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Zeitschrift/Journal: Arachnologische Mitteilungen

Jahr/Year: 2011

Band/Volume: 40

Autor(en)/Author(s): Deltschev [Deltshev] Christo

Artikel/Article: The faunistic diversity of cave-dwelling spiders (Arachnida, Araneae) of Greece 23-32 1

© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;

Arachnologische Mitteilungen 40:23-32 Nuremberg, January 201

The faunistic diversity of cave-dwelling spiders (Arachnida, Araneae) of Greece

Christo Deltshev

doi: 1 0.543 1/aramit4004

Abstract: Until today, from Greek caves a total of 1 09 species of spiders belonging to 25 families are known. One species, the linyphiid Porrhomma convexum (Westring, 1861) was recorded here for the first time in Greece. The 109

species are distributed in caves of different geographic territories as follows:Thrace - 8 species, Macedonia - 1 8,

- 1 - Central - Attiki-Saronic Islands - Peloponnese - 1 Evoia-Vories Sporades Epirus , Thessaly 6, Greece 3, 24, 5,

- 1, Eastern Aegean Islands - 5, Cyclades - 3, Dodecanese - 6, Ionian Islands - 23, Crete - 47. The largest fraction of

troglobite species were encountered mainly in the territories of Crete - 15 species (5 of which are anophthalmic),

the Ionian Islands - 4, Thrace - 2 (both anophthalmic), the Attiki-Saronic Islands - 2 (both anophthalmic), the Pe- loponnese - 2 (one anophthalmic), and Macedonia, Thessaly, and the Cyclades - each with 2 species. The richness of the troglobitic spiders in these regions strengthens the assumption that they were major centres of speciation

and evolution for the species of this group. According to their current distribution, the established 1 09 species can be classified into 12 zoogeograpical categories, grouped into 4 complexes (widely distributed, European, Mediterranean, endemics). The largest number of species belong to the endemic complex (53.2 %) and are also the most characteristic and reflect the local character of the cave-dwelling spiders.

Key words: cave-spiders fauna, endemics, troglobites, zoogeography

The earliest data on Greek cave-dwelling spiders were on the southern end of the Balkan Peninsula. The presented by SIMON (1885), KULCZYNSKI (1903), country has borders with Albania, the Republic of ROEWER (1928, 1959), DRENSKY (1936); Kra- Macedonia and Bulgaria to the north, and Turkey to TOCHVIL (1937, 1938), HADJISSARANTOS (1940), the east. The Aegean Sea lies to the east and south and FAGE (1945). More recent publications derive of mainland Greece, while the Ionian Sea lies to the from the investigations of BRIGNOLI (1968, 1971a, west. Both parts of the Eastern Mediterranean basin

1971b, 1972, 1974a, 1974b, 1974c, 1976, 1977, 1978, feature a vast number of islands, islets and rock is-

1979, 1984), Deeleman-Reinhold (1971, 1977, lands (Fig. 1). Two -thirds of the territory of Greece is 1983, 1985, 1989, 1993), DEELEMAN-REINHOLD dominated by limestone, many ofwhich are karstified & DEELEMAN (1988), SENGLET (1971, 2001), (CLENDENON 2009). Deltshev (1979, 1985, 1999, 2000, 2008), Beron The territory of Greece can be divided into 13 (1985, 1986), Beron & Stoev (2004), Thaler & geographical regions (BOSMANS & CHATZAKI 2005; Knoflach (1995), Wunderlich (1995), Bosse- Fig. 1). There are 7 geographical regions on the main- LAERS (1998), BOSSELAERS &HENDERICKX (2002), land: Thrace, Macedonia, Epirus, Thessaly, Central GASPARO (2003, 2004a, 2004b, 2005a, 2005b, 2006, Greece, Attica and the Peloponnese. The Ionian 2007, 2008, 2009), CHATZAKI et al 2002, BOSMANS Islands are situated on the western border of Greece dt CHATZAKI (2005), CHATZAKI &ARNEDO (2006), in the Ionian Sea. There are several island groups in and PLATNICK (2009). The critical incorporation of the Aegean Sea on the eastern side of Greece: Evoia all available literature records and the accumulation of and the Sporades, the Saronic Islands (grouped with new data are now sufficient to allow a critical analysis Attica), the Cyclades, the Eastern Aegean Islands, of the distribution of spiders established in the caves the Dodecanese and Crete (Fig. 1). of Greece. Results Study area and material Species composition

Greece is a country in south-eastern Europe, situated The spiders established in the caves of Greece (Main- land and Insular part) are represented by 109 species,

Christo DELTSHEV, Institute of Biodiversity and Ecosystem included in 52 genera and 25 families: Ctenizidae

Research, 1 Tsar Osvoboditel Blvd; 1 000 Sofia, Bulgaria - 1, Filistatidae - 1, Sicariidae — 1, Scytodidae - 1, E-Mail: [email protected] Leptonetidae - 9, Pholcidae - 10, Segestriidae - 3, - - - submitted: 30. 12.20 1 0, accepted: 9.4.20 1 0; online: 10.1.2011 Dysderidae 12, Oonopidae 1, Mimetidae 1, Ere- © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;

24 C. Deltshev

(6), Dysderidae (3), Nesticidae (2), and Linyphiidae

(1). The genera with the largest number of species

are Tegenaria (8), Lepthyphantes (6), Harpactea (5),

and Histopona (5). The species are distributed in caves belonging to the geographic territories of Greece as follows: Thrace

- 8 species, Macedonia - 18, Epirus - 1, Thessaly - 6,

Central Greece - 3, te Attiki- Saronic Islands - 24,

the Peloponnese - 15, Evoia-Voroies Sporades - 1,

the Eastern Aegean Islands - 5, the Cyclades - 7,

the Dodecanese - 6, the Ionian Islands - 23, Crete

- 47 (Table 1). We also have to emphasise that the

degree of exploration in these territories is not equal: the territories of Evoia- Sporades, the Cyclades, the

Dodecanese and Central Greece are less explored.

Cave-dwelling spiders can be categorised into four ecological groups (SKET 2008):

• troglobites: species limited to a life cycle in caves. Often they show a suite of characters, associated

with their adaptation to subterranean life: loss of LIV1KÖ8 f>ELAQOS pigment, loss of eyes and elongation of appenda- ges.

Figure 1 : Map of different geographical regions in Greece. • eutroglophiles: species which can live their entire

life in caves, but also occur in other environments.

sidae - 1, Uloboridae - 1, Nesticidae - 4, Theridiidae • subtroglophiles: species which utilise caves, but

- 4, Anapidae - 1, Linyphiidae - 16,Tetragnathidae must leave the caves to complete their life cycle.

- 4, Araneidae - 1, Lycosidae - 1, Agelenidae — 21, • trogloxenes: species which occur underground

Amaurobiidae — 4, Gnaphosidae — 6, Philodromidae sporadically.

- 1, Thomisidae - 2, Salticidae - 2 (Table 1). One The largest fraction of troglobite species was encoun-

species is new for the Greek spider fauna: Porrhomma tered mainly in the caves of the territories of Crete

convexum (Westring, 1861) (marked in the list with *), -15 (5 anophthalmic), the Ionian Islands - 4, Thrace a spider with a Holarctic distribution and widespread - 2 (2 anophthalmic), the Attiki- Saronic Islands 2 (2

in European caves. It is also well represented in the anophthalmic), the Peloponnese -2(1 anophthalmic), caves of the Balkan Penisula - and not only estab- and Macedonia, Thessaly and the Cyclades each by 2

lished in the caves of Croatia, Romania and Turkey species (Table 1). All troglobites are endemics for the

(DELTSHEV 2008). The number of species is high and territory of Greece or the Balkan Peninsula.

represents about 13 % of the Greek spiders. This is Very important is the presence of eutroglophiles also evident from a comparison with the number of (35 species), because together with troglobites (29 cave-dwelling spiders recorded from the other coun- species), they can be considered as dependent faunistic

tries of the Balkan Peninsula: Bulgaria - 99, Croatia elements of caves. The largest number of species is - 63, Serbia - 59, Bosnia and Herzegovina - 52, established in the caves of Crete (14 species), the Io- Macedonia - 44, Montenegro - 44, Slovenia - 43, Al- nian Islands (11 species), the Attiki- Saronic Islands (9

bania - 10, Turkey - 8, and Romania - 4 (DELTSHEV species), Macedonia (8 species), and the Peloponnese 2008). The established number of species, however, (8 species). Here, the endemics are represented by depends not only on the size of the regions, but also 14 species (35 %). A present day example of active on the degree of exploration. The most characteristic subterranean colonisation and cave penetration are are the families: Leptonetidae (8.2 %), Pholcidae the species Lepthyphantes centromeroides and Pallidup- (9.2 %), Dysderidae (11 %), Linyphiidae (14.6 %), hantes spelaeorum, widespread in the Balkan Peninsula and Agelenidae (19.2 %). The families with largest (DEELEMAN-REINHOLD 1978). Here, the species number of anophthalmic species are Leptonetidae Palliduphantes istrianus should also be included. 1 ;

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Cove-dwelling spiders of Greece 25

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Cave-dwelling spiders of Greece 27

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28 C. Deltshev

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Cave-dwelling spiders of Greece 29

Table 2: Zoogeographical composition of cave dwelling spiders in Greece

Complexes Chorotypes Species

Classification Code Number %

Cosmopolitian COS 7 6.4 Holarctic HOL 5 4.6 Widely distributed Palearctic PAL 9 8.2 West Palearctic WP 5 4.6 European- Central Asiatic ECA 1 0.9 Total 27 24.7 European E 8 7.3 Middle-Southeast European MSEE 1 0.9 European South East European SEE 1 0.9 Total 10 9.2 Mediterranean-Central Asiatic MCA 1 0.9 Mediterranean M 5 4.62 Mediterranean North East Mediteranean NEM 8 7.3 Total 14 12.8 Endemic for continental and island Greece EGR 55 50.4 Endemic for Balkan Penisnula EBP 3 2.7 Endemics Total 58 53.2

The group of subtroglophiles comprises 11 species humid Tertiary forests (DEELEMAN-REINHOLD occurring in dark places such as buildings, screes, rock 1977). However, due to a lack of data, it is difficult to crevices and caves. They can be considered as regular determine with certainty which cave spider endemics inhabitants of the caves. Three species are endemics of the Balkans are Tertiary, and which are Quaternary,

(Maimuna cretica Hoplppholcus minous and Tegenaria elements. , , labyrinthi). The complex of widely distributed species com- The trogloxenes are represented by 36 species. prises 36 species (29.2 %). Palearctic species are They are not truly cavernicolous faunistic elements, dominant (47.2 %), followed by Cosmopolitan (19.4 but their presence in caves should not be ignored. %) Holarctic (13.8 %), West Palearctic (13.8 %), and On the other hand some of them will probably be European-Central Asiatic (5.5 %). The complex considered subtroglophiles and even eutroglophiles includes mostly widespread species associated with once more information on their ecology is gathered. lowlands, buildings, caves, woodlands and high al- Here, the endemics are represented by 11 species. titude zones of mountains. Here characteristic for caves are Nesticus cellulanus Porrhomma convexum , Zoogeographical analysis and Tegenaria domestica. These species are also largely According to their current distribution, the established 109 species can be classified into 12 zoogeograpical categories, grouped into four complexes (widely dis- tributed, European, Mediterranean, endemics) (Tab.

1&2, Fig. 2).

Best represented is the complex of endemics with 58 species (53.2 %), which include 29 troglobites, 16 troglophiles, and 12 trogloxenes. The established number is high and reflects the local character of the cave fauna. The endemics are best represented in Crete - species 26 and 2 genera (.Minotauria, Rhodera), the Ionian Islands - 9 species, the Peloponnese - 8 species, Macedonia and the Eastern Aegean Islands - each Figure 2: Relative representation of the zoogeographical categories of cave dwelling spiders in Greece (for abbre- with 4 species. The recent cave spider fauna is formed viations see Table 1). after gradual changes in the fauna of the ancient © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; 30 C Deltshev widespread taxa in European caves where Porrhomma Acknowledgements convexum occurs in the deep parts. We are especially grateful to our colleagues P. Beron, E The European complex includes 10 species (9.2 Gasparo, S. Lazarov, M. Naumova, K. Paragamian, B. Pertrov, and P. Stoev for the collected material and valuable %). European species are dominant (8 %) among discussions. The research was also supported by the Project this complex. The Middle Southeast European and - Nos. B-1523/05 and BG051P0001-3.3.04/41. Southeast European species are represented by single species - trogloxenes. The complex comprises wide- References spread spiders in Europe and the Balkan Peninsula BERON P. (1985): On the cave fauna of the Greek islands which inhabit both lowlands and mountains. The of Santorin and Iraklia, with preliminary description most characteristic are Meta menardi and Metellina of a new pseudoscorpion. - Grottes Bulgares, Sofia 3: merianae, both widespread in European caves. 64-71 The Mediterranean complex includes 14 species BERON P. (1986): Results of the studies of the cave fauna of real representation this category is (12.8 %). The of Greece. - Biologia Gallo-Hellenica 12: 125-131 probably higher because part ofthe endemic fauna has BERON P. (2001): Etude comparative des faunes caverni- origin. are a Mediterranean Dominant here the spe- coles de la Bulgarie et de la Grece. - Historia naturalis cies widespread in the Mediterranean region or part bulgarica 13: 53-68

it, the real cavernicolous elements are Nesticus of but Beron P, B. Petrov & P. STOEV (2004): The inverte- eremita and Zangherella apuliae (characteristic mainly brate cave fauna of the Eastern Rhodopes (Bulgaria and for the superficial underground compartment) distrib- Greece). In: BERON P. & A. POPOV (eds): Biodiversity of uted in the caves of the north-west part of Greece. Bulgaria. 2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece). Pensoft & National Museum of of Natural Conclusions History. Sofia, pp. 791-822 BOSSELAERS (1998): Nesticus henderickxi (Araneae, Nesti- • The faunistic diversity of the reported 109 cave- J. cidae), a new anophthalmic troglobitic spider from Crete. dwelling spiders shows that Greek caves are - Bulletin of British Arachnological Society 11: 9-14 characterised by a considerable species richness.

BOSSELAERS J. &H. HENDERICKX (2002): A new Savignia This is also supported by comparing the number from Cretan caves (Araneae: Linyphiidae). - Zootaxa of cave-dwelling spiders recorded from other 109: 1-8 countries in the Balkan Peninsula: Bulgaria — 99, BOSMANS R. &M. CHATZAKI (2005): A catalogue of the Croatia - 63, Serbia - 59, Bosnia and Herzegovina spiders of Greece. A critical review of spider species - Macedonia - Montenegro - Slovenia 52, 44, 44, cited from Greece with their localities. - Newsletter - - - - 43, Albania 10, Turkey 8, and Romania 4 of the Belgian Arachnological Society 20 (Supplement

(Deltshev 2008). 2): 1-224

• The uneven species richness in the caves of different BRIGNOLI PM. (1968): Über griechische Leptonetidae regions of Greece is probably due mainly to the (Arachnida: Araneae). - Senckenbergiana biologica different degree of exploration by researchers. 49: 259-264 • Isolation and resulting endemism seems to be the BRIGNOLI PM. (1971a): Su alcuni Leptyphantes di Creta driving force for the cave faunal patterns obser- (Araneae, Linyphiidae). - Fragmenta entomologica 7: ved. 231-241

• Most characteristic, in a faunal and zoogeographical BRIGNOLI PM. (1971b): Beitrag zur Kenntnis der mediter- ranen Pholcidae (Arachnida, Araneae). - Mitteilungen respect, is the presence of 58 endemic species in aus dem Museum für Naturkunde in Berlin 47: 255-267 Greek caves. -doi: 10. 1002/mmnz. 19710470203 • The high percentage of the endemics (53.2 %) BRIGNOLI P.M. (1972): Su alcuni ragni cavernicoli di Corfu suggests a local speciation process with consequent (Arachnida, Araneae). - Revue suisse de Zoologie 79: formation of neo-endemics. 861-869 • All troglobitic spiders are endemic, which leads BRIGNOLI PM. (1974a): Ragni di Grecia VI. Specie nuove to the conclusion that the regions where they are o interessanti delle isole Ionie e della Morea (Araneae). currently distributed were major centres of specia- - Revue suisse de Zoologie 81: 155-175 tion. BRIGNOLI P.M. (1974b): Ragni di Grecia VII. Raccolte in

grotte dell’Attica del Dr P. Strinati (Araneae). - Revue suisse de Zoologie 81: 493-499 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/;

Cove-dwelling spiders of Greece 31

BRIGNOLI P.M. (1974c): Araignees de Grece VIII. Quel- especes mediteraneennes occidentales exceptees. - Ti- ques Leptonetidae de la Laconie et de Me de Crete jdschrift voor Entomologie 131: 141-269 - (Arachnida, Araneae). Annales de Speleologie 29: DELTSHEV C. (1979): A contribution to the study of cave 63-70 spiders (Araneae) in Greece. Four new species (Araneae, BRIGNOLI P.M. (1976): Ragni di Grecia IX. Specie nuove Nesticidae, Linyphiidae) from the islands of Crete and o interessanti delle famiglie Leptonetidae, Dysderidae, Thera. - Acta zoologica bulgarica 13: 53-63 - Pholcidae ed Agelenidae (Araneae). Revue suisse de DELTSHEV C. (1985): New data concerning cave spiders Zoologie 83: 539-578 (Araneae) in Greece with description of a new Lepton- BRIGNOLI P.M. (1977): Ragni di Grecia X. Nuovi dati sulla etela (Araneae, Leptonetidae). - Acta zoologica bulgarica Grecia continentale ed insulare (Araneae). - Revue suisse 27: 41-45

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