Content uploaded by Kirill Glebovich Mikhailov
Author content
All content in this area was uploaded by Kirill Glebovich Mikhailov
Content may be subject to copyright.
259
EuropEan arachnology 2005 (Deltshev, C. & Stoev, P., eds)
Acta zoologica bulgarica, Suppl. No. 1: pp. 259-268.
1
Institute for Biological Problems of the North, Portovaya Str. 18, Magadan 685000, Russia.
E-mail: yurmar@mail.ru
2
Zoological Museum, Moscow State University, B. Nikitskaya Str. 6, Moscow 125009, Russia.
E-mail: kmk2000@online.ru
3
Institute of Zoology, block 504, passage 1128, Baku 370073, Azerbaijan. E-mail: hun-vey-bin@rambler.ru
Advance in the study of biodiversity of Caucasian spiders
(Araneae)
Yuri M. Marusik
1
, Kirill G. Mikhailov
2
, Elchin F. Guseinov
3
Abstract: The history of investigation of the Caucasian areaneofauna can be divided into four periods:
1866-1938, 1939-1978, 1979-1998 and 1999 to the present. According to published data, over 1000 species
belonging to 46 families are known from the Caucasus. The species richest families are as follows: Linyphi-
idae (~180), Salticidae (122), Gnaphosidae (>100), Lycosidae (>100), Theridiidae (80), Dysderidae (70)
and Thomisidae (70). In the different families endemism values vary from 0 to 100%. The average level of
endemism in the Caucasus is about 22%, the highest level of endemism among species-rich families was
found in the Dysderidae, being around 60%.
Key words: spiders, Caucasus, fauna, zoogeography, endemism
Introduction
The Caucasus is a territory lying between the Black and the Caspian seas. There are no strict
geographical borders that separate the mountain from the areas lying to the north and south.
Conventionally, the northern border of the Caucasus coincides with the northern foothills of the
Caucasus Major, and the southern border is formed by the southern borders of Georgia, Armenia
and Azerbaijan (Fig. 1).
The study of Caucasian arachnids was initiated by L. Koch (1866) who described the
gnaphosid Melanophora caucasia (= Zelotes c.) from this region. The history of the study of
Caucasian spiders can be conveniently divided into 4 periods: 1) beginning (1866-1938); 2) pre-
Dunin (1939-1978); 3) Dunin (1979-1998); and 4) modern (1999 to the present). In the rst period
the greatest contribution to the knowledge of Caucasian spiders was made by the Russian and
foreign authors: A.I. Kroneberg, L. Koch, W. Kulczyński, E. Simon, T. Thorell, E. Werzbitski and
A.M. Zavadski (Fig. 2). During this period many new species were described or recorded from
the area. According to charitonov’s (1932) catalogue, 178 species of spiders were known from
the Caucasus in 1926. A decade later, the number of Caucasian species reached 206 (charitonov
1936). It is worth mentioning that all these arachnologists lived far from the Caucasus.
We date the beginning of the second period from the series of papers published by chari-
tonov which dealt with the cave fauna of the region (charitonov 1939, 1941a, b) (Fig. 3). At the
same time, a Georgian arachnologist, Tamara S. Mkheidze began her career. During the second
period important contributions to the study of Caucasian arachnids were made by T.S. Mkheidze
(Tbilisi), S.A. Spassky (Novocherkassk), D.E. Charitonov, A.S. Utotchkin (Perm), V.E. Pichka
(Kiev) and several other authors (Fig. 3). During this period the rise of knowledge of the taxonomy
and faunistics of Caucasian spiders was somewhat slow.
260
EuropEan arachnology 2005
The third period began when Peter M. Dunin started working at the Institute of Zoology
of Azerbaijan, Baku (Fig. 3). The time of Dunin’s employment coincides with the activation of
arachnological studies in the entire Soviet Union. During this period many young arachnologists
such as A.A. Zyuzin, V.I. Ovtsharenko, A.V. Ponomarev, K.Yu. Eskov, A.V. Tanasevitch, K.G.
Mikhailov, Yu.M. Marusik and D.V. Logunov started to study spiders, including those from the
Caucasian region. This period was characterized by a great growth in the number of described and
reported species. We name this period after Dunin because he made the most important contribu-
tion, publishing over 30 papers on Caucasian spiders and describing over 60 species from the area.
Fig. 1. Conventional borders of Caucasus.
Fig. 2. Portraits of the rst generation of arachnologists, who had taken part in the study of the Caucasian fauna.
261
Yu. Marusik et al.: Study of Caucasian spiders
Besides Dunin, an important contribution was made by A.V. Tanasevitch in his revisional studies
of Caucasian linyphiids (tanasEvitch 1987, 1990). He described about 40 species and reported
over 100 species from the Caucasus. Besides this, Tanasevitch revised the Central Asian fauna,
and many species from Central Asia were later found in the Caucasus. After the collapse of the
Soviet Union, the study of Caucasian arachnids nearly ceased because of military conicts, lack
of nancing and other reasons. During this period most studies were based on museum materials
collected earlier. Most of the works from this period were published by Ovtsharenko and co-au-
thors (ovtsharEnKo et al. 1992, 1994, 1995).
In the fourth, or modern period, arachnological studies intensied in Azerbaijan. During
this period papers were published by E.F. Guseinov (=Huseynov), D.V. Logunov, Yu.M. Marusik,
K.G. Mikhailov, S. Koponen, P.T. Lehtinen, G.N. Azarkina, M.M. Kovblyuk and several other
authors. Although the number of species described was not high, this period was marked by the
large number of new supraspecic taxa (4 families, about 25 genera) reported from the Caucasus
for the rst time (MarusiK, gusEinov 2003, MarusiK et al. 2005). During all periods of investiga-
tion of Caucasian spiders there were taxonomical, faunistic and mixed papers. Faunistic papers
published during the 2
nd
and 3
rd
periods have many deciencies: 1) many species, genera and even
families were incorrectly identied; 2) many descriptions and redescriptions were inadequate, and
it is impossible to identify spiders based on these papers. Inaccuracy of identications during this
period was related to the lack of appropriate literature in Azerbaijanian and Georgian libraries,
the lack of revisional studies and the lack of access to comparative material stored in Moscow,
St. Petersburg and abroad.
Difculties in the study of Caucasian spiders
There are several difculties in the study of Caucasian spiders. They are related to the following
factors: 1) lack of access to the types described by Mkheidze in 1940-1990 (more than 30 species);
2) lack of the types of species described by L. Koch (9 species) and V. Kulczyński; 3) the materials
on which faunistic papers by Mkheidze, Kulczyński, Werzbitski were based are not accessible
or were lost during World War II. Some materials collected by Guseinov were also lost. Until
recently, the study of Caucasian spiders was hampered by the lack of revisions and redescriptions
of old materials from adjacent areas like Turkey, Asia Minor and Near East, Bulgaria, Greece and
the Crimea. Many new species were described from these areas at the end of 19
th
century and the
Fig. 3. Portraits of the second and third generations of arachnologists, who had taken part in the study of the
Caucasian fauna.
262
EuropEan arachnology 2005
beginning of 20
th
. Most of these species had long been known only from the original publications.
Signicant progress in the study of Caucasian spiders was achieved because of revisions of vari-
ous families from Israel made by G. lEvy (1985, 1986, 1987, 1991, 1992, 1995, 1996, etc.). Levy
revised and redescribed many spider species previously described from the eastern Mediterranean
and northern Africa. Besides Levy, large contributions were made by K. Thaler and B. Knoach
(Knoflach 1996, 1999, Knoflach, thalEr 2000, thalEr, Knoflach 1998, thalEr et al. 2000, etc.)
on the Theridiidae and several other groups from southern Europe. Near the end of 20
th
century,
the spiders of the family Salticidae were almost completely revised in Central Asia and partially
in the Caucasus by Logunov and his co-authors (logunov 1996, 1999 a, b, logunov, MarusiK
1999, 2003, logunov et al. 1999, raKov, logunov 1996, etc.)
Recent state of knowledge
According to MiKhailov’s (1997) catalogue, 886 species of spiders were known from the Cau-
casus in 1997. In the following years over 100 additional species were reported from Georgia
(MKhEiDzE 1997) and Azerbaijan. Of the Transcaucasian regions, the most thoroughly studied
country is Azerbaijan with over 600 species (MarusiK, gusEinov 2003). In Georgia, 456 species
of spiders are known to date, and only 127 species from Armenia (MiKhailov 2002). We do not
have exact data about the number of species from the Russian Caucasus.
Studies conducted by us in Azerbaijan after 2001 revealed that the fauna of the Caucasus was
inadequately known. During a short-term expedition to the Absheron Peninsula and Lenkoran we
found 16 genera and 4 families (Desidae, Mysmenidae, Palpimanidae and Prodidomidae) new to
the Caucasus as a whole, including 5 genera new to the fauna of the former Soviet Union (MarusiK,
gusEinov 2003). Subsequent expeditions to Nakhchivan and other parts of Azerbaijan revealed
several additional genera new to Azerbaijan, the whole of the Caucasus and even the former USSR,
e.g. Siwa grasshoff, 1970 (MarusiK et al. 2004). The number of new species reported for Azer-
baijan, the Caucasus, and all of the former Soviet Union is several dozen. It is worth mentioning
that the species new to the Caucasus were found not only among poorly studied families such as
the Lycosidae and Gnaphosidae, but also in the well studied Araneidae. For example, Cyclosa
sierrae siMon, 1870, Singa neta (O. P.-caMbriDgE, 1872) and Siwa atomaria (O. P.-caMbriDgE,
1876) found in Nakhchivan (MarusiK et al. 2005b) were new to the former Soviet Union. Two
days of collecting in Sukhum, Abkhazia, and several hours of collecting in Adler (near Sochi)
revealed a family new to the Caucasus (Zoropsidae) (MarusiK, KovblyuK 2004), and two families
new to the European part of Russia (Mysmenidae, Oonopidae) (MarusiK 2005).
Because of collecting efforts during the last 5 years, the spider fauna of Azerbaijan, with
44 families, became the most family-rich of all the regions of the former Soviet Union. The total
number of families known from the Caucasus is now 46. Two of them, Cybaeidae and Zorop-
sidae, have not yet been found in Azerbaijan. There is no doubt that the diversity of families in
the Caucasus, and particularly in Azerbaijan, will be increased. The presence of representatives
of the Anapidae, Cithaeronidae, Hersiliidae, Phyxelididae, Sicariidae and Synaphridae is likely.
Comaroma simony bErtKau, 1889, belonging to the rst mentioned family, is common in Europe
in thick leaf litter. Cithaeronids, hersiliids and sicariids are known from adjacent Turkmenistan and
Iran. Phyxelidids are known from Turkey and synaphrids have been reported from Turkmenistan,
the Crimea and several Mediterranean countries (MarusiK et al. 2005b). It is worth mentioning
that most of the undescribed and newly reported taxa were found not in mountains, or other poorly
accessible areas, but in coastal lowlands: the Absheron Peninsula, Lenkoran and Sukhum. These
areas have a semi-arid or subtropical climate.
The thoroughness with which the various Caucasian spider families have been studied dif-
fers greatly. Among species-rich families the best studied ones are the Clubionidae, Dysderidae,
Linyphiidae and Salticidae. Several special publications are devoted to these families. The least
263
Yu. Marusik et al.: Study of Caucasian spiders
studied families are the Agelenidae, Dictynidae, Gnaphosidae, Lycosidae, Philodromidae, Theri-
diidae and Thomisidae. The study of the Azerbaijanian Agelenidae reveals that, among 19 species
found in the republic, 14 are new to the science and one is new to the Caucasus (gusEinov et al.
2005). A very high number of new taxa were found in the Gnaphosidae. Many species found in
Azerbaijan belong to genera unknown to us. Among other families from Azerbaijan, such as the
Lycosidae, Dictynidae, Thomisidae and Theridiidae, the proportion of new taxa is lower than in
the Agelenidae. For instance, the percentage of new species among the theridiids is about 20%.
Although the Linyphiidae is the most species-rich family and one of the best studied in the re-
gion, we recognized, among new material from Azerbaijan, several species new to the science
or to Azerbaijan. New species were found also among other well studied families such as the
Dysderidae.
The species diversity of all families represented in the Caucasus is summarized in Table
1. According to the literature and unpublished personal data the fauna of the region includes at
least 970 species. For some of the families we provide estimated data, which is slightly higher
than the number of reported species.
Table 1. Number of species in each family found in the Caucasus, number and percentage of endemic spe-
cies. * evaluation data; ** % from reported/ known species.
Family Number of species Number of endemics %**
1. Agelenidae 35 18 51
2. Amaurobiidae 5 1 20
3. Anyphaenidae 2 0 0
4. Araneidae 48 0 0
5. Argyronetidae 1 0 0
6. Atypidae 1 0 0
7. Clubionidae 29 4 14
8. Cheiracanthidae 8 0 0
9. Cybaeidae 2 1 50
10. Desidae 1 1 100
11. Dictynidae 18 1 6
12. Dysderidae* 70 >59 91
13. Eresidae 3 1 33
14. Filistatidae 3 1 33
15. Gnaphosidae* 100 >10 >12
16. Hahniidae 6 1 17
17. Heteropodidae 2 0 0
18. Leptonetidae 2 2 100
19. Linyphiidae* 180 >45 >27
20. Liocranidae 6 3 75
21. Corinnidae 6 3 50
22. Lycosidae* 100 16 >20
23. Mimetidae 3 1 50
24. Mysmenidae 2 0 0
25. Nemesiidae 4 4 100
26. Nesticidae 9 7 78
27. Oecobiidae 6 0 0
264
EuropEan arachnology 2005
Family Number of species Number of endemics %**
28. Oonopidae 5 3 60
29. Oxyopidae 4 0 0
30. Palpimanidae 1 1 100
31. Philodromidae* 30 1 >4
32. Pholcidae 6 2 33
33. Pisauridae 3 0 0
34. Prodidomidae 2 1 50
35. Salticidae 122 17 14
36. Scytodidae 1 0 0
37. Segestriidae 3 0 0
38. Tetragnathidae 18 0 0
39. Theridiidae 80 6 8
40. Theridiosomatidae 1 0 0
41. Thomisidae* 70 >10 >15
42. Titanoecidae 7 1 14
43. Uloboridae 6 0 0
44. Zodariidae 7 5 71
45. Zoridae 4 0 0
46. Zoropsidae 1 0 0
1022 >226 >22
The most diverse spider families in Caucasus are as follows: Linyphiidae (~180), Salticidae
(122), Gnaphosidae (>100), Lycosidae (>100), Theridiidae (80), Dysderidae (70) and Thomis-
idae (70). The spider fauna of the Caucasus comprises about 1/3 of the species diversity of the
former Soviet Union and about 2.5% of the world species diversity. In terms of family diversity,
the fauna of Caucasus encompasses 42% of the families of the world. The value of endemism
in different families ranges from 0 to 100%. The highest level of endemism was found not only
in the families with few species like the Nemesiidae, Leptonetidae or Desidae, but also in the
species-rich families like the Dysderidae. In total, the level of endemism in the Caucasian fauna
is not less than 22%, and probably this value will increase when several families like the Gna-
phosidae, Philodromidae and Linyphiidae are properly revised. We think that the largest growth
in species, new to the science and new to the region, will be in such families as the Gnaphosidae,
Lycosidae, and Linyphiidae. There is no doubt that in Armenia and Georgia numerous new spe-
cies of Agelenidae and Dysderidae will be found as well.
The most promising areas for nding species new to the fauna of the Caucasus are the arid
regions of Azerbaijan, Georgia and Armenia, the subtropical coasts of the south-east, the whole
western Caucasus and the high mountains. High altitudes are promising only in terms of new spe-
cies with limited ranges, but subtropical and arid territories should produce many new or newly
reported supraspecic taxa. At present we are working in collaboration with colleagues from different
countries on revisions of the Corinnidae, Liocranidae, Lycosidae, Oonopidae, Philodromidae and
Theridiidae. In their general species diversity, spiders of the Caucasus are similar to those of oribatid
mites. According to tarba (2002) the number of oribatids in the Caucasus reaches 770 species. This
number comprises 10% of the world species diversity (2.5% in spiders). Possibly, the high value
of species diversity of mites in the Caucasus reects a poor level of study of Oribatida in the rest
of the world. The value of endemics among spiders (22%) and oribatids (17%) is very similar.
Table 1. Continued.
265
Yu. Marusik et al.: Study of Caucasian spiders
Caucasus – Far East disjunctions
One of the most interesting and unique characteristics of the Caucasian spider fauna is the presence
of about a dozen species with Caucasus-Far East disjunctions. When we began to study spiders
of this area in 1980 we were faced with ve species (Octonoba yesoensis, Phintella castriesiana,
Myrmarachne formicaria, Rhomphea sagana and Tmarus horvathi) that were known in Azerbaijan
and/or Georgia and in the Far East (MarusiK 1987, logunov, MarusiK 1991). Now, the number
of such species has increased to 9 (MarusiK et al. 2004) due to the recent discovery of Caucasian
species in the Far East (Rhomphea hyrcana, Larinia bonneti) and of Far Eastern species in Cau-
casus (Yaginumena maculosa, Howaia mogera).
Why do we call this situation unique? It is because such long disjunctions are unknown in
other groups of animals. We were able to nd one species of feather beetle, one species of saw-y
and one terrestrial mollusc with the same disjunction pattern. All these groups are taxonomically
difcult in comparison to those of the spider species with disjunctions. We asked several experts
in species-diverse groups such as Curculiniodae, Carabidae, Rhopalocera, Arctiidae, Noctuidae,
Heteroptera, but none of them knew of disjunctive ranges at the species level. Although they are
more common in spiders, supraspecic disjunctions are also known in insects. The high percent-
age of spider species with disjunctive ranges indicates that evolution, or at least morphological
evolution, occurs more slowly in them than in other arthropods.
Ethological studies
Despite the almost 140 years of faunistic and taxonomic studies of the Caucasian araneofauna,
investigation of the biology of spiders in this region began only recently. It started with a paper
by gusEinov (1997), who gave preliminary information on the natural prey of some species of
wandering spiders occurring in the Absheron Peninsula, Azerbaijan. Later, the diets of 20 species
from six families (Salticidae, Thomisidae, Philodromidae, Oxyopidae, Gnaphosidae, Filistatidae)
were studied in detail, and this formed an important part of the Ph.D. thesis of the third author (gu-
sEinov 1999). Some of these results are already published in a series of separate papers (gusEinov
2004a, b, 2005), and some are in press. Moreover, in cooperation with Robert Jackson (Canterbury,
New Zealand) and his students, some ethological aspects, such as predatory behaviour and prey
preference, of a few Azerbaijanian jumping spiders have been investigated (cErvEira et al. 2003,
gusEinov et al. 2004). In addition to these studies on the natural prey and predatory behaviour of
spiders in Azerbaijan, investigation of their microhabitat preferences is also in progress.
Acknowledgements: We wish to thank Seppo Koponen for checking the draft of this work and making
valuable comments. The English of the nal manuscript was kindly checked by D.J. Buckle. This work was
supported partly by the RFBR grant (No. 04-04-48727) and by the Academy of Finland (No. 211596).
Received: 30.11.2005
Accepted: 04.04.2006
References
cErvEira A. M., R. R. JacKson, E. F. gusEinov 2003. Stalking decisions of web-invading araneophagic jumping
spiders from Australia, Azerbaijan, Israel, Kenya, Portugal, and Sri Lanka: opportunistic smokescreen
tactics of Brettus, Cocalus, Cyrba, and Portia. - New Zealand Journal of Zoology, 30: 21-30.
c
haritonov D. E. 1932. Katalog der russischen Spinnen. - Yezhegodnik Zoololocheskogo Muzeya Akademii
Nauk SSSR, 32: 1-207.
c
haritonov D. E. 1936. Addendum to the catalogue of Russian spiders. - Uchenyye zapiski Permskogo
universiteta, 2 (1): 167-225. (In Russian)
c
haritonov D. E. 1939. On the cave spiders of Abkhasia. - In: Materialy k faune Abkhazii. Academie des
Sciences l’URSS, Filiale Georgienne - Section de Zoologie: 197-210. (In Russian)
266
EuropEan arachnology 2005
charitonov D. E. 1941a. On the cave fauna of Caucasus. - Bulletin de l’Institut des recherches Biologiques
(Molotov), 12: 67-71. (In Russian)
c
haritonov D. E. 1941b. New materials on the Arachnoidea of Abkhasian caves. - Travaux de l’Institute de
Zoologie, Academie des Sciences l’URSS, Filiale Georgienne, 4: 165-176. (In Russian)
g
usEinov E. F. 1997. Preliminary data on prey composition of some wandering spider species (Araneae)
inhabiting Apsheron Peninsula, Azerbaijan. - Deposited in AzNIINTI, № 2500-Az, 5 p. (In Russian)
g
usEinov E. F. 1999. Spiders of Lenkoran nature area and Absheron Peninsula, Azerbaijan. Ph.D. thesis,
Institute of Zoology, Baku. (In Russian)
g
usEinov E. F. 2004a. Prey composition of three Thanatus species (Philodromidae, Araneae): indication of
relationship between psammophily and myrmecophagy. - In: saMu F., Cs. szinEtár (eds): European
Arachnology 2002. Proceedings of the 20th European Colloquium of Arachnology, Szombathely,
22-26 July 2002: 103-108.
g
usEinov E. F. 2004b. Natural prey of the jumping spider Menemerus semilimbatus (Hahn, 1827) (Araneae:
Salticidae), with notes on its unusual predatory behaviour. - In: logunov D. V., D. pEnnEy (eds):
European Arachnology 2003. Proceedings of the 21st European Colloquium of Arachnology, St.-
Petersburg, 4-9 August 2003: 93-100.
g
usEinov E. F. 2005. Natural prey of the jumping spider Salticus tricinctus (Araneae: Salticidae). - Bulletin
of the British Arachnological Society, 13: 130-132.
g
usEinov E. F., A. M. cErvEira, R. R. JacKson 2004. The predatory strategy, natural diet, and life cycle
of Cyrba algerina, an araneophagic jumping spider (Salticidae: Sparaeinae) from Azerbaijan. - New
Zealand Journal of Zoology, 31: 291-303.
K
noflach B. 1996. Die Arten der Steatoda phalerata-Gruppe in Europa (Arachnida: Araneae, Theridiidae). -
Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 69: 377-404.
K
noflach B. 1999. The comb-footed spider genera Neottiura and Coleosoma in Europe (Araneae, Theridi-
idae). - Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 72: 341-371.
K
noflach B., K. thalEr 2000. Notes on Mediterranean Theridiidae (Araneae) - I. - Memorie della Società
Entomologica Italiana, 78: 411-442.
K
och L. 1866. Die Arachniden-Familie der Drassiden. Heft 6. Nurnberg, Bauer und Raspe, 257-304.
l
Evy G. 1985. Araneae: Thomisidae. - In: Fauna Palaestina, Arachnida II. Israel Academy of Sciences and
Humanities, Jerusalem, vi+114 p.
l
Evy G. 1986. Spiders of the genera Siwa, Larinia, Lipocrea and Drexelia (Araneae: Araneidae) from
Israel. - Bulletin of the British Arachnological Society, 7: 1-10.
l
Evy G. 1987. Spiders of the genera Araniella, Zygiella, Zilla and Mangora (Araneae, Araneidae) from
Israel, with notes on Metellina species from Lebanon. - Zoologica Scripta, 16: 243-257.
l
Evy G. 1991. On some new and uncommon spiders from Israel (Araneae). - Bulletin of the British Arach-
nological Society, 8: 227-232.
l
Evy G. 1992. The spider genera Palaestina, Trygetus, Zodarion and Ranops (Araneae, Zodariidae) in Israel
with annotations on species of the Middle East. - Israel Journal of Zoology, 38: 67-110.
l
Evy G. 1995. Revision of the spider subfamily Gnaphosinae in Israel (Araneae: Gnaphosidae). - Journal
of Natural History, 29: 919-981.
l
Evy G. 1996. The agelenid funnel-weaver family and the spider genus Cedicus in Israel (Araneae, Agel-
enidae and Cybaeidae). - Zoologica Scripta, 25: 85-122.
l
ogunov D. V. 1996. A review of the genus Phlegra Simon, 1876 in the fauna of Russia and adjacent coun-
tries (Araneae: Salticidae: Aelurillinae). - Genus, 7: 533-567.
l
ogunov D. V. 1999a. Redenition of the genus Habrocestoides Prószyński, 1992, with establishment of a
new genus, Chinattus gen. n. (Araneae: Salticidae). - Bulletin of the British Arachnological Society,
11: 139-149.
l
ogunov D. V. 1999b. Two new jumping spider species from the Caucasus (Aranei: Salticidae). - Arthropoda
Selecta, 7: 301-303.
l
ogunov D. V., Yu. M. MarusiK 1990. The spider genus Argyrodes (Aranei, Theridiidae) in the USSR. -
Zoologicheskiy zhurnal, 69 (2): 133-136. (In Russian)
l
ogunov D. V., Yu. M. MarusiK 1999. A brief review of the genus Chalcoscirtus Bertkau, 1880 in the faunas
of Central Asia and the Caucasus (Aranei: Salticidae). - Arthropoda Selecta, 7: 205-226.
267
Yu. Marusik et al.: Study of Caucasian spiders
logunov D. V., Yu. M. MarusiK 2003. A revision of the genus Yllenus Simon, 1868 (Arachnida, Araneae,
Salticidae). Moscow, KMK Scientic Press, 168 p.
l
ogunov D. V., yu. M., MarusiK, s. y. raKov 1999. A review of the genus Pellenes in the fauna of Central
Asia and the Caucasus (Araneae, Salticidae). - Journal of Natural History, 33: 89-148.
M
arusiK Yu. M. 1987. Systematics and biology of the orb-weaving spider, Octonoba yesoensis (Aranei, Ulo-
boridae). - Zoologicheskiy zhurnal, 66 (4): 613-616. (In Russian)
M
arusiK Yu. M. 2005. New family and new interesting records of spiders (Aranei) from the European part
of Russia. - Arthropoda Selecta, 14 (1): 89-91.
M
arusiK Yu. M., G. gusEinov 2003. Spiders (Arachnida, Aranei) of Azerbaijan. I. New families and genera
records. - Arthropoda Selecta, 12 (1): 29-46.
M
arusiK Yu. M., M. M. KovblyuK 2004. New and interesting cribellate spiders from Abkhazia (Aranei:
Amaurobiidae, Zoropsidae). - Arthropoda Selecta, 13 (1-2): 55-61.
M
arusiK Yu. M., V. A. gnElitsa, M. M. KovblyuK 2005a. A new species of Synaphris (Araneae, Synaphridae)
from Ukraine. - Bulletin of the British Arachnological Society, 13 (4): 125-130.
M
arusiK Yu. M., F. G. gusEinov, H. A. aliEv 2005b. Spiders (Arachnida, Aranei) of Azerbaijan 4. Fauna of
Nakhchivan. - Arthropoda Selecta, 13 (3): 135-149.
M
arusiK Yu. M., E. F. gusEinov, S. KoponEn, H. yoshiDa 2004. A new case of Caucasus-Far East disjunctive
range in spiders (Araneae). - Acta Arachnologica, 53 (2): 125-129.
M
iKhailov K. G. 1997. Catalogue of the spiders of the territories of the former Soviet Union (Arachnida,
Aranei). Moscow, Zoological Museum of the Moscow State University, 416 p.
M
iKhailov K. G. 2002. The spider fauna of Russia and other post-Soviet republics: a 2000 update. - In: toft
S., N. scharff (eds.), European Arachnology 2000: Proceedings of the 19th European Colloquium of
Arachnology. Aarhus University Press, 255-259.
M
KhEiDzE T. S. 1941. About study of spiders in Georgia. - Proceeding of the Tbilisi State University, 21:
99-104.
M
KhEiDzE T. S. 1997. Spiders of Georgia: Systematics, Ecology, Zoogeographic Review. Tbilisi, Tbilisi
University, 390 p. (In Georgian)
o
vtsharEnKo V. I., N. I. platnicK, D. X. song 1992. A review of the North Asian ground spiders of the
genus Gnaphosa (Araneae, Gnaphosidae). - Bulletin of the American Museum of natural History,
212: 1-88.
o
vtsharEnKo V. I., G. lEvy, N. I. platnicK 1994. A review of the ground spider genus Synaphosus (Araneae,
Gnaphosidae). - American Museum Novitates, 3095: 1-27.
o
vtsharEnKo V. I., N. I. platnicK, Yu. M. MarusiK 1995. A review of the Holarctic ground spider genus
Parasyrisca (Araneae, Gnaphosidae). - American Museum Novitates, 3147: 1-55.
r
aKov S. Y., D. V. logunov 1996. A critical review of the genus Heliophanus C. L. Koch, 1833, of Middle
Asia and the Caucasus (Aranei, Salticidae). - Arthropoda Selecta, 5 (3/4): 67-104.
t
anasEvitch A. V. 1987. The linyphiid spiders of the Caucasus, USSR (Arachnida: Araneae: Linyphiidae). -
Senckenbergiana biologica, 67 (4-6): 297-383.
t
anasEvitch A. V. 1990. The spider family Linyphiidae in the fauna of the Caucasus (Arachnida, Aranei). -
In: striganova B. R. (ed.): Fauna nazemnykh bespozvonochnykh Kavkaza. Moscow, Akademia Nauk,
5-114. (In Russian)
t
arba Z. M. 2002. Taxonomic diversity of oribatids in the Caucasian fauna. - Biological diversity of Cau-
casus. Proc. 2
nd
regional conference, Sukhum: 241-242. (In Russian)
t
halEr K., J. buchar, B. Knoflach 2000. Notes on wolf spiders from Greece (Araneae, Lycosidae). - Linzer
Biologische Beitrage, 32: 1071-1091.
t
halEr K., B. Knoflach 1998. Zoropsis spinimana (Dufour), eine für Österreich neue Adventivart (Ara-
neae, Zoropsidae). - Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck, 85:
173-185.
268
EuropEan arachnology 2005
Възход в изследванията на паяците на Кавказ (Araneae)
Ю. Марусик, К. Михайлов, Е. Гусейнов
(Резюме)
Направен е преглед на проучванията на кавказката аранеофауна, които според авторите
могат да бъдат разделени на четири периода: Начален – от 1866 до 1938 г., преди Дунин – от
1939 до 1978 г., по времето на Дунин – от 1979 до 1998 г., и Съвременен – от 1999 г. досега.
Понастоящем от територията на Кавказ са известни над 1000 вида паяци, принадлежащи
към 46 семейства. Най-богати на видове са: Linyphiidae (около 180), Salticidae (122), Gna-
phosidae (над 100), Lycosidae (над 100), Theridiidae (80), Dysderidae (70) и Thomisidae (70).
Ендемизмът сред семействата варира в широки граници – от 0% до 100%, като средно е
около 22%. Сред семействата с по-голям брой видове най-висок ендемизъм се наблюдава
при Dysderidae – около 60%.
