me-
dian bulbus sclerites and simultaneous reduction of
the extensible membrane - has occurred independ-
ently in many groups of spiders. It was observed in
orthognath (e.g. Theraphosidae), haplogyne (Sicari-
idae, Scytodidae, Pholcidae), and entelegyne spiders
{Castianeira, Corinnidae) (Kraus 1984, Huber 1994).
These observations suggest that there is a tendency
in male spider palp evolution for the “pyriform male
palp organ” to develop into the “glandular bulb”
(Kraus 1984). Gnaphosid spiders also follow this
major evolutionary trend, by which the embolus of
their bulb fuses with the tegulum and thus changes
the tripartite male palp into the bipartite. This proc-
ess took place independently many times in different
groups of the gnaphosid spiders. We observe this in
the present study in Herpyllus^ and it was previously
seen in Drassodes and Intruda (Zakharov &. Ovtch-
arenko 2011). Another phenomenon of the parallel
transformation of the male bulb happens with those
ground spiders whose male palpal organ undergoes
an increase of external construction complexity. Thus,
we have found the intercalary sclerite in the bulb of
Callilepis pluto. The topology, special similarity, and
function of this sclerite are the same as the intercalary
sclerite in Zelotes. However, these two genera belong
to different subfamilies of gnaphosid spiders and im-
ply that they may have developed independently.

Acknowledgments

The present study was supported in part by an American
Museum of Natural History (New York). We thank the
National Science Foundation PEET (Partnerships for
Enhancing Expertise in Taxonomy) program for providing
funds through grant DEB-9521631 for revision of Australa-
sian ground spiders (particularly the PI. of the project - Dr.
Norman Platnick), and Research Foundation of the City
University of New York for providing funds through grant
PSC-CUNY 43-582. Authors thank Ms. Patricia Malkin
for her excellent drawings. Dr. Sandra Dickinson for her
valuable help with the text and two anonymous reviewers
whose advice helped in preparation of this article.

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CAlO-91.1

Arachnologische Mitteilungen 45: 21-24

Karlsruhe, Juni 2013

Nachweise für Mecklenburg-Vorpommern neuer und seltener Spinnenarten
(Arachnida, Araneae)

Dieter Martin

doi: 1 0.543 l/aramit4505

Abstract. Records of new and rare spider species from Mecklenburg-Western Pomerania (Arachnida, Ara-
neae). The first records for Mecklenburg-Western Pomerania, Germany, of the species Mermessus trilobotus, Po-
rasteatoda tobulata and Araniello inconspicua are provided, together with noteworthy occurences of the rare species
Ero tuberculata, Jacksonella falconeri, Philodromus histrio and Oxyopes ramosus.

Keywords: faunistics, Germany

Mit der Neufassung der Roten Liste der Spinnen
Mecklenburg-Vorpommerns (Martin 2012) wurde
auch die Gesamtartenliste des nordöstlichen Bun-
deslandes aktualisiert. Sie umfasst 572 Arten von
Webspinnen (Araneae). Mittlerweile konnten vor al-
lem durch die Sammeltätigkeit von K. Rudnick (Ber-
gen auf Rügen), aber auch durch eigene Aufsamm-
lungen weitere drei Arten nachgewiesen werden, die
über die oben genannte Checkliste hinaus neu für
Mecklenburg-Vorpommern sind. Die Gesamtzahl
der Spinnenarten Mecklenburg-Vorpommerns er-
höht sich somit auf 575.

Besonders bemerkenswert sind die Erstnach-
weise von Mennessus trilobatus und Parasteatoda ta-
bulata in Mecklenburg- Vorpommern. Sie markieren
die jeweils nordöstlichsten Fundpunkte der offenbar
in Arealausweitung befindlichen Adventivarten in
Deutschland.

Das Belegmaterial befindet sich in der Samm-
lung des Verfassers. Die Nomenklatur richtet sich
nach Platnick (2012).

Mermessus trilobatus (Emerton, 1882)

Synonym Eperigone trilobata

1(5, Meesiger, Naturpark Mecklenburgische Schweiz
und Kummerower See, (MTB 2143, 53°48’46“ N,
12°54’06“ E, 6 m ü. NN), Bodenfalle 14.-30.8.2012,
Sandmagerrasen (leg. D. Martin).

Der erste europäische Nachweis der aus Nord-
amerika stammenden Art gelang 1981 in einem Bu-
chenwald bei Karlsruhe (Dumpert Platen 1985).
Seither erfolgte eine stetige Arealerweiterung über

Dr. sc. Dieter MARTIN, Lindenweg 11, 17213 Untergöhren,
e-mail: dieter_martin.untergoehren@t-online.de

eingereicht 7.1 0.201 2, angenommen 21 .2.201 3, online 13.5.2013

mehrere mittel- und südeuropäische Länder (Hels-
dingen 2009, Dolansky et al. 2009, Rozwalka 2010,
Nentwig et al. 2012). In Deutschland ist die Art weit
in den Norden vorgedrungen (Staudt 2012). Der
vorliegende Fund belegt die Art erstmalig für das
Bundesland Mecklenburg-Vorpommern und stellt
gleichzeitig den bislang nördlichsten Fundpunkt in
Deutschland dar (Abb. 1).

Die Art besiedelt ein breites Spektrum an Lebens-
räumen (Hänggi et al. 1995). Nach Kielhorn (2007)
werden Offenlebensräume (Grünland, Brachen,
nicht aber bewirtschaftete Äcker) präferiert, wobei
keine Bindung an bestimmte Feuchtigkeitsverhält-
nisse erkennbar ist. Andererseits stammen mehrere
Nachweise aus Waldlebensräumen (Buchenwälder,
Aue- und Feuchtwälder) (Dumpert & Platen 1985,
Kielhorn 2011). Auch bezüglich der Lichtverhält-
nisse verhält sich die Art also offenbar tolerant. Der
vorliegende Fund in einem süd-exponierten Sand-
magerrasen sowie teilweise die begleitende Araneo-
fauna {Alopecosa schmidti, Alopecosa barbipes, Cheira-
canthium virescens) lassen eine Thermophilie der Art
vermuten.

Parasteatoda tabulata (Levi, 1980)

Synonym Achaearanea tabulata
1(5, Rügen, Bergen, Plattenbau-Wohngebiet (MTB
1546, 54°24’N, 13°25’ E, 30 m ü. NN), 26.6.2012, auf
Gehwegplatten, Handfang (leg. K. Rudnick).

Die eusynanthrop an Häusern im städtischen
Bereich lebende Art (Komposch 1993) ist nach Plat-
nick (2012) holarktisch verbreitet und wurde bislang
in Nordamerika, Ostasien und Europa gefunden.
Die ersten europäischen Nachweise erfolgten durch
Moritz et al. (1988) aus Deutschland (Berlin-Mar-
zahn und Brandenburg) sowie Knoflach (1991) aus
Österreich (Innsbruck). Mittlerweile wurde die ihr

22

D. Martin

Mermessus triiobal US

i ■■

i . ■ ^

-

3

* Nn

' s

i

fl ^.3:

-r

'x

*

•

• '

• 1 1 ^ '

<0^ ♦ i A ' /

^ f

♦ J

♦ M

. •

f ’ ■ A ,
k *i V {yTp >

V- • ' • •, •• V . -

.f

•• '

^ *i

'

W

#

,#

i * •> • 7 r-t-

^ • ••• • i / d ^

= lA

— H

i -tw

i

Abb. 1: Nachweise von Mermessus trilobatus (nach Staudt 201 2).
Fig. 1 : Records of Mermessus trilobatus (after Staudt 201 2).

► vorliegender Neunachweis/new record

Areal expandierende Art in zahlreichen weiteren
mittel- und osteuropäischen Ländern nachgewiesen
(Nentwig et al. 2012, Sestakovä & Gajdos 2011).

Staudt (2012) verzeichnet neben den o. a. Fund-
orten in Berlin und Brandenburg zusätzlich zwei
Funde aus Sachsen und Thüringen sowie einen wei-
teren Fundpunkt aus Brandenburg (MTB 3046).

Der vorliegende Fund der sehr seltenen Kugel-
spinnenart ist damit nicht nur der Erstnachweis
für Mecklenburg-Vorpommern, sondern zugleich
der bislang nördlichste Fundpunkt in Deutschland
(Abb. 2).

Araniella inconspicua (Simon, 1874)

IB, Groß Molzahn (MTB 2231, 53°52’23“ N,
10°58’06“ E, 1 m ü. NN), Seggenwiese, Kescherfang,
9.5.2012 (leg. K. Rudnick).

Araniella inconspicua ist eine der seltensten Ara-
niella-Kcte.r\ Europas (Sacher 1984). Sie fehlte bis-
lang in Mecklenburg-Vorpommern, wurde aber in
Grenznähe in Niedersachsen (Lemke 2010) sowie

Abb. 2: Nachweise von Porasteatoda tobulata (nach Staudt 201 2).
Fig. 2: Records of Parasteatodo tabulato (after Staudt 2012).

► vorliegender Neunachweis/new record

in Schleswig-Holstein (M. Lemke, nach Staudt
2012) gefunden. Der vorliegende Nachweis aus
Nordwest-Mecklenburg schließt sich letzterem un-
mittelbar an.

Ero tuberculata (De Geer, 1778)

1(5, Göhren-Lebbin, OT Untergöhren (MTB 2541,
Sr2^’’2T N, 12°29’38“ E, 70 m ü. NN), 31.7.2012,
Aeronaut, an Hauswand ca. 3 m von einer Kiefer
entfernt (leg. D. Martin).

Die selten gefundene Spinnenfresserart ist in
Schleswig-Holstein, Niedersachsen und Branden-
burg nachgewiesen (Staudt 2012), fehlte aber bislang
in Mecklenburg-Vorpommern. Der vorliegende, in
der aktuellen Checkliste (Martin 2012) bereits be-
rücksichtigte Fund schließt diese Lücke.

Von Broen (1993) verweist auf eine Bevorzugung
trocken-warmer Standorte. Dem entsprechen auch
die Angaben bei Hänggi et al. (1995). Nach Lemke
(2008) lebt die Art auf den unteren Zweigen von Na-
delbäumen. Eine Nachsuche (Klopfen) an den Ästen

Neue und seltene Spinnenarten in Mecklenburg-Vorpommern

23

Abb. 3: Nachweise von Jacksonella falconeri (nach Staudt 2012).
Fig. 3: Records of Jacksonella falconeri (after Staudt 201 2).

► vorliegender Neunachweis/new record

der in der Nähe des Fangortes stehenden Kiefer blieb
allerdings erfolglos.

Jacksonella falconeri (Jackson, 1908)

l9, Rügen, Halbinsel Drigge, Boddenufer
(MTB 1745, 54T7’20“ N, 13T0’25“ E, 1 m ü.
NN), Hochstaudenflur, Kescherfang, 25.5.2010 (leg.
K. Rudnick).

Die Erstmeldung für die deutsche Fauna erfolg-
te durch Wunderlich (1972) sowie Moritz (1973).
Nach letzterem beflnden sich in der Sammlung des
Naturkundemuseums in Berlin allerdings Belege be-
reits aus dem Jahr 1902, die F. Dahl in Schleswig-
Holstein sammelte.

Obwohl bei Staudt (2012) mittlerweile zahlrei-
che Funde dokumentiert sind, fehlt die Art bislang
im gesamten Nordosten Deutschlands. Der vorlie-
gende Fund an der Südküste Rügens erweitert somit
das Nachweisgebiet beträchtlich (Abb. 3). Jacksonella
falconeri wurde in der aktuellen Checkliste Mecklen-
burg-Vorpommerns (Martin 2012) bereits berück-
sichtigt.

Die ökologischen Ansprüche der Art sind noch
unklar. Während Wunderlich (1973) Xerotherm-
hänge verzeichnet, nennt Moritz (1973) vor allem
verschiedene Feuchtlebensräume. Der vorliegende
Fund reiht sich hier ein. Nach Hänggi et al. (1995)
dagegen beziehen sich die meisten Nennungen auf
Waldlebensräume.

Philodromus histrio (Latreille, 1819)

1(5, Grabower Heide (MTB 2735,53T5’N, 11°35’E,
35 m ü. NN), Calluna-Wtidt, auf trockenem Heide-
kraut, 9.5.2012, leg. K.Rudnick).

Der vorliegende Fund ist der zweite Nachweis
der Art in Mecklenburg-Vorpommern. Erstmalig
wurde sie durch Buchholz &c Schirmei (2011) in den
Küstendünenheiden der Insel Hiddensee gefunden.
In Schleswig-Holstein existiert bislang auch nur ein
Nachweis im Lübecker Raum in unmittelbarer Nähe
zur Landesgrenze von Mecklenburg-Vorpommern
(M. Lemke, nach Staudt 2012). Südlich des aktuellen
Fundortes schließt sich ein Vorkommen in Sandtro-
ckenrasen in Niedersachsen an (Merkens 2000).

Oxyopes ramosus (Martini & Goeze, 1778)

2 subad., Grabower Heide (MTB 2735, 53T5’ N,
11°35’ E, 35 m ü. NN), Callm^a-Yieide^ auf trocke-
nem Heidekraut, 9.5.2012, leg. K.Rudnick).

Die zwei bislang existierenden Nachweise der
Art in Mecklenburg-Vorpommern - gleichzeitig die
nördlichsten in Deutschland - liegen bereits Jahr-
zehnte zurück (Rabeler 1931: Göldenitzer Hoch-
moor; Martin 1983: NSG „Ostufer der Müritz bei
Waren - jetzt Müritz Nationalpark). Der vorliegen-
de Fund bestätigt das aktuelle Vorkommen der Art
in Mecklenburg-Vorpommern.

Danksagung

Für seine unermüdliche Sammeltätigkeit danke ich Herrn
Kurt Rudnick, der mir neben zahlreichen anderen interes-
santen Spinnenfunden auch den Großteil der hier aufge-
führten Arten zur Bearbeitung überließ. Theo Blick sowie
den Gutachtern der Arachnologischen Gesellschaft danke
ich für ihre wertvollen Hinweise.

Literatur

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fährdeter Spinnen (Araneae) in der Mark Brandenburg.

- Arachnologische Mitteilungen 6: 12-25
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nen (Araneae) in Berlin und Brandenburg. - Märkische
Entomologische Nachrichten 9(1): 99-108

Kielhorn K-H 2011 Bemerkenswerte Spinnenfunde aus
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reich neue Kugelspinne (Arachnida, Aranei:Theridiidae).

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aus Schleswig-Holstein der Jahre 2004 bis 2007. -
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aramit3506

Lemke M 2010 Araniella inconspicua beim GEO-Tag der
Artenvielfalt, Preten 04.06. bis 05.06.2010. Arachnida:
Araneae und Opiliones. - Internet: http://spinnen-
forum.de/smf/index.php?topic=3994,0 (15.9.2012)
Martin D 1983 Die Spinnenfauna des Naturschutzgebietes
,Ostufer der Müritzf - Zoologischer Rundbrief für den
Bezirk Neubrandenburg 3: 1-40
Martin D 2012 Rote Liste der Spinnen (Araneae) Meck-
lenburg-Vorpommerns. Ministerium für Landwirtschaft,

Umwelt und Verbraucherschutz Mecklenburg- Vorpom-
mern, Schwerin. 64 S.

Merkens S 2000 Die Spinnenzönosen der Sandtrockenrasen
im norddeutschen Tiefland im West-Ost-Transekt -
Gemeinschaftsstruktur, Habitatbindung, Biogeographie.
Dissertation, Univ. Osnabrück. 165 S.

Moritz M 1973 Neue und seltene Spinnen (Araneae) und
Weberknechte (Opiliones) aus der DDR. - Deutsche
Entomologische Zeitschrift N.F. 20: 173-210 - doi:
10.1002/mmnd.l9730200106
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367 - doi: 10.1002/mmnd.l9880350428

Nentwig W, Blick T, Gloor D, Hänggi A & Kropf C 2013
araneae - Spinnen Europas, Version 1.2013. - Internet:
http://www.araneae.unibe.ch (4.1.2013)

Platnick NI 2012 The world spider catalog, version 13.0.

- Internet: http://research.amnh.org/iz/spiders/catalog
(15.9.2012)

Rabeler W 1931 Zur Fauna des Göldenitzer Hochmoores
in Mecklenburg (Mollusca, Isopoda, Arachnoidea,
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BF00406497

Rozwalka R 2010 Mermessus trilobatus (Emerton, 1882)
(Araneae: Linyphiidae) - nowy gatunek paj^ka dla fauny
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Araniella Chamberlin &Ivie, 1942 aus dem Naturhisto-
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des Naturhistorischen Museums Wien B 86: 243-249
Sestakovä A & Gajdos P 2011 Expansny druh snovacky
Parasteatoda tabulata Levi, 1980 (Araneae, Theridiidae)
na Slovensku. - Folia faunistica Slovaca 16: 169-172
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Wunderlich) 1972 Zur Spinnenfauna Deutschlands, XII.

Neue und seltene Arten der Linyphiidae und einige
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Arachnologische Mitteilungen 45: 25-29

Karlsruhejuni 2013

First record of a schizomid, Stenochrus portoricensis (Schizomida: Hubbardiidae),
in Slovakia

Jana Christophoryova, Anna Sestakova, Miroslav Krumpal & Peter Fenda

doi: 1 0.5431 /aramit4506

Abstract. The discovery of Stenochrus portoricensis Chamberlin, 1922 is the first record of the order Schizomida in
Slovakia. Juvenile specimens and adult females were found in heated greenhouses in the Botanical Garden of the
Comenius University in Bratislava, Slovakia. The main morphological characters of juveniles and adult females were
studied and photographed.

Keywords: Central Europe, faunistics, greenhouse, introduced species, short-tailed whipscorpion

The order Schizomida is a small group of arachnids,
currently represented by two recent families: Proto-
schizomidae Rowland, 1975 with two genera and
12 species; and Hubbardiidae Cook, 1899 with 48
genera and 271 species worldwide (Harvey 2007,
Armas 2010, Monjaraz-Ruedas 2012). Schizomids
can be found in leaf litter, under rocks, in caves and
other subterranean voids, in many tropical and sub-
tropical regions of the world. Some species have been
recorded from greenhouses and other artificial envi-
ronments in Europe, demonstrating their ability to
withstand long-distance transport by human com-
merce (Harvey 2003). Three species have previously
been recorded from greenhouses in Europe. Scbi-
zomus crassicaudatus (O. P.-Cambridge, 1872) was
introduced from Sri-Lanka into France. Zomus bag-
nallii (Jackson, 1908) was introduced from South-
east Asia, the Seychelles or Mauritius into Great
Britain (Harvey 2003, Blick 2006). Stenochrus por-
toricensis Chamberlin, 1922 has been introduced into
Great Britain, Spain (Canary Islands) and the Czech
Republic (Harvey 2003, Blick 2006, Korenko et al.
2009). All of the specimens from Europe were col-
lected from heated greenhouses. Undetermined schi-
zomids have been collected from two greenhouses in
Germany (Cokendolpher et al. 2006).

Material and methods

The schizomids were found in two heated green-
houses (temperature about 26 °C) situated in the
same part of the Botanical Garden of the Comen-

Jana CHRISTOPHORYOVÄ, Anna SESTÄKOVÄ, Miroslav KRUMPÄL, Peter
FENDA, Department of Zoology, Faculty of Natural Sciences, Comenius
University, Mlynska dolina B-1, SK-84215 Bratislava, Slovak Republic, e-
mails: christophoryova@gmail.com, asestakova@gmail.com, krumpal@
fns.uniba.sk, fenda@fns.uniba.sk

submitted 4. 12.2012, accepted 22.3.2013, online 13.5.2013

ius University in Bratislava (Fig. lA), Slovakia (grid
reference number of the Databank of Slovak Fauna
7868; 48°08’49.2”N; 17°04’20.97”E; 148 m a.s.L;
Igt. J. Christophoryova, P. Fenda, K. Krajcovicova
&c A. Sestakova). Thirteen juvenile specimens were
extracted from soil samples using Tullgren funnels
(9 May 2012: 3 juveniles, 7 June 2012: 2 juveniles,
22 November 2012: 8 juveniles). Seven females were
individually collected (22 November 2012) under
the stones (Fig. IB) and two were extracted from soil
samples (22 November 2012).

Two juveniles and five females were dissected,
studied as permanent slide mounts, and photo-
graphed using a Feica DM1000 stereoscopic mi-
croscope with an ICC50 Camera Module (FAS EZ
application 1.8.0). Other specimens were preserved
in 80 % or 96 % ethanol and were photographed
using a digital camera CANON EOS HOOD con-
nected to a Zeiss Stemi 2000-C stereomicroscope.
The female genitalia were dissected in 96 % etha-
nol and macerated using NaOH solution, after
which they were mounted on a permanent slide in
Swan’s fluid. Digital images were montaged using
the “CombineZP” image stacking software. The
specimens were identified by J. Christophoryova
and M. Krumpal. The material is deposited in the
collection of the first author in the Comenius Uni-
versity, Bratislava.

Results and discussion
Main morphological features

The adult female of Stenochrus portoricensis (Fig. 2A)
is characterised by the following morphological fea-
tures (Rowland & Reddell 1980, Reddell & Coken-
dolpher 1995,Tourinho 6cKury 1999, Armas 2010):
propeltidium with two apical setae on an anterior
process arranged one behind the other and two pairs

26

J. Christophoryovä, A. Sestäkovo, M. Krumpol & P. Fend'o

Fig. 1 : Heated greenhouses in the Botanical Garden of the Comenius University in which Stenochrus portoricensis occurs. A. Interior
view of greenhouse. B. Detail of the stones under which females were collected. - Photos; Jana Christophoryovä

of dorsal setae (Fig. 3A); corneate eyes completely
absent; metapeltidium entire; movable cheliceral fin-
ger without accessory teeth (Fig. 3B), guard tooth
present at end of serrula; short mesal spur present on
pedipalpal trochanter (Fig. 3C); body without cla-
vate setae; abdominal tergite II with two posterior
setae; flagellum short, with three segments (Fig. 3D);
spermathecae (Fig. 3E) with two pairs of asymmetric
lobes, the laterals clearly shorter than the medians,
both lobes with a wrinkled wall covered with small
bumps, the median lobes heavily sclerotized and vis-
ible without dissection through the genital sternite,
gonopod short.

At first, only juvenile specimens were collected
during our research in the Botanical Garden of the
Comenius University. Their identification was dif-
ficult because the majority of recent schizomid de-
scriptions are based on the characters of adults. The
juveniles were smaller and markedly less sclerotized
(Fig. 2B) and were characterized by features typical
for the genus (Fig. 4). The identification of the spe-
cies was equivocal prior to finding the females.

Fig. 2: Stenochrus portoricensisfrom Slovakia. A. Female. B. Juve-
nile. Scales: 1 mm. - Photos: Anna Sestakova

Stenochrus portoricensis new to Slovakia

27

Fig. 3: Morphological

characters of Stenochrus
portoricensis female (mi-
croscope slides). - Photos:
Jana Christophoryova

A. Detail of anterior part
of propeltidium (dorsal
view). Arrows point to
apical propeltidial setae.

B. Chelicera (lateral view).
Scales: 0.2 mm (A, B)

C. Pedipalp with detail
of trochanter (prolateral
view). Arrow points to
mesal spur on trochanter.
Scales: 0.5 mm (C), 0.1
mm (C detail)

D. Flagellum (dorsal view).

E. Spermathecae (dor-
sal view). Scales: 0.1 mm
(D, E)

Natural history

Stenochrus portoricensis naturally occurs in Mexico
and the Caribbean, but has been introduced to
many other countries (e.g. the USA, Rio de Janeiro
in Brazil, Colombia, Ecuador, the Canary Islands,
Great Britain and the Czech Republic) (Reddell &
Cokendolpher 1995, Tourinho & Kury 1999, Har-
vey 2003, Korenko et al. 2009, Kury et al. 2010).
The majority of the males are known from southern
Mexico, but some have been found on Caribbean
Islands (Tourinho &Kury 1999). The species is fac-

ultatively parthenogenetic (Martin 5c Oromi 1984)
and only females and juveniles have been found in
heated greenhouses (Cloudsley-Thompson 1949,
Korenko et al. 2009). Parthenogenesis in this species
probably facilitates its importation. This species has
been reported from caves, under rocks, in leaf litter
and soil, in both synanthropic and disturbed habi-
tats (Rowland 5c Reddell 1980, Martin 5c Oromi
1984, Reddell 5c Cokendolpher 1995, Tourinho 5c
Kury 1999, Santos et al. 2008, Armas 2010). Some
specimens have been collected in association with

28

J. Christophoryovä, A. Sestakova, M. Krumpa! & P. Fendb

Fig. 4: Morphological

characters of Stenochrus
porfor/cens/s juvenile (mi-
croscope slides). - Photos;
Jana Christophoryovä

A. Pedipalp with detail
of trochanter (prolateral
view). Arrow points to the
mesal spur on trochanter.
Scales: 0.2 mm (A), 0.1
mm (A detail)

B. Leg I (lateral view). Ar-
row points to one of the
trichobothria on tibia.

C. Chelicera (lateral view).
Scales: 0.5 mm (B), 0.2
mm (C)

D. Detail of anterior part
of propeltidium (dorsola-
teral view). Arrows point
to apical propeltidial se-
tae. E. Flagellum (dorsal
view). Scales; 0.1 mm
(D, E)

ants and termites (Martin &c Oromi 1984, Red-
dell &. Cokendolpher 1995, Santos et al. 2008). In
Tenerife, only females are known from volcanic pit
at low altitude (Martin & Oromi 1984, Oromi Sc
Martin 1992). In the Czech Republic, the species
was found under pieces of bark lying on wet soil in
a greenhouse in the Botanical Garden of Masaryk
University, Brno (Korenko et al. 2009).

The records provided in the current study are
consistent with known natural history patterns.
Stenochrus portoricensis occurs in Slovakia in heated

greenhouses, with a temperature of about 26 °C,
in the soil and under stones. There have not been
importations of plants from tropical areas or other
botanical gardens for several years and some insec-
ticides have been regularly applied one to two times
per month (Vertimex, Confictor, Omite). According
to these facts, the population of S. portoricensis can
probably survive in the greenhouses. Only females
and juveniles were collected, which implies that the
Comenius University population of this species is
probably parthenogenetic.

Stenochrus portoricensis new to Slovakia

29

Acknowledgements

The authors would like to thank the director of the Bo-
tanical Garden of the Comenius University, Dr. Jaroslav
Bella, and his staff for permission to collect the schizomids.
We would also like to express our thanks to Alica Christo-
phoryova for technical assistance with the figures, Katarina
Krajcovicova for help with collecting schizomids and Viera
Janoskova for translating Spanish papers. The advice and
help of our colleagues Stanislav Korenko and Mark Har-
vey are also appreciated.

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405: 1-6

A ra c h n o I ogische Mitteilungen 45: 30-3 5

Karlsruhe, Juni 2013

Do really all wolf spiders carry spiderlings on their opisthosomas? The case of
Hygrolycosa rubrofasciata (Araneae: Lycosidae)

Petr Dolejs

doi: 1 0.5431 /aramit4507

Abstract. Wolf spider females are characterised by carrying cocoons attached to their spinnerets. Emerged spi-
derlings are carried on the females' opisthosomas, with the exception of three Japanese lycosid species who car-
ry spiderlings on empty cocoons. Here, the same behaviour is recorded in a European spider: the drumming wolf
spider Hygrolycosa rubrofasciata. Spiderlings of this species do not try to climb on the female's opisthosoma, even
when they are adopted by a female of a species with a normal pulli-carrying behaviour. This behaviour occurs in
Trechaleidae and four unrelated species of Lycosidae inhabiting wet habitats and is therefore regarded as an adap-
tation to the unsuitable environment.

Keywords: Cocoons, female abdominal knobbed hairs, humid habitats, pulli-carrying behaviour, spiderling clus-
ters

Female wolf spiders are known for their care of both
cocoons and spiderlings (Foelix 2011). They carry
their cocoons attached onto the spinnerets (cocoon-
carrying behaviour) and their spiderlings on the
opisthosoma (pulli-carrying behaviour) (Fujii 1976).
All lycosids show cocoon-carrying behaviour, there
are, however, three exceptions concerning pulli-
carrying. Fujii (1976) found that females of Arctosa
ebicha Yaginuma, 1960 and Arctosa fujiii Tanaka,
1985 (sub Lycosa sp.) do not carry their spiderlings
on the opisthosoma. Juveniles of these species stay
for several hours on the cocoon surface and then
disperse within one to four days (Fujii 1976). This
behaviour is also typical for members of the puta-
tive sister family Trechaleidae, which also carry co-
coons attached to the spinnerets like lycosids (Carico
1993). The same behaviour was also recorded for fe-
males of Hygrolycosa umidicolaT2iCi2k2i., 1978 (Suwa in
litt. 1977, Yaginuma 1991). Females of both ebicha
A. fujiii lack the abdominal knobbed hairs (Fujii
1983) that enable the spiderlings to climb onto the
mother’s opisthosoma (Rover et al. 1973). Kronest-
edt (1984, 1996) recorded that these knobbed hairs
are also absent in females of Hygrolycosa rubrofasciata
(Ohlert, 1865) and suggested that this may be associ-
ated with the pulli-carrying behaviour. Ahtiainen et
al. (2002) noted that “the offspring (of H. rubrofas-

Petr DOLEJS, Department of Zoology, National Museum - Natural His-
tory Museum, Cirkusova 1 740, CZ - 1 92 00 Praha 9 - Horni Pocernice,
Czech Republic; e-mail: petr_dolejs@nm.cz. Department of Zoology,
Charles University in Prague, Faculty of Science, Vinicna 7, CZ - 1 28 44
Praha 2, Czech Republic; e-mail: petr.dolejs@natur.cuni.cz

submitted 10.2.2013, accepted 1 1.4.2013, online 13.5.2013

data) usually remain on the female’s abdomen or on
top of the empty egg sac for a day to chitinise their
exoskeleton, after which they disperse”. Thus, this
species was chosen for the present study to clarify its
pulli-carrying behaviour.

Hygrolycosa Dahl, 1908 is still of uncertain sub-
familial affinities. It belongs either to Piratinae (Zy-
uzin 1993) or Venoniinae (Murphy et al. 2006), and
currently contains four species (Platnick 2013). The
drumming wolf spider, Hygrolycosa rubrofasciata is a
rare, Palaearctic, Eurosiberian wolf spider inhabiting
reed swamps, alder forests and marshy pine forests
(Buchar & Ruzicka 2002). It is a diurnal species
(Kronestedt 1984, 1996), well known for its audible
acoustic courtship behaviour (Helversen in: Rovner
1975, Kronestedt 1984, 1996, Köhler & Tembrock
1987) and for the lowest male diploid chromosome
number (2nc5 = 20) among European wolf spiders
(Gorlov et al. 1995). Both males and females ma-
ture in autumn (Kronestedt 1984) and it takes two or
three years to complete their life cycle (Kotiaho et al.
1996, Vertainen et al. 2000). The life cycle is a transi-
tion between diplochronous - with mating periods in
autumn and spring - and stenochronous with a mat-
ing period occurring only in spring (Braun 1976 and
references therein). Males die during or immediately
after the mating season (in spring); however, females
can survive until the next mating season (Kotiaho et
al. 1999). In northern Europe, females produce more
than one cocoon (Vertainen et al. 2000), and in cen-
tral Europe, females can make up to three cocoons
per year (Dolejs pers. obs.). The cocoons contain
about 60 eggs (Wiebes 1959). The pulli-carrying be-
haviour of this species has, however, not been thor-

Spiderlings on the opisthosomos? Not in Hygrolycosa rubrofasciata

31

oughly investigated previously. Therefore, this study
aims to investigate (1) the pulli-carrying behaviour
of H. rubrofasciata^ (2) the behaviour of H. ubrofas-
ciata spiderlings on “substitute” mothers and “adop-
tive” spiderlings on H. rubrofasciata mothers after
exchange of cocoons, and (3) the surface of the H.
rubrofasciata cocoon in an attempt to determine the
possible responsible structures for its pulli-carrying
behaviour.

Methods

Females were collected at their first cocoon-carrying
stage in the Kokofinsko Protected Landscape Area
during a field course “Fauna of the Czech Republic”.
There were six females collected at the locality Cerny
dul Natural Monument (50°29’N, 14°37’E; grid
mapping square 5553; 310 m a.s.l.) on 10.5.2011, six
females were collected at the locality Mokfady Horni
Libechovky Nature Reserve (50°31’N, 14°32’E; grid
mapping square 5453; 260 m a.s.l.) on 11.5.2011, and
two females were collected at the locality Jestfebske
slatiny (50°36’N, 14°37’E; grid mapping square 5453;
250 m a.s.l.) on 23.5.2011 (Igt. Antonin Kürka).
Voucher specimens are deposited in the National
Museum in Prague (N-s P6A 5070-5072).

Pulli-carrying behaviour was studied in the labo-
ratory. In order to study differences in behaviour in
test tubes and in simulated natural conditions, seven
females were held in plastic test tubes (length 10 cm,
diameter 15 mm) with cotton wool as a source of
water and seven females were kept in four diagonally
divided glass terraria (14x11x8 cm) with 3 cm of
leaves. The programme NCSS 2007 (Hintze 2006)
was used to test normality, to calculate descriptive
statistics on the length of the pulli-carrying period,
and to compare (Two- Sample T-Test) the length of
the pulli-carrying period of females kept in test tubes
with those kept in terraria.

After the females of H. rubrofasciata made the sec-
ond cocoon, the behaviour of females with exchanged
cocoons were tested. Cocoons of three females (in
terraria) were replaced by cocoons of Pardosa amen-
tata (Clerck, 1757), a species that displays a normal
pulli-carrying behaviour (Vlijm et al. 1963). Three
females of the latter species were collected at their
second cocoon-carrying stage from Zleby (49°53’N,
15°29’E; grid mapping square 6158; 240 m a.s.l.).
They were kept in terraria as described above for H.
rubrofasciata. After the spiderlings of both species
emerged from the cocoons, the behaviour of spider-

lings on their “adoptive” mothers was examined and
documented (digital camera Olympus C-7070 WZ).
In order to compare possible structural differences
in cocoon surface, empty cocoons of both species
were inspected under a scanning electron microscope
JEOL 6380 LV.

Results

Pulli-carrying behaviour of Hygrolycosa
rubrofasciata

After emergence from the cocoon, spiderlings of H.
rubrofasciata did not climb onto the opisthosoma of
their mother. Instead, they stayed on the surface of
the cocoon (Fig. 1). Spiderlings of females kept both
in terraria and test tubes behaved in the same way.
However, time spent on the cocoon surface differed
significantly in both groups (p = 0.0031%). Spider-
lings of females kept in terraria dispersed within 1-3
days (0 = 2.29, SD = 0.756, n = 7), but those of fe-
males kept in test tubes within 4-6 days (0 = 5.00,
SD = 0.816, n = 7). In cases of larger clutch size, a
few spiderlings stepped on the mother’s opisthosoma
using their forelegs, however they always remained in
contact with the cocoon or with lower layer of spid-
erlings on the cocoon surface. Females detached their
empty cocoons after the spiderlings had dispersed.
Behaviour of spiderlings on substitute females
All females (of both H. rubrofasciata and P amentata)
accepted, sooner or later, cocoons of the other species.
After emergence, spiderlings of H. rubrofasciata did
not try to climb on the opisthosomas of P amentata
females and remained on the cocoon surface (Fig. 2).
By contrast, spiderlings of P amentata tried to climb

Fig. 1: Pulli-carrying behaviour of Hygrolycosa rubrofasciata.
Newly emerged spiderlings occupy the surface of the cocoon
instead of climbing onto the opisthosoma of their mother.

32

P. Dolejs

onto the opisthosoma of H. rubrofasciata females, yet
only few spiderlings were successful in settling on a
substitute mother. The majority of spiderlings formed
a cluster either beside the empty cocoon or on the
ventral surface of the female’s opisthosoma - in the
area where the cocoon was touching the opisthosoma.
Therefore, H. rubrofasciata females were carrying two
pellets’: the empty cocoon and the cluster of P amen-
tata spiderlings (Fig. 3). However, the females lost
the clusters within the same day as the spiderlings
emerged and the spiderlings dispersed the following
day. Examination of the cocoon surfaces revealed no
differences between the cocoon structures of both
species (Figs 4, 5).

Fig. 2: Behaviour of Hygrolycosa rubrofasciata spiderlings on a
substitute Pardosa amentata mother. The spiderlings remained
on the cocoon surface and did not try to climb onto the female's
opisthosoma.

Discussion

A generally accepted statement that females of all
wolf spiders carry spiderlings on their opisthosomas
was contested. Fujii (1976) was the first author to
report “abnormal” behaviour in two Arctosa species
which carried spiderlings on their cocoons instead of
on their opisthosomas. Suwa (in litt. 1977) and Yagi-
numa (1991) reported similar behaviour for Hygroly-
cosa umidicola, and Kronestedt (1984) and Ahtiainen
et al. (2002) suggested such a possibility in Hygro-
lycosa rubrofasciata too. In the present study, it was
confirmed unequivocally that females of H. rubrofas-
ciata do not carry spiderlings on their opisthosomas,
but instead on their empty cocoons. Until now, this

Fig. 3: Behaviour of Pardosa amenfafa spiderlings on substitute
Hygrolycosa rubrofasciata mother. Most of the spiderlings were
not successful in settling on the opisthosoma of a substitute fe-
male. Instead, they formed a cluster beside the empty cocoon.
The female was carrying both the spiderling cluster and the em-
pty cocoon for a while, until she lost the cluster.

unusual behaviour was only known from four lycosid
species: A. ebicha^ A. fujiii, H. rubrofasciata and H.
umidicola.

The time which the spiderlings spent on the co-
coon surface differed between females kept in terraria
and those in the test tubes. In natural conditions, the
spiderlings probably disperse within two days - this
is maybe the reason why pulli-carrying behaviour has
not yet been observed in the field. In the test tubes,
spiderlings probably did not have enough space to
disperse and therefore remained longer on the co-
coon surface. A similarly short pulli-carrying period
was reported {or A. fuj Hi ^ but three times longer for
A. ebicha (Fujii 1976). However, the females of these
species were kept in small glass vials and therefore
the length of the period could be affected. There are
no data about the length of pulli-carrying period in
H. umidicola.

After cocoon exchange, spiderlings behaved on
the substitute mother as they would on their own
mother: H. rubrofasciata spiderlings remained on
the cocoon surface, although they were carried by
a Pardosa female. Similarly, Pardosa spiderling tried
to climb on the H. rubrofasciata female, but only
few of them were successful — presumably due to
the absence of abdominal knobbed hairs. Tke ma-
jority of spiderlings tended to aggregate elsewhere.
It seems therefore that the primary behaviour of the

Spiderlings on the opisthosomas? Not in Hygrolycosa rubrofosciata

33

X100 100Mm

15kU

X100 100Mm

Figs 4a-c: Cocoon surface of Hygrolycosa rubrofosciata. Scale
bars: a) and b) 1 00 |jnn, c) 50 urn

Figs 5a-c: Cocoon surface of Pardosa amentata. Scale bars: a)
and b) 1 00 pm, c) 50 pm

spiderlings is species-specific and does not depend
on the mother. The same results were also obtained
by Fujii (1980). He tested behaviour of Pardosa as-
trigera L. Koch, 1878 which carry spiderlings nor-
mally on the opisthosoma, 2ccvd A. fiijiii (sub Arctosa
sp.), which carry spiderlings on the cocoon (see Fu-

jii 1980: figs. IG and IH). Both Pardosa spiderlings
failed to settle on the substitute mother {A.fujin, H.
rubrofasciata) and formed a cluster. This behaviour
disagrees with observations in Schizocosa crassipes
(Walckenaer, 1837) and Rabidosa rabida (Walck-
enaer, 1837) in which the spiderlings never clus-

34

P. Dolejs

tered when separated from their mothers (Higashi
&c Rovner 1975).

Spiderlings settle only on suitable surfaces (Engel-
hardt 1964, Rovner et al. 1973, Fuji! 1983). Therefore,
it is curious that there are no proper structures’ on
the cocoon surface of H. rubrofasciata. Unfortunately,
only a few workers examined cocoon microstructure
(e.g. De Bakker et al. 2006), but none of them consid-
ered lycosid and trechaleid cocoons. For this reason, it
would be of value to study the mechanics underlying
lycosid and trechaleid spiderling behaviour.

The clustering of spiderlings on an empty co-
coon for a short time - instead of on their mother’s
opisthosoma — in A. ebicha and A.fujiii is peculiar be-
cause at least A. ebicha is related neither to the genus
Hygrolycosa nor to other Arctosa species (Murphy et
al. 2006), and probably belongs (together with the re-
lated Arctosa kwangreungensis Paik & Tanaka, 1986)
to a separate, undescribed genus (Framenau in litt.
5. xii. 2007). Interestingly, members of Hygrolycosa,
both A. ebicha and A. fujiii, and the most members
of the family Trechaleidae live in very humid or even
wet habitats (Tanaka 1978, 1991, Carico 2005). Fit-
tie drops of water remaining on the opisthosomal
surface and hairs probably create unsuitable condi-
tions for clustering the spiderlings. The dragline
threads eventually forming a layer over the surface
and providing a good means of attachment for the
spiderlings (Rovner et al. 1973) probably cannot be
attached to the moist surface or hairs either. Mois-
ture can even paste the knobbed hairs together and
disable the spiderlings’ ability to cling onto their
mother’s opisthosoma.

For this reason, the cocoon surface seems to be
more water-repellent (Hieber 1984 in: Hieber 1992)
and presents better place to cluster, despite the ab-
sence of any auxiliary structures. Fujii (1983) reported
the absence of knobbed hairs, but normal pulli-car-
rying behaviour, m Arctosa subamylacea (Bösenberg &
Strand, 1906) and Arctosa depectinata (Bösenberg &
Strand, 1906). These two species live mainly in fields
(Tanaka 1991) so clustering on the cocoon has not
developed. On the other hand, Fujii (1980), who also
tested spiderling behaviour after cocoon exchange in
Pardosa astrigera and Pirata subpiraticus (Bösenberg 6c
Strand, 1906), showed that spiderlings of P astrigera
climbed indeed onto the opisthosoma of wet-habi-
tat-dwelling P subpiraticus but all of them dropped
from her till the next day and clustered elsewhere.
Correspondingly, spiderlings of most of Pirata (and

Piratuld) species do not mount the opisthosoma of
the female, but remain in the silken retreat (Nielsen
1932). Therefore, living in wet habitats rather than
lacking knobbed hairs seems to be the main reason
for modifying pulli-carrying behaviour.

This contribution shows how little is known
about basic biological data for wolf spiders. It is nec-
essary to study the pulli-carrying behaviour and fe-
male abdominal knobbed hairs of other wet-habitat
lycosids {Hygrolycosa, Arctosa, Pirata, Piratula) and
trechaleids. As Fujii (1976) already stated, more data
must be obtained to clarify the evolution of pulli-
carrying behaviour in lycosids and related families.

Acknowledgements

My great thanks belong to Hozumi Tanaka who informed
me about the behaviour of H. umidicola, provided me with
unpublished information from Masayoshi Suwa, and en-
couraged me to study H. rubrofasciata. I would like to thank
the participants of the field course “Fauna of the Czech Re-
public”, namely Lucie Jarosova, who collected females of H.
rubrofasciata in Kokofinsko PLA, and to Antonin Kurka who
collected them at Jestfebske slatiny. I wish to thank Volker
W. Framenau for providing me with his unpublished data. I
am indebted to Jan Buchar and Hozumi Tanaka for valuable
discussion and critical reading of the manuscript. I also thank
Torbjörn Kronestedt and an anonymous reviewer for impor-
tant comments and valuable suggestions that improved the
earlier version of the manuscript. This work was financially
supported by Ministry of Culture of the Czech Republic
(DKRVO 2013/13, National Museum, 00023272).

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Arachnologische Mitteilungen 45: 36-39

Karlsruhejuni 2013

Bestätigung von Homalenotus quadridentatus (Opiliones: Sclerosomatidae)
für die Fauna Deutschlands

Antje Deepen-Wieczorek & Axel L. Schönhofer

doi: 1 0.5431 /aramit4508

Abstract. Confirmation of Homalenotus quadridentatus (Opiliones: Sclerosomatidae) for the fauna of Ger-
many. The sclerosomatid harvestman Homalenotus quadridentatus (Cuvier, 1795), long known close to the border
of Germany, is hereby confirmed for the country. Several specimens were observed and collected in a small urban
garden area in the town Aachen, suggesting a considerable population. The finding is briefly discussed, and general
remarks on the distribution, ecology and conservation of the species are provided.

Keywords: Central Europe, first record, Germany, harvestmen, oceanic climate

Bereits in der Bearbeitung der mitteleuropäischen
Fauna wies Martens (1978) auf die grenznahen Nach-
weise des in Westeuropa weit verbreiteten Scleroso-
matiden Homalenotus quadridentatus (Cuvier, 1795)
und damit auf die Wahrscheinlichkeit seines Vorkom-
mens in Deutschland hin. Stabile Vorkommen waren
zu diesem Zeitpunkt aus Süd-Limburg (Niederlan-
de) bekannt (Spoek 1957, 1975), mit einigen Fund-
punkten nur 5 km von der niederländisch-deutschen
Grenze entfernt. In weiser Voraussicht wurde bereits
eine deutsche Nachweiskarte für Homale?totus quad-
ridentatus vorbereitet (Staudt 2013) und tatsächlich
wurde die Art bereits von Petto (1991) aus dem Itter-
bachtal südöstlich Aachens gemeldet, der Nachweis
Mangels prüfbarer Belege allerdings nicht in deut-
sche Faunenlisten übernommen (T. Blick in litt.). Im
November 2012 gelangen nun eindeutige Nachweise
von H. quadridentatus (Abb. 1) auf deutschem Gebiet,
in einem Aachener Garten. Sie werden nachfolgend
dokumentiert und diskutiert.

Fundort, Material und Methoden

Der Fundort aller Tiere ist ein privater Garten:
Deutschland, Nordrhein-Westfalen, Aachen, Brand,
Niederforstbach, Münsterstr. 207, 248 m ü. N.N.
(WGS84: 50,7395°N, 5,1583°E, TK25: 5202). Im
Zeitraum vom 9. November bis Ende November
2012 wurden täglich ein bis zwei Exemplare durch
manuelle Suche nachgewiesen, am 9. und 28.11.
sogar vier Exemplare (Fotobelege mit Panasonic

Antje DEEPEN-WIECZOREK, Münsterstr. 207, 52076 Aachen, Deutsch-
land. E-Mail: antje.wieczorek@online.de

Axel L. SCHÖNHOFER, Institut für Zoologie, Abteilung Evolutionsbiolo-
gie, Joh.-v.-Müller-Weg 6, 55128 Mainz, Deutschland.

E-Mail: axel.schoenhofer@gmx.net

eingereicht 4.2.201 3, angenommen 12.4.2013, online 13.5.2013

Lumix G2 mit Leica DG Macro-Elmarit 1:2.8/45
asph.vom 9., 12., 14., 18., 20. und 30.11). Von diesen
wurden zwei Männchen am 20.11.2012 als Belege
in 70 % EtOH konserviert und in der Sammlung
Schönhofer unter der Nummer 1534 archiviert
(Abb. lA). Zwei Exemplare wurden vom 18. bis
28.11. in mit Erde, Steinen und Rindenstückchen
ausgelegten Petrischalen beobachtet. Die Binokula-
raufnahme wurde mit einem Wild-Heerbrugg M5A
und einer Nikon D90 (Weitwinkelobjektiv mit Ret-
roadapter) erstellt. Die Determination erfolgte nach
Martens (1978) und Wijnhoven (2009), siehe dort
weitere Informationen zur Spezies.

Geografische Einordnung des Fundortes und
Anmerkungen zum Areal

Nach Durchsicht der relevanten Literatur bleibt,
bis auf die neuen Nachweise aus den Niederlanden
(Noordijk & Wijnhoven 2009, Wijnhoven 2009)
und Deutschland, das von Martens (1978: Abb. 747)
grob umrissene Verbreitungsgebiet aktuell (Abb. 2).
Probleme ergeben sich aber mit der Zuordnung von
Homalenotus-N3.chweisen am südlichen Arealrand.
So wurde H. quadridentatus auch für die Südost- und
Südwestküste der Iberischen Halbinsel gemeldet
(Rambla 1960, 1967) - Angaben, die bereits Martens
(1978) nicht übernahm. Es sei hier auf die Schwie-
rigkeiten bei der Abgrenzung der südeuropäischen
Homalenotus- Asien hingewiesen, die eine Zuord-
nung von Literaturnachweisen ohne Sichtung des
Materials nicht immer zweifelsfrei erlaubt. Nördlich
der Pyrenäen scheint H. quadridentatus hingegen die
einzige Art der Gattung zu sein.

Nicht abschließend geklärt scheint auch der ge-
naue Verlauf der östlichen Arealgrenze von Homale-
notus quadridentatus^ die sich im Süden Frankreichs

Homalenotus quadridentatus in Deutschland

37

Abb. 1: Homalenotus quadridentatus. A. Männchen von dorsal; B-D. Adulte Exemplare unbestimmten Geschlechts; E. Subadultes
Exemplar.

Fig.1 : Homalenotus quadridentatus. A. Male, dorsal view; B-D. Adult specimen of unknown gender; E. Subadult specimen.

etwa entlang der Grenze zu den Tulpen hinzieht (Les-
sert 1917: Dep. Haute-Savoie: Gaillard, Petit-Saleve;
Delfosse 2004: Dep. Isere: Le Sappey, Dep. Alpes-
de-Haute-Provence: Digne-les-Bains) und in der
Schweiz ab dem Genfer See am Nordalpenrand bis
knapp östlich Luzern (Lessert 1917: Weggis; Becker
1896: Vallee de la Reuss) Vordringen soll. Martens
(1978) bezieht diese östlichsten Schweizer Funde als
wahrscheinlich zu unsicher, nicht in seine Verbrei-
tungskarte mit ein. Nördlich davon lässt sich keine
genaue Ostgrenze ziehen. Es scheint jedoch bisher
keine Funde in den Vogesen oder östlich davon zu
geben. Eine Einwanderung über eine Südroute nach
Deutschland ist wegen der dazwischen liegenden
Gebirgszüge mit für Homalenotus ungünstigerem
Klima weniger wahrscheinlich. Nennungen des süd-
westeuropäischen Homalenotus monoceros C.L. Koch,
1839. aus Süddeutschland (Hoc 1966: Regensburg)
beruhen sicher auf Verwechslungen.

Betrachtet man die weitere Verbreitung der Art,
so fügt sich der Fundort bei Aachen fast nahtlos an
die bisher bekannte Verbreitung und vor allem an die
schon länger bekannten Vorkommen in Limburg an.

Mit dem Nachweis von Ho7nale7totus quadridentatus
steigt die Zahl der in Deutschland im Freiland vor-
kommenden Weberknechtarten auf 52 (Muster et al.
im Druck).

Ökologie

Der Fundort zeichnet sich durch dichte Kraut-
schicht, Grasbewuchs durchsetzt mit Stauden wie
z.B. Goldrute aus und liegt im Bereich eines mit
Efeu überwachsenen Wiesen-Weidenzauns. Ein
künstlich angelegter Teich sorgt für höhere Boden-
feuchtigkeit. Die Tiere wurden hier vorwiegend un-
ter flachen Steinen aufgefunden, und sind aufgrund
ihrer Tarnung mit Erde oft schwer zu entdecken. Als
Begleitarten wurden im Jahre 2012 beobachtet: Opi-
lio canestrinii (Thoreil, 1876) (23.12.-24.12.: 1 juv.;
2.11.: 7 Ex.), Paroligolophus agrestis (Meade, 1855)
(21. und 29.11.: 499), Nemastoma dentigerum Canes-
trini, 1873 (22.12.-23.12.: 6 ?id.)^Rilaena triangularis
(Herbst, 1799) (14. und 16.11.: 12 juv.). Die weitere
Umgebung ist vorwiegend landwirtschaftlich ge-
nutzt, in der Nähe beflndet sich ein aufgelassener
und mit Wald bestandener Kalksteinbruch, was in

38

A. Deepen-Wieczorek & A. L Schönhofer

Abb. 2: Verbreitung von Homalenotus quadridentatus basierend auf Martens (1978) und modifiziert nach Delfosse (2004), Noordijk&
Wijnhoven (2009), Vanhercke (2004), Wijnhoven (2009) und dem hier publiziertem Nachweis. Der Pfeil zeigt den Fundort in Deutsch-
land. Fragezeichen zeigen unsichere Schweizer Fundorte nach Becker (1896) und Lessert (1917) und die nicht geklärte Verbreitungs-
grenze in Frankreich. Abkürzungen nach Internationalem Ländercode.

Fig. 2: Distribution of Homalenotus quadridentatus based on Martens (1978) and modified according to Delfosse (2004), Noordijk&
Wijnhoven (2009), Vanhercke (2004), Wijnhoven (2009) and the record herein. The arrow indicates the locality in Germany. Question
marks indicate problematic Swiss records according to Becker (1896) and Lessert (1917) and the unverified distribution border in
France. Abbreviations follow international country code.

etwa den Fundorten in Limburg entspricht. Noor-
dijk & Wijnhoven (2009) fanden die Art mehrfach
in Ackerrandstreifen (Seeland, Niederlande). Petto
(1991) meldete 24 Individuen gefangen mit Boden-
fallen in Gehölzstrukturen des Itterbachtals.

Die regelmäßig erbrachten Nachtveise von Ho-
malenotus aus einem kleinflächigen urban-anthropo-
gen geprägten Lebensraum, sowie die durch unsere
Funde wahrscheinlich zu bestätigenden Nachweise
von Petto (1991, ebenfalls TK 5202), lassen auf eine
relativ stabile Population im Raum Aachen schlie-
ßen. Trotz des lokalen Nachweises halten wir eine
Gefährdung daher für unwahrscheinlich.

In Gefangenschaft nahm Homalenotus quadri-
dentatus verschiedene Wirbellose (Annelida, Acari,
Isopoda, Diptera, Collembola, Diplopoda, vorwie-
gend frisch getötet) als Nahrung an. Gelegentlich
wurde ein Abbürsten abgestorbener Blätter mit den

Palpen beobachtet. Bei Kontaktaufnahme betasteten
sich die Tiere mit den Vorderbeinen. In Ruhephasen
saßen sie beieinander, hielten aber Individualabstand
ohne Kontakt. Teilweise wurde kräftiges Häkeln mit
den Beinen oder Wegschieben eines anderen Tieres
beobachtet.

Ausbreitung und Ausblick

Die von Noordijk 8c Wijnhoven (2009) und Wijnho-
ven (2009) berichteten, rezenten Funde von Homale-
notus quadridentatus verschieben die vormals bekannte
Verbreitungsgrenze der Art innerhalb der Niederlan-
de (Süd-Limburg) scheinbar deutlich nach Norden.
Zusammen mit den Funden in Aachen stellt sich die
Frage, ob auch dieser Weberknecht sein Areal aktuell
erweitert. Arten für die eine deutliche Expansion an-
zunehmen ist, sind beispielsweise Nemastoma dentige-
rum^ Odiellus spinosus (Bose, 17 92), Astrobunus laevipes

Homalenotus quadridentatus in Deutschlond

39

(Canestrini, 1872) und Nelima sempronii Szalay, 1951.
Diese Arten des kontinentalen oder mediterranen
Verbreitungstyps besiedelten in Deutschland zuerst
lokale, kontinental geprägte Wärmeinseln (Stand
Martens 1978) und breiteten sich in den letzten Jahr-
zehnten, wahrscheinlich im Zuge der Erwärmung,
sehr deutlich aus (vgl. Staudt 2013).

Homalenotus quadridentatus hingegen besiedelt
den ozeanisch geprägten Raum, der durch relativ
kühle Sommer, milde Winter und ganzjährigen Nie-
derschlag charakterisiert ist (vgl. Heinze & Schrei-
ber 1984). Die Jahresdurchschnittstemperatur in
Aachen-Brand liegt bei 10 °C, die minimale Tages-
durchschnittstemperatur bei -12,2 °C, und ist damit
im deutschen Vergleich sehr gemäßigt (siehe http://
www.wetter-aachen-brand.de). Auch der am Fund-
ort nachgewiesene Paroligolophus agrestis besitzt ein
mit dem atlantischen Klimaeinfluss korrespondie-
rendes Areal, dringt allerdings deutlich weiter in
den kontinentalen Bereich als Homalenotus (Martens
1978). Im Vergleich mit den kontinental-mediterra-
nen Arten ist daher nicht von einer starken rezen-
ten Ausbreitung von Homalenotus auszugehen. Auch
die neuen Funde schließen wahrscheinlich lediglich
bestehende Nachweislücken, die weitere Funde ent-
lang der niederländischen Grenze (Noordijk & Wi-
jnhoven 2009, Wijnhoven 2009) und in Fuxemburg
(Vanhercke 2004) möglich erscheinen lassen. Auch
das unveränderte Homalenotus- in England
(British arachnological Society 2013) spricht nicht
für eine rezente Ausbreitung der Art.

Danksagung

Die gemeinschaftliche Bestimmung oder Bestätigung der
Weberknechte erfolgte auf http://insektenfotos.de/forum
zusammen mit Katrin Jäckel, Arp Kruithof und Jörg Pageler,
wofür wir uns herzlich bedanken. Theo Blick ermöglichte
die unkomplizierte und kompetente Betreuung des Manu-
skripts. Wertvolle Kommentare steuerten Peter Bliss (Halle/
Saale) und ein anonymer Gutachter bei.

Literatur

Becker L 1896 Les Arachnides de Belgique. Part 3. Opilio-
nes. - Annales du Musee Royal d’Histoire Naturelle de
Belgique 12: 339-369 - doi: 10.5962/bhl.title.48721

British arachnological Society 2013 Spider and harvestman
recording scheme website. Summary for Homalenotus
quadridentatus (Opiliones). - Internet: http;//srs.british-
spiders.org.uk/portal.php/p/Summary/s/Homalenotus%20
quadridentatus (aufgerufen am 12.04.2013)

Delfosse E 2004 La faune de France et dbutre-mer. Catalo-
gue preliminaire des Opilions de France metropolitaine
(Arachnida, Opiliones). - Bulletin de Phyllie 20: 34-56
Heinze W & Schreiber D 1984 Fine neue Kartierung der
Winterhärtezonen für Gehölze in Mitteleuropa. - Mit-
teilungen der Deutschen Dendrologischen Gesellschaft
75: 11-56

Hoc S 1966 Die Weberknechte des Regensburger Gebietes.

- Acta Albertina Ratisbonensia 26: 107-122

Bessert R de 1917 Opilions. - Catalogue des Invertebres
de la Suisse 9: 1-80

Martens J 1978 Spinnentiere, Arachnida - Weberknechte,
Opiliones. - Die Tierwelt Deutschlands 64: 1-464
Muster C, Blick T & Schönhofer A (im Druck) Rote Liste
der Weberknechte Deutschlands (Arachnida: Opiliones).

- Naturschutz und Biologische Vielfalt 70(4)
NoordijkJ öcWijnhoven H 2009 Hooiwagenwaarnemingen

uit Zeeuwse akkerranden (Opiliones). - Entomologische
Berichten 69: 78-82

Petto R 1991 Struktur und Dynamik der Spinnengemein-
schaften von Saumbiotopen. Dissertation, Aachen. 169

S.

Rambla M 1960 Contribuciön al estudio de los Opiliones
de la Fauna Iberica - La Nota sobre Opiliones de
Andalucia. - Publicaciones del Instituto de Biologia
Aplicada 31: 5-16

Rambla M 1967 Opiliones de Portugal. - Revista de Bio-
logia, Lisboa 6: 1-34

Spoek GL 1957 De hooiwagens (Arachnoidea, Opiliones)
van de Sint-Pietersberg en van andere delen van de
Provincie Limburg. - Natuurhistorisch Maandblad 46:
40-50

Spoek GL 1975 Spinachtigen - Archnoidea III. De hooiwa-
gens (Opilionida) van Nederland. Tweede,herziene druk.

- Wetenschappelijke Mededelingen KNNV 50: 1-32
Staudt A 2013 Nachweiskarten der Spinnentiere Deutschlands

(Arachnida: Araneae, Opiliones, Pseudoscorpiones). - Inter-
net: http:/ / spiderling. de/ arages, Homalenotus quadride?itatus\
http://spiderling.de/ arages/Verbreitungskarten/ species.
php?name=homqua (aufgemfen am 02.02.2013)
Vanhercke L 2004 Opiliones in Belgium - Species list and
distribution maps: Homalenotus quadridentatus (Cuvier).

- Internet: http://www.elve.net/opilio/hq.htm (vers.
08.2004; aufgerufen am 02.02.2013)

Wijnhoven H 2009 De Nederlandse hooiwagens (Opi-
liones). - Entomologische Tabellen 3: 1-118

Arachnologische Mitteilungen 45: 40-44

Karlsruhe, Juni 2013

A tropical invader, Coleosoma floridanum, spotted for the first time in Slovakia
and the Czech Republic (Araneae, Theridiidae)

Anna Sestakova, Jana Christophoryova & Stanislav Korenko

doi: 10.543 l/aramit4509

Abstract. The pantropical theridiid spider Coleosoma floridanum Banks, 1900 was recorded for the first time in Slo-
vakia and in the Czech Republic. Both sexes and juveniles were collected in some numbers in heated greenhouses
with high humidity. A description and photographs of the species are provided.

Keywords: botanical garden, comb-footed spider, faunistics, first record, greenhouse, introduced species

The small genus Coleosoma consists of nine tropical
species distributed mostly in the Indo-Malayan eco-
zone (Platnick 2012). Except for the largest species,
C. matinikum Barrion ScLitsinger, 1995 - known only
from males, with a total length of ca. 4.8 mm - the re-
maining species are of small size (ca. 2 mm). They are
thus easily accidently imported to other countries on
plants carried by ships. Despite this fact, only C.flori-
danum has so far spread to Europe. This species is
commonly found in packages arriving from tropics,
thus it has been exported over the globe and may be
expected to occur in any sort of tropical greenhouse.
The type locality for C. floridanum is situated in Flor-
ida; the species was also found on many islands in
the Caribbean, inferring that the Neotropical region
is probably its place of origin. However, some authors
pointed out that the Oriental region should be con-
sidered instead due to the presence of its related spe-
cies there (Levi 1967, Spoczynska 1969).

Species of this genus have a distinct sexual di-
morphism. Females have a basically oval abdomen, in
some species protruding as a tubercle above the spin-
nerets (C. acutiventer (Keyserling, 1884), C. blandum
O. P.-Cambridge, 1882).

Methods

Specimens were collected in the greenhouses of bo-
tanical gardens in Bratislava, Brno and Prague. They
were collected predominantly (ca. 90 %) on the un-

Anna SESTÄKOVÄ, Jana CHRISTOPHORYOVÄ, Department of Zoology,
Faculty of Natural Sciences, Comenius University, Mlynska dolina B-1 ,
SK-842 1 5 Bratislava, Slovakia, e-mail: asestakova@gmail.com, christo-
phoryova@gmail.com

Stanislav KORENKO, Department of Agroecology and Biometeorology,
Faculty of Agrobiology, Food and Natural Resources, Czech University of
Life Sciences, Kamycka 1 29, CZ-1 65 21 Prague 6, Suchdol, Czech Repub-
lic, e-mail: korenko.stanislav@yahoo.com

submitted 18. 12.2012, accepted 13.4.2013, online 13.5.2013

derside of plant leaves; some of them (ca. 10 %) were
extracted from soil samples using Tullgren funnels.
They were identified using Nentwig et al. (2012) and
compared to the original description (Banks 1900)
and to the other species of the genus through the
detailed description and figures provided by several
authors, e.g. Bryant (1940, 1944), Levi (1959), Bar-
rion dcLitsinger (1995) and Saaristo (2006).

Microphotographs were made using EOS Utility
software and a digital camera (Canon EOS HOOD)
connected to a Zeiss Stemi 2000-C stereomicroscope.
Microslides of epigynes were photographed using a
Leica ICC50 camera connected to a Leica DM1000
stereomicroscope using LAS EZ 1.8.0. Digital im-
ages were combined using CombineZP image stack-
ing software. Description of the species is based on
all mature specimens obtained in Slovakia. All meas-
urements are in millimeters, and were obtained using
AxioVision 4.8.2; M = median, x = arithmetic mean.
Material is deposited in 70 % ethanol in the collec-
tions of the first and the last author.

Results and discussion
Coleosoma floridanum Banks, 1900

The species has been described under various names;
a list of junior synonyms is given below. Many of
them were recognized by Levi (1959), although he
also pointed out that there should be further syno-
nyms within the genus Theridion.

Theridion interruptum Banks, 1908: 205, fig. 9 (de-
scribed 9). (Bryant 1944)

Bathyphantes semicincta Banks, 1914: 640 (undefined
sex). (Levi 1972)

Lithyphantes oophorus Petrunkevitch, 1930: 170, fig.
8-9 (described 9). (Levi 1959)

Theridion delebile Petrunkevitch, 1930: 206, fig. 53-56
(described 9). (Levi 1959)

Coleosoma floridanum in Slovakia and the Czech Republic

41

Jheridion rapanae Berland, 1942: 15, fig. 6a— f (de-
scribed (5$). (Levi 1959)

Theridium albovittatum Caporiacco, 1955: 334, fig.
25a-c (described 9). (Levi 1959)

Theridion aleipata Marples, 1955: 483, pi. 58, fig. 9,
13, 19 (described d9). (Levi 1959)

Coleosoma saispotum Barrion & Litsinger, 1995: 432,
fig. 258a-l (described c5). (Knoflach 1999)

Theridion antheae Barrion & Litsinger, 1995: 447, fig.
268a-d (described 9). (Knoflach 1999)

For the full list of references, see Platnick (2012).

Material examined

SLOVAKIA: Id (22.XI.2012); 1399, 2dd, 3subaddd,
5juv (12.XII.2012) Bratislava, greenhouse. Botanical
Garden of the Comenius University (average tempera-
ture 26 °C), 48°8’49.2”N; 17°4’20.97”E, 148 m a.s.l. (grid
square 7868, Fig. 1); leg. J. Christophoryova, M. Holecova,
K. Krajcovicova & A. Sestakova.

CZECH REPUBLIC: Id, 2599, 16juv (21.XI.2006) Brno,
greenhouse. Botanical Garden and Arboretum of Men-
del University (average temperature 26 °C), 49°12‘57“N,
16°36‘52“E, 245 m a.s.l. (grid square 6765, Fig. 1), leg. S.
Korenko, E. Liznarova & L. Sentenska.

Id (3.IV.2012) Prague, greenhouse, Prague Botanical
Garden (average temperature 26 °C/23 °C - day / night)
50°07’20”N, 14°24’50”E, 248 m a.s.l. (grid square 5852,
Eig. 1), leg. S. Korenko & B. Korenkova.

Diagnosis

The male can be easily distinguished from the other
males of the genus by an oval and markedly narrower
bulbus with a shorter semi-circular embolus - this is
nearly circular in the other species - and by a project-
ing terminal apophysis. Unlike other species, the con-
striction of the male abdomen is indistinct. Females
have a much simpler vulva with short ducts and a
markedly wider atrium than those of other species.

Description

Male: Total length 1.67-1.94 (M = 1.73; x = 1.77),
tibia + patella I 0.96-1.12 (M = 1.10; x = 1.06).

Males are ant-mimics. Carapace pale without
markings, dark bordered. Sternum pale with dark
narrow hem. Abdomen oval, more than twice as
long as wide, with slight median constriction; an-
teriorly stridulating, sclerotized scutum protruding
into blunt, bilobate projection over carapace, with
two long setae; the scutum continues ventrally half
the length of the abdomen. Colouration pale, distally
black; dorsally in the middle with irregularly distrib-
uted white spots and two longitudinal black stripes.
Legs long, thin, yellow; distal end of tibia IV black, in
some specimens all other tibiae distally dark (Fig. 2).
Bulbus oval, longer than wide; projecting strongly-
sclerotized terminal apophysis; embolus filiform and
semicircular (Fig. 4).

12“ 13“ 14“ 15“ 16“ 17“ 18“

42

A. Sestäkova, J. Christophoryovä & S. Korenko

Fig. 2: Male habitus of Coleosoma floridanum from Slovakia, a)
lateral, b) dorsal and c) ventral view. Arrows point to the biloba-
te sclerotization of the abdomen (Scale = 0.5 mm).

Female: Total length 1.34-1.87 (M = 1.62; x =
1.60), tibia + patella I 0.84-1.07 (M = 0.96; x = 0.96).

Female with similar colouration to the male, but
paler. Abdomen globular, pale; dorsally with irregular
white spots and two longitudinal interrupted stripes
reaching spinnerets; ventrally transverse black spot
above spinnerets. Legs yellow with dark distal end of
femur and tibia, mostly visible on the first and fourth
pair of legs (Fig. 3). Epigyne weakly sclerotized,
semitransparent; wide, transverse epigynal atrium;
spermathecae visible through cuticle, vulva with rela-
tively short copulatory ducts (Fig. 5).

Records in Europe

In September 1964, specimens of C. floridanum were
collected in Europe for the first time by Clark in the
heated greenhouse of the Kew Botanical Garden in
Britain. This record was published seventeen years
later by Hillyard (1981). The first published record of
the species in Europe is by Spoczynska (1969), who

collected in the same greenhouse in Britain on the
14.IX.1966. Additional first records from other Eu-
ropean countries are summarized in Tab. 1.

Tab. 1 : Current distributions and the first records of Coleosoma
floridanum in Europe.

State

Date of
collection

Reference of the
first record

Austria

29.VIII.1999

Knoflach (1999)

Czech Republic

21.XI.2006

present work

Finland

26.11.1990

Koponen (1990)

France

10.VIII.2006

Emerit & Ledoux
(2008)

Germany

1.1995

Broen et al. (1998)

Great Britain

IX.1964

Hillyard (1981)

Slovakia

22.XI.2012

present work

Switzerland

19.III.1999

Knoflach (1999)

The Netherlands

8.II.1995

Helsdingen (1995
dcpers. comm.)

Natural history

Coleosoma floridanum is a pantropical species. It
seems to be native to the American tropics (especial-
ly the Caribbean biozone). It has also been recorded
from Africa (Ghana, Togo, Seychelles), India, Poly-
nesia, New Hebrides, Hawaii, Galapagos Isl., Tai-
wan, Japan and China (Levi 1967, Spoczynska 1969,
Saaristo 1978,Tanikawa 1991, Knoflach 1999). Eu-
ropean populations are strictly synanthropic, being
recorded from highly humid and heated green-
houses (about 20-30 °C) where they can occur in
high numbers (Saaristo 2006). Similarly to natural
populations, they occupy analogous microhabitats
such as crevices in walls and tree bark, under stones,
beneath abandoned dry flowerpots and on vegeta-
tion (Cutler 1972, Platen dcBroen 2005). Spoczyn-
ska (1969) observed tiny webs - no more than 8 mm
in diameter - usually on the base of leaves; however
she found the majority of specimens outside their
webs. The egg sac (Fig. 3c) is attached to the spin-
nerets and guarded by the female until hatching; the
few eggs (ca. 10—12) are sparsely wrapped in white
threads (Knoflach 1999).

We observed numerous specimens under the
leaves of plants. Some were between stones and un-
der the flowerpots, and only few specimens were col-
lected within the soil. Our records are thus consist-
ent with known natural history patterns described by
several authors (e.g. Levi 1967, Cutler 1972, Platen
& Broen 2005, Harvey et al. 2002).

Coleosoma floridonum in Slovakia and the Czech Republic

43

Fig. 3: Female habitus of Coleosoma fJoridanum from Slovakia, and its cocoon, a) dorsal and b) ventral view, c) cocoon (Scale = 0.5
mm).

Fig. 4: Male palp of Coleosoma
floridanum from Slovakia, a) prola-
teral, b) ventral and c) retrolateral
view (Scale = 0.2 mm). CO = con-
ductor; EM = embolus; TA = termi-
nal apophysis.

Fig. 5: Epigyne of Coleosoma flo-
ridanum from Slovakia, a) exter-
nal and b) internal view (Scale =
0.2 mm).

44

A. Sestäkovä, J. Christophoryovd & 5. Korenko

Acknowledgements

We are indebted to personnel of the botanical gardens of
the Comenius University in Bratislava, Mendel University in
Brno, and Prague Botanical Garden who kindly allowed us
to collect spiders. Our thanks also go to Peter van Helsdin-
gen, Seppo Koponen and Jean-Claude Ledoux for sending
their publications, Katarina Krajcovicova,Milada Holecova,
Barborka Korenko va, Lenka Sentenska, and Eva Liznarova
for their help with collecting of spiders. Our special thanks
to JozefMertlik for providing the grid map. We are grateful
to Peter Dolejs and two other anonymous reviewers for their
valuable comments. This study was supported by the grant
VEGA 1/0176/09 and by the project of European Science
Foundation and Ministry for Education and Youth of the
Czech Republic CZ.1.07/2.3.00/30.0040.

References

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Ent3296-4

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Broen B von. Thaler- Knoflach B & Thaler K 1998 Nach-
weis von Coleosoma floridanum in Deutschland (Ara-
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Bryant EB 1940 Cuban spiders in the Museum of Compara-
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Cutler B 1972 Notes on the behavior of Coleosoma flori-
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- Nieuwsbrief SPINED 9: 4-6
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(Araneae, Theridiidae), found in the Botanical Garden of
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pro Fauna et Flora Femtica 66: 106-107
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Microscopical Society 91: 533-538
Nentwig W, Blick T, Gloor D, Hänggi A & Kropf C
2012 Spiders of Europe. Version 12.2012. - Internet:
http://www.araneae.unibe.ch (accessed at November

29.2012)

Platen R & Broen B von 2005 Gesamtartenliste und Rote
Liste der Webspinnen und Weberknechte (Arachnida:
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ten der gefährdeten Pflanzen und Tiere von Berlin.
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de/ natur_gruen/ naturschutz/ downloads/ artenschutz/
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3.2012)

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(accessed at November 25, 2012)

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Seychelle Islands, with notes on taxonomy. - Annales
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14: 49-89

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98/99: 1-7

Arachnologische Mitteilungen 45: 45-53

Karlsruhe, Juni 2013

Spider records from East Macedonia and Thrace (NE Greece)

Sascha Buchholz

doi: 1 0.543 l/aramit45 10

Abstract. The present study summarises spider records from East Macedonia and Thrace (NE Greece). Spiders were
mainly collected by hand sampling and to a smaller extent by pitfall trapping during field trips in 2002, 2004, 2007,
2008 and 2009. 171 species from 23 families were sampled. Pelecopsis pavida (O. P.-Cambridge, 1872) (Linyphiidae)
and Xysticus kaznakovi Utochkin, 1 968 (Thomisidae) were new to the European spider fauna and ten further species,
Diaea livens Simon, 1876, Herioeus grominicola (Doleschall, 1852), Meioneta ress// Wunderlich, 1973, Oedothorax re-
tusus (Westring, 1851), Philodromus albidus Kuiczynski, 1911, Pocadicnemis pumila Keyserling, 1880, Sitticus saltator
(O. P.-Cambridge, 1 868), Stemonyphontes lineatus (Linnaeus, 1 758), Synageles scutiger Proszynski, 1 979 and Thanatus
coloradensis Keyserling, 1880, could be added to the Greek fauna for the first time. 21 and 38 species were new to
East Macedonia and Thrace, respectively. Based on previous work, plus the present study, the Greek spider fauna

now includes 1,108 species.

Keywords: Araneae, Balkan, Nestos,Thassos

The Balkan Peninsula is a biodiversity hotspot
(Hubenov 2004, Krystufek & Reed 2004, Popov &c
Fet 2007) which comprises a very species rich spi-
der fauna (Deltshev 1999, 2005). Northern Greece
- part of the Balkan Peninsula - is situated in the
transition zone of the European, Mediterranean and
the Ponto-Anatolian biogeographical regions, which
results in a high biodiversity (Jerrentrup et al. 1989,
Schröder et al. 2011). Despite an increasing number
of faunistic surveys (Buchholz 2007, Van Keer et
al. 2010, Schröder et al. 2011) as well as taxonomic
works (for an extensive compilation see Bosmans &c
Chatzaki 2005 and Van Keer et al. 2010) the spider
fauna of Northern Greece, and especially Thrace, is
still poorly investigated and thus remains underesti-
mated (Chatzaki 2008). The present study summa-
rises spider data which were recorded in the Eastern
part of Northern Greece, namely East Macedonia
and Thrace, and thus is intended to enhance current
knowledge concerning spider distribution in this re-
gion.

Study area

Data were collected in the region of East Macedo-
nia (prefectures Drama, Kavala, Thassos) and Thrace
(prefectures Evros, Rhodopi, Xanthi) (Tab. 1, Fig. 1)
which are situated in the north-eastern part of Greece
and separated by the Nestos river. The study area is

Sascha BUCHHOLZ, TU Berlin, Institut für Ökologie, Rothenburgstraße
12, 12165 Berlin, Germany, e-Mail: sascha.buchholz@tu-berlin.de

submitted 28.2.2013, accepted 13.4.2013, online 13.5.2013

delimited by the Bulgarian border to the north and
the Turkish border or Evros river to the east, while
the coastline of the Thracian Sea forms the southern
border.

The northern part of north-east Greece is char-
acterised by mountain ranges such as the Greek
Rhodope massif along the Greek-Bulgarian border
reaching altitudes of about 1950 m above sea lev-
el as well as the Pangaion, Menoikon, Lekani and
Falakron mountains; the latter reaches the highest
altitudes (2232 m a.s.k). Towards the east, a lower
mountainous area with altitudes not exceeding 650
m a.s.l. exists in the area of the Dadia nature re-
serve. In a southern and south-eastern direction the
landscape becomes gradually lower and, via a belt of
cultivated land, changes into the coastal plain of the
Aegean Sea. Near the coast the landscape is formed
by level to slightly undulated lowlands about 20 m
above sea level. In the study area two rivers (Nestos,
Evros) form deltas which are largely cultivated, but
also still contain many natural habitats such as la-
goons, salt marshes and remnants of floodplain for-
ests. The Nestos delta includes the lagoon areas of
Keramoti and Agiasma, and the eastward situated
Lakes Vistonis and Mitrikou form the east Mac-
edonian-Thracian wetland belt, which comprises
a considerable number of natural lagoons and salt
marshes.

The climate of the coastal plains is Mediterra-
nean, although continental impacts become effective.
Impacts manifest themselves as huge fluctuations in
yearly temperature, with summer maxima of 40 °C

46

S. Buchholz

_d.

Tab. 1: Geographical information on the sampling localities.

No.

N

E

m a.s.l.

Location /Area

Region

Prefecture

1

41°18’01.36”

24°04’42.31”

1950

Falakron

East Macedonia

Drama

2

41°00’46.62”

24°17’03.30”

70

Philippi

East Macedonia

Kavala

3

4r02’51.14”

24°38’41.66”

190

Stenopos

East Macedonia

Kavala

4

40°51’13.53”

24°43’30.33”

0

Nestos Delta

East Macedonia

Kavala

5

40°50’58.25”

24°47’48.94”

0

Nestos estuary

East Macedonia

Kavala

6

40°46’40.86”

24°43’12.16”

120

Thassos

East Macedonia

Thassos

7

41°00’27.72”

24°42’24.39”

20

Aladjagiola

East Macedonia

Kavala

8

4r01’37.76”

24°38’04.75”

360

N Zarkadia

East Macedonia

Kavala

9

4r05’49.18”

24°45’10.85”

60

Nestos canyon

Hirace

Xanthi

10

4r08’56.73”

24°42’18.21”

460

Ano Livera

Thrace

Xanthi

11

41°06’57.70”

24°44’52.37”

760

Galani/Nestos view

Thrace

Xanthi

12

4r08’46.16”

24°53’04.24”

160

N Xanthi

Thrace

Xanthi

13

4ri5’58.72”

24°48’45.96”

430

W Oreo

Thrace

Xanthi

14

41°02’00.92”

25°04’16.25”

20

Nea Kessani

Thrace

Xanthi

15

41°00’29.09”

25°08’48.95”

0

Lake Vistonis

Thrace

Xanthi

16

40°59’28.32”

25°18’23.91”

10

Lake Mitrikou

Thrace

Rhodopi

17

41°00’30.27”

25°07’13.74”

0

Porto Lagos

Thrace

Xanthi

18

4r00’12.62”

25°10’29.98”

5

SW Glykoneri

Thrace

Rhodopi

19

4r08’23.80”

25°12’37.69”

40

Kompsatos river

Thrace

Rhodopi

20

4ri7’56.73”

26°01’16.01”

350

Roussa

Thrace

Evros

21

4r07’19.97”

26°13’40.10”

100

Dadia

Thrace

Evros

Fig. 1 : Map of study area and location of sampling sites in East Macedonia (west of the Nestos river) and Thrace (east of the Nestos
river) (pale area = lowland, grey area = mountain range). For further geographical information see Tab. 1 .

Spiders from NE Greece

47

and winter minima of -20 °C. The annual average
amount of precipitation is 600-700 mm (Jerrentrup
et al. 1989), whereas the average annual temperature
is 11 °C (Lienau 1989). The climate of the mountain
ranges shows typical Central European characte-
ristics with snow from November to May (Volpers

1988) . The potential natural vegetation along the
coast is the Ostryo-Carpinion followed by Quercion
frainetto-zone up to altitudes of 1000 meters above
sea level (Horvat et al. 1974). At about 1000 meters
the oak-zone changes into a beech-zone with spru-
ce {Picea abies) and fir {Abies) (Volpers 1988, Lienau

1989) .

Methods

Spiders were mainly collected by hand sampling and
to a lesser extent by pitfall traps during field trips in
2002, 2004, 2007, 2008 and 2009. A broad variety of
habitat types were sampled in 21 locations in East
Macedonia and Thrace (Tab. 1, Fig. 1). Nomencla-
ture and information on zoogeographical distribu-
tion of species were taken from Helsdingen (2012)
and Platnick (2013) and more detailed literature on
local elements. Bosmans & Chatzaki (2005), Buch-
holz (2007), Chatzaki (2008), Bosmans (2009),
Bosmans et al. (2009), Van Keer et al. (2010) and
Schröder et al. (2011) were consulted to identify
those new records (indicated by ‘nr’ in Tab. 2) that
the present study added to the list of known species
in Greece, Macedonia and Thrace. By contrast, the
‘+’ sign in Tab. 2 indicates species that were already
recorded in the area.

Results and discussion

In total, 171 species from 23 families were record-
ed (Tab. 2). Two species - Pelecopsis pavida (O. R-
Cambridge, 1872) (Linyphiidae) (det. R. Bosmans)
and Xysticus kaznakovi Utochkin, 1968 (Thomisidae)
(det. D. Logunov) - proved to be new for the Eu-
ropean spider fauna. Pelecopsis pavida was hitherto
only known from its type locality in Palestine (Bos-
mans 1994). During the present study, one male was
sampled in a grey dune habitat in the Nestos delta.
Xysticus kaznokovi has been recorded in Central Asia
(Marusik 6c Logunov 1990) and Turkey, where it was
collected under stones and on the ground in grass
(Demir et al. 2009). One male was caught in a dry
and sparse vegetated habitat.

Twenty-one and thirty-eight species were new to
East Macedonia and Thrace, respectively. In addition

to Pelecopsis pavida and Xysticus kaznakovi^ ten fur-
ther species could be added to the Greek fauna for
the first time:

Oedothorax retusus^ Pocadicnemis pumila and Ste-
monyphantes lineatus are widespread across Europe
(Nentwig et al. 2013) and inhabit a broad variety of
habitats (Hänggi et al. 1995).

Sitticus saltatoris also an element of the European
fauna (Nentwig et al. 2013), but its occurrence seems
to be restricted to dry habitats such as dry grassland,
heathland and dunes (Bauchhenß 1995, Merk-
ens 2002, Buchholz 6c Kreuels 2009, Buchholz 6c
Schirmel 2011) and accordingly three females were
found in a white dune of the Nestos delta.

Meioneta ressli (det. R. Bosmans) and Thana-
tus coloradensis (det. C. Muster) are high mountain
species (Muster 2001, Nentwig et al. 2013). While
the latter was found in meadows of the European
mountains (Alps, Pyrenees, Carpathians, Caucasus)
(Nentwig et al. 2013), Meioneta ressli was hitherto
considered an endemic species of the Alps (Muster
2001) inhabiting alpine grass heaths and as far down
as the valley (Thaler 1995, Muster 2001, Höfer et al.
2010). In the present study, Thanatus coloradensis was
sampled in dry grasslands of the Falakron mountain
while Meioneta ressli was found at the same habitat
type in the Nestos canyon.

Philodromus albidus (det. C. Muster), which was
sampled from rocks, is mainly distributed in Western
and Central Europe (Nentwig et al. 2013, Platnick
2013) but was also recorded in Bulgaria (Lazarov
2007) and Turkey (Bayram et al. 2013).

Diaea livens has been found in Western, Central
and Southern Europe (Lazarov 2007, Bayram et al.
2013, Nentwig et al. 2013) as well as in the Cauca-
sus (Otto 6c Tramp 2011, Nentwig et al. 2013) but
was also introduced into the USA (Platnick 2013).
Diaea livens is arboreal (mainly oaks) (Nentwig et
al. 2013) and accordingly this species was caught in
dwarf- shrubs.

Heriaeus graminicola is distributed in Central,
Eastern and South-Eastern Europe (Deltshev et al.
2004, Nentwig et al. 2013) and according to Hänggi
et al. (1995) and Nentwig et al. (2013) it inhabits
dense vegetated and humid habitats (e.g., reedy
marsh). By contrast, a number of individuals were
sampled in different and mostly dry habitats.

Synageles scutiger (det. D. Logunov) has been
rarely recorded. According to Logunov 6c Marusik
(2000) and Platnick (2013) it was only found in

48

S. Buchholz

Tab. 2: Species list (nomenclature follows Platnick 2013). Abbreviations; biogeogr = biogeographical type: BIk = Balkan, Ci-Me =
Circum-Mediterranean, EaMe = East-Mediterranean, Eu = European, Gr = Greek, Tur = Turanian (species with western limits of dis-
tribution formed by the Italian and Balkan peninsulas, respectively, and ranging as far as Central Asia), Tur-Eu = European species
whose ranges extend eastward to Central Asia, Tur-Me = Mediterranean species whose ranges extend eastward to Central Asia, Wi =
widely distributed species (cosmopolitan, palaearctic, holarctic); Gr = Greece, Ma = East Macedonia, Th =Thrace (nr = new record, -i- =
species already recorded, ; m/f = number of sampled males and females, respectively; habitat types: A = building, B = dry grassland,
C = dry habitat with sparse vegetation, D = dwarf shrub, E = fallow land, F = floodplain forest, G = fringe, H = gravel bank, 1 = grey
dune, J = humid meadow, K = meadow, L = P/nus-forest, M = pseudo maquis, N = reed, 0 = riparian strip, P = rocks, Q = salt meadow,
R = sandy shore, S = semi-dry grassland, T = tall-forb vegetation, U = white dune.

species

biogeogr

Gr

Ma

Th

m/f

sampling

location

habitat

type

Scytodidae

Scytodes thoracica (Latreille, 1802)

Pholcidae

Wi

+

+

+

1/0

21

P

Holocnemus pluchei (Scopoli, 1763)

Dysderidae

Eu

+

+

+

0/1

9

P

Harpactea babori (Nosek, 1905)

Mimetidae

EaMe

+

+

1/0

4

F

Mimetus laevigatas (Keyserling, 1863)

Eresidae

Tur-Me

+

+

1/0

4

F

Eresus kollari Rossi, 1846

Tur-Eu

+

+

+

5/0

2, 3, 7, 9

B

Oecobiidae

Oecobius maculatus Simon, 1870

Tur-Me

+

+

nr

0/1

21

P

Uloboridae

Uloborus walckenaerius Latreille, 1806

Wi

+

+

+

1/1

4

I

Theridiidae

Enoplognatha penelope Hippa & Oksala, 1982

Blk

+

+

0/2

7

E

Euryopis episinoides (Walckenaer, 1847)

Tur-Me

+

+

0/1

4

a

Euryopis sexalbomaculata (Lucas, 1846)

Ci-Me

+

+

3/0

6

M

Kochiura aulica (C. L. Koch, 1838)

Wi

+

+

1/0

7

M

Latrodectus tredecimguttatus (Rossi, 1790)

Tur-Me

+

+

+

1/0

4

I

Paidiscura dromedaria (Simon, 1880)

Wi

+

nr

0/2

4

U

Phylloneta impressa (L. Koch, 1881)

Wi

+

+

+

1/1

7

E

Steatoda triangulosa (Walckenaer, 1802)

Linyphiidae

Wi

+

+

+

0/1

21

A

Erigone atra Blackwall, 1833

Wi

+

nr

1/0

12

A

Erigone dentipalpis (Wider, 1834)

Wi

+

+

1/0

4

a

Frontinellina frutetorum (C. L. Koch, 1834)

Wi

+

+

+

0/7

4, 20

K,P

Gnathonarium dentatum (Wider, 1834)

Wi

+

+

0/1

7

N

Gongylidium rufipes (Linnaeus, 1758)

Wi

+

+

0/1

4

F

Maso gallicus Simon, 1894

Wi

+

+

0/1

4

K

Maso sundevalli (Westring, 1851)

Wi

+

+

7/0

4

F

Mecopisthes peusi Wunderlich, 1972

Eu

+

+

0/9

4,9

K,R

Meioneta fuscipalpa (C. L. Koch, 1836)

Wi

+

+

nr

6/0

4, 9,21

K, R,T

Meioneta ressli Wunderlich, 1973
Metopobactrus prommulus

Eu

nr

nr

1/0

9

B

(O. R-Cambridge, 1872)

Wi

+

+

0/1

4

J

Oedothorax apicatus (Blackwall, 1850)

Wi

+

+

1/3

7

N,S

Oedothorax retusus (Westring, 1851)

Wi

nr

nr

0/1

2

P

Pelecopsis elongata (Wider, 1834)

Eu

+

+

0/1

4

F

Pelecopsis inedita (O. P.-Cambridge, 1875)

Ci-Me

+

nr

0/2

4

F,I

Pelecopsis pavida (O. P.-Cambridge, 1872)

Tur

nr

nr

1/0

4

I

Spiders from NE Greece

49

species

biogeogr Gr

Ma 111

m/f sampling

location

habitat

type

Prinerigone vagans (Audouin, 1826)

Wi

+

+

+

1/6

4,6,9,15

F,K,M,Q,T

Pocadicnemis juncea Locket ScMillidge, 1953

Wi

+

+

0/28

4

a

Pocadicnemis pumila (Blackwall, 1841)

Wi

nr

nr

1/0

7

N

Stemonyph antes lineatus (Linnaeus, 1758)

Wi

nr

nr

0/1

4

a

Styloctetor romanus (O. P.-Cambridge, 1872)

Wi

+

+

0/9

4

I

Tenuiphantes tenuis (BlackwaU, 1852)

Wi

+

+

nr

0/1

20

c

Trichoncus hackmani Millidge, 1955

Eu

+

+

0/1

7

E

Walckenaeria alticeps (Denis, 1952)

Tur-Eu

+

+

0/1

4

F

Walckenaeria vigilax (Blackwall, 1853)

Tetragnathidae

Wi

+

+

1/0

4

J

Tetragnatha extensa (Linnaeus, 1758)

Wi

+

+

+

1/0

20

O

Tetragnatha montana Simon, 1874

Wi

+

+

1/0

7

s

Tetragnatha obtusa C. L. Koch, 1837

Araneidae

Wi

+

+

nr

1/0

15

T

Araneus angulatus Clerck, 1757

Wi

+

+

+

3/19

7

G,M,S

Araniella cucurbitina (Clerck, 1757)

Wi

+

+

+

0/2

3

K

Araniella opisthographa (Kulczynski, 1905)

Tur-Eu

+

+

+

2/4

7,19,20

H,M,0

Argiope bruennichi (Scopoli, 1772)

Wi

+

+

+

8/9

7

E, G,M,N,S

Argiope lobata (Pallas, 1772)

Wi

+

+

+

0/3

17

K

Cyclosa sierrae Simon, 1870

Tur-Eu

+

nr

+

0/12

7,20,21

K, L,M

Gibbaranea bituberculata (Walckenaer, 1802)

Wi

+

+

+

0/3

7,20

B,C

Hypsosinga albovittata (Westring, 1851)

Wi

+

+

+

0/4

6,7

E,P,T

Hypsosinga pygmaea (SundevaU, 1831)

Wi

+

+

0/1

7

S

Larinioides cornutus (Clerck, 1757)

Wi

+

+

nr

1/9

4,7,16

E,I,J,

K,N,S

Mangora acalypha (Walckenaer, 1802)

Wi

+

+

+

8/46

7,10, 20,21

B,C,E, G,
K,M,N,0,S

Neoscona adianta (Walckenaer, 1802)

Lycosidae

Wi

+

+

+

29/76

7,16,18

E,G,J,K,

M,N,S

Alopecosa albofasciata (Brülle, 1832)

Tur-Me

+

+

+

6/8

7,9,18,19,

20,21

B,E,H,K,

L,P

Alopecosa cuneata (Clerck, 1757)

Wi

+

+

+

0/2

1

B

Arctosa cinerea (Fabricius, 1777)

Wi

+

+

+

2/2

5,7,20

O, R,U

Arctosa leopardus (SundevaU, 1833)
Aulonia kratochvili Dunin, Buchar &.

Wi

+

+

+

3/0

7

J,N

Absolon, 1986

Tur-Me

+

+

+

1/0

7

J

Geolycosa vultuosa (C. L. Koch, 1838)

Tur-Me

+

+

+

0/10

3,7,9

B,S

Pardosa agricola (Thorell, 1856)

Tur-Eu

+

+

nr

1/4

20

K

Pardosa atomaria (C. L. Koch, 1847)

Blk

+

+

+

0/2

20

O

Pardosa blanda (C. L. Koch, 1833)

Wi

+

+

3/3

1

B

Pardosa cribrata Simon, 1876

Ci-Me

+

+

4/3

7

J,N

Pardosa hortensis (Thorell, 1872)

Wi

+

+

+

0/5

7, 10,20

B,K

Pardosa monticola (Clerck, 1757)

Wi

+

+

0/1

1

B

Pardosa paludicola (Clerck, 1757)

Wi

+

+

1/1

1,7

J,s

Pardosa prativaga (L. Koch, 1870)

Tur-Eu

+

+

1/2

7

J,N

Pardosa proxima (C. L. Koch, 1847)

Wi

+

+

+

2/0

7

J

Pirata latitans (Blackwall, 1841)

Tur-Eu

+

+

+

0/2

4

F

Trochosa ruricola (De Geer, 1778)

Wi

+

+

+

3/2

7

J,N

Xerolycosa miniata (C. L. Koch, 1834)

Wi

+

+

nr

0/1

20

K

50

S. Buchholz

species

biogeogr

Gr

Ma

Th

m/f

sampling

location

habitat

type

Pisauridae

Pisaura mirabilis (Clerck, 1757)

Wi

+

+

+

0/13

7,9,20,21

b,e,j,k,

p,s

Pisaura novicia (L. Koch, 1878)

Tur-Me

+

nr

0/1

3

K

Oxyopidae

Oxyopes heterophthalmus (Latreille, 1804)

Wi

+

+

+

4/8

3,7,20,21

e,g,j,k,

R,T

Oxyopes lineatus Latreille, 1806

Wi

+

+

+

6/6

7,9,20,21

B,C,K, R,T

Oxyopes nigripalpis Kulczyhski, 1891

Ci-Me

+

+

nr

5/0

21

T

Agelenidae

Agelena orientalis C. L. Koch, 1837

Tur-Me

+

+

0/1

7

M

Maimuna vestita (C. L. Koch, 1841)

EaMe

+

+

+

0/4

6,21

L,M,P

Tegenaria angustipalpis Levy, 1996

EaMe

+

+

0/1

6

P

Tegenaria parietina (Fourcroy, 1785)

Wi

+

+

nr

0/1

13

A

Dictynidae

Devade indisticta (O. P.-Cambridge, 1872)

Ci-Me

+

nr

4

a

Dictyna arundinacea (Linnaeus, 1758)

Wi

+

+

+

12/22

7,20

E, G,J,N,
O, R,S

Titanoecidae

Nurscia albomaculata (Lucas, 1846)

Tur-Eu

+

+

0/1

6

P

Zodariidae

Zodarion blagoevi Bosnians 2009

Blk

+

+

10/0

7

R

Zodarion epirense Brignoli, 1984

Blk

+

+

0/4

7

G,P

Zodarion frenatum Simon, 1884

EaMe

+

+

2/12

4

B, K,p,au

Zodarion granulatum Kulczyhski, 1908

EaMe

+

nr

1/0

7

G

Zodarion hauseri Brignoli, 1984

Gr

+

+

1/0

7

R

Zodarion morosum Denis, 1935

Eu

+

+

2/0

4,7

E

Zodarion pirini Drensky, 1921

Blk

+

nr

3/0

4

K,R

Zodarion thoni Nosek, 1905

Tur-Eu

+

nr

4/0

4,7

B,U

Gnaphosidae

Callilepis cretica (Roewer, 1928)

Tur-Me

+

+

+

1/0

19

H

Drassodes lapidosus (Walckenaer, 1802)

Wi

+

+

+

1/1

8,19

K,P

Drassodes lutescens (C. L. Koch, 1839)

Tur-Me

+

+

nr

0/4

6, 19,20,21

C,P

Gnaphosa lucifuga (Walckenaer, 1802)

Wi

+

+

+

2/4

4, 20

o,a

Gnaphosa lugubris (C. L. Koch, 1839)

Tur-Eu

+

+

0/1

1

B

Haplodrassus dalmatensis (L. Koch, 1866)

Wi

+

+

nr

1/0

14

A

Haplodrassus signifer (C. L. Koch, 1839)

Wi

+

+

+

0/1

21

A

Nomisia exornata (C. L. Koch, 1839)

Tur-Eu

+

+

nr

7/8

6,9,19,20,21

B,C,H,L,

M,S

Nomisia ripariensis (O. P.-Cambridge, 1872)

Tur

+

+

+

4/0

4, 6,18

K,P,S

Trachyzelotes barbatus (L. Koch, 1866)

Tur-Me

+

+

0/1

9

B,P

Trachyzelotes lyonneti (Audouin, 1826)
Trachyzelotes malkini Platnick &

Tur-Me

+

+

0/1

4

a

Murphy, 1984

Eu

+

+

2/1

4,7

B, K,S

Zelotes argoliensis (C. L. Koch, 1839)

Blk

+

+

+

0/1

9

R

Zelotes caucasius (L. Koch, 1866)

Tur-Eu

+

+

nr

3/0

21

P

Zelotes cingarus (O. P-Cambridge, 1874)

EaMe

+

+

nr

0/1

21

P

Zelotes tenuis (L. Koch, 1866)

Ci-Me

+

+

1/1

4

I, a

Spiders from NE Greece

51

species biogeogr Gr Ma Th m/f sampling habitat

location type

Sparassidae

Micrommata ligurina (C. L. Koch, 1845) Tur-Me

Philodromidae

Philodromus albidus Kulczyhski 1911 Eu

Philodromus glaucmus Simon, 1870 Ci-Me

Philodromus lunatus Muster 6c Thaler, 2004 EaMe
Philodromus pulchellus Lucas, 1846 Ci-Me

Thajiatus atratus Simon, 1875 Wi

Thanatus coloradei^sis Keyserling, 1880 Wi

Thanatus pictus L. Koch, 1881 Wi

Thanatus vulgaris Simon, 1870 Wi

Tibellus oblongus (Walckenaer, 1802) Wi

Tbomisidae

Diaea livejis Simon, 1876 Tur-Eu

Heriaeus graminicola (Doleschall, 1852) Tur-Eu

Tieriaeus setiger (O. P.-Cambridge, 1872) Wi

Misumena vatia (Clerck, 1757) Wi

Mojiaeses israeliensis Levy, 1973 Tur-Me

Ozyptila confluens (C. L. Koch, 1845) EaMe

Runcinia grammica (C. L. Koch, 1837) Wi

Synema globosum (Fabricius, 1775) Wi

Synema plorator (O. P.-Cambridge, 1872) Tur-Me

Thomisus onustus Walckenaer, 1805 Wi

Tmarus piger (Walckenaer, 1802) Wi

Xysticus caperatus Simon, 1875 Ci-Me

Xysticus kaznakovi Utochkin, 1968 Tur

Xysticus kochi Thorell, 1872 Wi

Xysticus laetus Thorell, 1875 Tur-Me

Xysticus lanio C. L. Koch, 1835 Wi

Xysticus tristrami (O. P.-Cambridge, 1872) Tur-Me

Salticidae

Aelurillus luctuosus (Lucas, 1846) Tur-Me

Bianor albobimaculatus (Lucas, 1846) Tur-Me

Chalcoscirtus nigritus (Thorell, 1875) Wi

Cyrba algerina (Lucas, 1846) Wi

Evarcha arcuata (Clerck, 1757) Wi

Evarcha jucunda (Lucas, 1846) Ci-Me

Heliophanus equester L. Koch, 1867 Tur-Me

Heliophanus flavipes (Hahn, 1832) Wi

Heliophanus kochii Simon, 1868 Wi

Heliophanus melinus L. Koch, 1867 Wi

Heliophanus tribulosus Simon, 1868 Tur-Eu

Icius hamatus Wi

Macaroeris nidicolens (Walckenaer, 1802) Tur-Eu

Menemerus semilimb atus (Hahn, 1829) Wi

Menemerus taeniatus (L. Koch, 1867) Tur-Me

Mogrus neglectus (Simon, 1868) Tur-Me

Neaetha membrosa (Simon, 1868) Eu

+

+

+

0/1

16

K

nr

nr

0/1

8

P

+

+

0/1

7

T

+

+

nr

2/1

7, 20,21

C,M,P

+

+

0/1

4

E,S

+

+

nr

10/11

4,6,7, 10,20

B,E,N,0,

P,S

nr

nr

1/2

1

B

+

nr

nr

3/0

4,16

K,U

+

+

0/3

4,7

s,u

+

+

nr

8/20

7,9

B, F, G, K,

N,0,S

nr

nr

1/0

8

D

nr

nr

nr

8/0

7,20

C,G,K,S

+

+

nr

2/0

4,21

B,T

+

+

nr

0/4

7,21

S,T

+

+

1/0

7

G

+

+

1/0

4

A

+

+

+

20/1

7,21

E,G,N,S,T

+

+

+

5/3

15,21

K,T

+

+

0/1

8

P

+

+

+

8/0

9,20,21

B, K,P,T

+

+

+

2/0

4,7

F,R

+

+

1/0

6

M,P

nr

nr

1/0

4

C

+

+

+

1/3

7,9

B, E, G,J

+

+

nr

4/1

4,21

F,T

+

+

0/2

20

c,o

+

nr

3/1

21

P

+

nr

0/1

21

P

+

+

+

0/1

9

R

+

+

1/0

4

K

+

nr

1/1

9,19

B,P

+

+

+

3/2

7

N,S

+

+

nr

5/0

9,19,21

B, H, L, P

+

+

+

1/5

7,16,20

E,N,S,T

+

+

+

1/0

7

N

+

+

+

0/1

20

K

+

+

+

8/2

9,10, 19,20,21B,C,P

+

+

1/0

6

M

+

+

nr

1/1

18

K

+

+

nr

2/0

20,21

C,L

+

+

nr

3/2

2,4, 6,15

P,T,U

+

+

nr

0/1

18

K

+

+

nr

2/4

4,7,9

B,G,P,S,U

+

+

1/0

9

B,P

52

S. Buchholz

species

biogeogr Gr

Ma Th

m/f sampling

location

habitat

type

Pellenes brevis (Simon, 1868)

Eu

+

+

1/0

7

R

Pellenes diagonalis (Simon, 1868)

EaMe

+

+

0/5

6, 7,8

B,P,S

Pellenes ßavipalpis (Lucas, 1853)

EaMe

+

nr

1/0

19

P

Pellenes geniculatus (Simon, 1868)

Wi

+

nr

2/4

9, 19,20

B,H,K,P

Pellenes nigrociliatus (Simon, 1875)

Wi

+

+

+

1/0

9

R

Pellenes seriatus (Thoreil, 1875)

Tur-Me

+

+

0/3

7

E

Philaeus chrysops (Poda, 1761)

Wi

+

+

+

9/1

3, 6, 8, 9,
19,21

B,H,K,P

Phintella castriesiana (Grube, 1861)

Wi

+

+

1/0

4

F

Phlegra fasciata (Hahn, 1826)

Wi

+

+

+

4/2

3,7, 9,21

E, L,N,P

Pseudeuophrys obsoleta (Simon, 1868)

Wi

+

+

+

1/0

7

M

Pseudicius picaceus (Simon, 1868)

Tur-Me

+

nr

1/0

9

P

Saitis tauricus Kulczynski, 1905

Tur-Me

+

+

nr

1/2

6,21

A,P

Salticus mutabilis Lucas, 1846

Wi

+

+

nr

1/0

18

K

Salticus noordami Metzner, 1999

Gr

+

nr

1/0

9

B,P

Sitticus atricapillus (Simon, 1882)

Eu

+

+

0/1

1

B

Sitticus saltator (O. P. -Cambridge, 1868)

Wi

nr

nr

0/3

4

U

Synageles dalmaticus (Keyserling, 1863)

Ci-Me

+

+

3/0

5,6

A,P,U

Synageles scutiger Pröszynski, 1979

Tur-Eu

nr

nr

0/1

4

U

Ukraine and Azerbaijan. One female was caught in a
white dune of the Nestos delta.

Knowledge of the distribution patterns and ecol-
ogy of several of the species caught is still poor, and
more faunistic and especially ecological studies are
needed to gain a better understanding of the Greek
but also of the Eastern Mediterranean spider fauna.
However, based on previous work (Bosmans & Chat-
zaki 2005, Bosmans et al. 2009, Van Keer et al. 2010,
Schröder et al. 2011), together with the present study,
the Greek spider fauna now includes 1,108 species.

Acknowledgements

I would like to thank Mareike Breuer, Lars Gaedicke, Nils
Hein, Peter Krech, Jonas Linke, Tomasz Luzar, Hermann
Mattes, Dorothee Rolfsmeyer, Anne Schulze-Niehoff,
Johanna Siewers and Maria Schröder for hand sampling
species. Many thanks to Robert Bosmans, Maria Chatza-
ki, Barbara Knoflach-Thaler, Dmitri Logunov, Christoph
Muster and Stano Pekar for identifications and checking
of doubtful species.

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passen.

Legenden (in deutschen Artikeln deutsch und englisch)

sind in normaler Schrift über den Tabellen (Tab. 1: ...) bzw.
unter den Abbildungen (Abb. 1/Fig. 1: ...) anzuordnen.
Fußnoten können nicht berücksichtigt werden.
Literaturzitate: Im Text wird ab drei Autoren nur der Erst-
autor zitiert (Schulze et al. 1969), mehrere Zitate sind durch
Komma getrennt (Koponen 2000, 2008, Huber 2010). Im
Literaturverzeichnis werden die Arbeiten alphabetisch nach
Autoren geordnet und die Zeitschriftentitel ausgeschrieben.
Arbeiten mit identischem/n Autor/en und Jahr werden mit
a, b, c ... gekennzeichnet.

Beispiele:

Broen B von 1993 Nachweise selten gefundener oder gefährdeter
Spinnen (Araneae) in der Mark Brandenburg. - Arachnologi-
sche Mitteilungen 6: 12-25

Chatzaki M 2008 A critical review of the spider family Gnaphosidae
in Greece. In: Makarov SE &Dimitrijevic RN (eds.) Advances
in arachnology and developmental biology. Pensoft, Sofia, pp.
355-374

Clayhills T, Rinne V &c Koponen S 2008 On insect and spider fauna
of Jungfruskär in Houtskär, 2007. Unpublished report to the
Finnish Forest and Park Service, Turku. 80 pp. (in Finnish)
Lindgren L 2000 Island pastures. Metsähallitus and Edita Ltd.
Helsinki. 203 pp.

Merkens S 2000 Die Spinnenzönosen der Sandtrockenrasen im
norddeutschen Tiefland im West-Ost-Transekt - Gemein-
schaftsstruktur, Habitatbindung, Biogeographie. Dissertation,
Univ. Osnabrück. 165 pp.

Nentwig W, Blick T, Gloor D, Hänggi A & Kropf C 2013 araneae
- Spiders of Europe, version 1.2013. - Internet: http://www.
araneae.unibe.ch (4.1.2013)

Platnick NI 20 12 The world spider catalog, version 13.0. - Internet:
http://research.amnh.org/iz/spiders/catalog (15.9.2012)
Doi-Links sind, wenn möglich, mit anzugeben. Beispiel:
Kovblyuk MM & Marusik YM 2012 First description of the
female of the theridiid spider Robertus golovatchi (Araneae:
Theridiidae). - Arachnologische Mitteilungen 44: 17-19 - doi:
10.543 l/aramit4405

Gliederung:

Auf den präzise gehaltenen Titel folgt in der nächsten Zeile
der/die Verfasser, jeweils mit vollem Vornamen und Nachna-
men, bei mehreren Autoren der letzte mit &c angehängt.
Darunter ein englischsprachiges Abstract. ..., das bei deut-
schen Artikeln mit der Übersetzung des Titels beginnt und
die wesentlichen Ergebnisse der Arbeit kurz zusammenfasst.
Anschließend wenige, präzise Keywords (alphabetisch sor-
tiert), die Titel und englischen Titel ergänzen (sinnvoll für
die bibliographische Erfassung).

Für die Kopfzeilen ist ein Kurztitel anzugeben.

Der Text ist wie folgt zu gliedern: einführendes Kapitel (ohne
Titel), Material und Methoden, Ergebnisse und Diskussion.
Eine Danksagung steht vor der Literatur.

Für die Autorenftißzeile bitte den vollen Namen (Vorname
NACHNAME), Anschrift der Verfasser und E-Mail angeben.

Hinweise zu digitalen Formaten:

Das Manuskript soll in Microsoft-kompatibler Form, vor-
zugsweise als MS-WORD-Datei (name.doc) oder im Rich-
Text-Format (name.rtf) per E-Mail eingereicht werden. In
MS-Office erstellte Tabellen können in das WORD-Do-
kument eingebettet sein oder als getrennte Dateien geliefert
werden. Letzteres ist für alle Bilder erwünscht. Es ist darauf
zu achten, dass MS-EXCEL-Grafiken auch als Datei einge-
reicht werden, da bei der Vorbereitung für den Satz häufig die
zugrundeliegende Datentabelle benötigt wird. Fotos werden
in Halbton (8bit) gedruckt, sollten aber in Farbe eingereicht
werden (RGB, 4-5 MegaPixel), da sie in den digitalen Sepa-
rata (.pdf) auch farbig eingebettet werden. Alle Zeichnun-
gen, sowohl flächenhafte (z.B. Bleistiftzeichnungen) als auch
Strichzeichnungen (z.B. Tusche), sind als Halbton-Bilddatei-
en (8bit, 4-5 MegaPixel, TIEF- oder PSD-Datei) zum Satz
einzureichen. Fotos können auch als JPG-Datei (etwa Origi-
naldateien der Kamera) beigelegt werden. Sie werden anhand
der Abbildungszusammenstellung des Autors und den Le-
genden arrangiert und beschriftet. Alternativ kann der Autor
die Abbildungen auch fertig arrangiert digital einreichen. In
diesem Fall ist die Beschriftung der Einzelabbildungen so zu
legen, dass sie beim Satz zur Anpassung an die Schriften der
Zeitschrift entfernt werden können (im Photoshop etwa auf
einer getrennten Textebene).

Instructions to Authors

The journal “Arachnologische Mitteilungen” publishes
scientific papers about Western Palaearctic arachnids (ex-
cluding mites and ticks) in German or British English.

Manuscripts should be written double-spaced, in 12-
point font and submitted to either of the two editors as
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Title, author line, main text, all headings, legends, etc.
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cepted.

Literature citations: In the text, if there are three or more
authors only the first author is cited (Schulze et al. 1969),
citations are separated by comma (Koponen 2000, 2008,
Huber 2010). In the literature cited the citations are ar-
ranged alphabetically by author and journal names are not
abbreviated. Papers with the same author(s) and years are
identified by a, b, c, etc.

Examples:

Broen B von 1993 Nachweise selten gefundener oder gefährdeter
Spinnen (Araneae) in der Mark Brandenburg. - Arachnolo-
gische Mitteilungen 6: 12-25

Chatzaki M 2008 Ä critical review of the spider family Gna-
phosidae in Greece. In: Makarov SE & Dimitrijevic RN
(eds.) Advances in arachnology and developmental biology.
Pensoft, Sofia, pp. 355-374

Clayhills T, Rinne V & Koponen S 2008 On insect and spider
fauna of Jungfruskär in Houtskär, 2007. Unpublished report
to the Finnish Forest and Park Service, Turku. 80 pp. (in
Finnish)

Lindgren L 2000 Island pastures. Metsähallitus and Edita Ltd.
Helsinki. 203 pp.

Merkens S 2000 Die Spinnenzönosen der Sandtrockenrasen im
norddeutschen Tiefland im West-Ost-Transekt - Gemein-
schaftsstruktur, Habitatbindung, Biogeographie. Dissertation,
Univ. Osnabrück. 165 pp.

Nentwig W, Blick T, Gloor D, Hänggi A Sc Kropf C 2013 ara-
neae — Spiders of Europe, version 1.2013. - Internet: http://
www.araneae.unibe.ch (4.1.2013)

Platnick NI 2012 The world spider catalog, version 13.0. - Inter-
net: http://research.amnh.org/iz/spiders/catalog (15.9.2012)

Doi-Links should be given. Example:

Kovblyuk MM Sc Marusik YM 2012 First description of the
female of the theridiid spider Robertus golovatchi (Araneae:
Theridiidae). - Arachnologische Mitteilungen 44: 17-19 -
doi: 10.543 l/aramit4405

Structure: Following a concise title, the next line is the
author(s) full name(s) (Prename Surname), the last au-
thor linked by Sc. After this comes an Abstract. . . . briefly
summarising the main results. Next come a few precise
Keywords:..., sorted alphabetically, which supplement
the title and abstract (for the purposes of bibliographic
databasing). A short title should be provided. The text is
to be arranged in an introductory chapter (without title).
Material and methods (or Study area. Methods, etc.).
Results and Discussion. An Acloiowledgements comes
before the References. For the authors address block give
the full name (Prename SURNAME), address(es) and e-
mail.

Advice for electronic formats

The manuscript should be submitted by e-mail in a Micro-
soft-compatible format, preferably as a WORD- (name,
doc) or a Rich-Text- (name.rtf) document. All MS Of-
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as JPG-files (e.g. originals from the digital camera). They
wiU be arranged according to their legends along with all
other figures submitted with the manuscript. Alternatively
the author(s) can submit ready-formatted digital figures.
However, in such cases authors should consult the editors
with respect to file formats, etc.

Arachnologische

Mitteilungen

Volume 45

Contents

SMITHSONIAN LIBRARIES

Karlsruhe, June 2013

Ma^az Kuntner: Preface to the section of the Proceedings of the 27th European Congress of

Arachnology, Ljubljana, 2012 Sept. 2-7 (pp. 4-20) 1-3

Christo Deltshev: On the identity of the poorly known spider species Zelotes strandi (Araneae:

Gnaphosidae) 4-7

Seppo Koponen: Ground-living spiders in wooded habitats under human influence on an island in

Finland 8-14

Boris Zakharov & Vladimir Ovtcharenko: Male palp organ morphology of three species of ground

spiders (Araneae, Gnaphosidae) 15-20

Dieter Martin: Nachweise für Mecklenburg-Vorpommern neuer und seltener Spinnenarten

(Arachnida, Araneae)

Records of new and rare spider species from Mecklenburg-Western Pomerania (Arachnida,

Araneae) 21-24

Jana Christophoryova, Anna Sestakovä, Miroslav Krumpäl & Peter Fend’a: First record of a

schizomid, Stenochrus portoricensis (Schizornida: Hubbardiidae), in Slovakia 25-29

Petr Dolejs: Do really all wolf spiders carry spiderlings on their opisthosomas? The case of Hygro-

lycosa rubrofasciata (Araneae: Lycosidae) 30-35

Antje Deepen-Wieczorek & Axel L. Schönhofer: Bestätigung von Homalenotus quadridentatus

(Opiliones: Sclerosomatidae) für die Fauna Deutschlands

Confirmation of Homalenotus quadridentatus (Opiliones: Sclerosomatidae) for the fauna of
Germany 36-39

Anna Sestakovä, Jana Christophoryova & Stanislav Korenko: A tropical invader, Coleo-
soma floridanum^ spotted for the first time in Slovakia and the Czech Republic (Araneae,
Theridiidae) 40-49

Sascha Buchholz: Spider records from East Macedonia and Thrace (NE Greece) 45-53

ISSN 1018-4171

www.AraGes.de