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

2011·7.5 MB·English

by  Christo Deltshev

1
Arachnologische Mitteilungen40:23-32 Nuremberg,January201
The faunistic diversity of cave-dwelling spiders (Arachnida,Araneae) of Greece
Christo Deltshev
doi:10.5431/aramit4004
Abstract:Untiltoday,fromGreekcavesatotalof109speciesofspiders belongingto25familiesareknown.One
species,the linyphiid Porrhommaconvexum(Westring, 1861) was recorded hereforthefirsttime in Greece.The 109
speciesaredistributed in cavesofdifferentgeographicterritoriesasfollows:Thrace-8species,Macedonia- 18,
Epirus - 1,Thessaly- 6,Central Greece- 3,Attiki-Saronic Islands - 24,Peloponnese - 15,Evoia-Vories Sporades
-1,EasternAegeanIslands-5,Cyclades-3,Dodecanese-6,Ionian Islands-23,Crete-47.Thelargestfractionof
troglobitespecieswereencountered mainlyintheterritoriesofCrete- 15species(5ofwhichareanophthalmic),
theIonian Islands-4,Thrace-2(bothanophthalmic),theAttiki-Saronic Islands-2 (bothanophthalmic),the Pe-
loponnese-2(oneanophthalmic),andMacedonia,Thessaly,andtheCyclades-eachwith2species.Therichness
ofthetroglobiticspidersintheseregionsstrengthenstheassumptionthattheyweremajorcentresofspeciation
and evolution forthe species ofthis group.According totheircurrent distribution,the established 109 species
can be classified into 12 zoogeograpical categories, grouped into 4 complexes (widely distributed, European,
Mediterranean,endemics).The largest numberofspecies belong totheendemiccomplex (53.2%) and arealso
the mostcharacteristicand reflectthelocal characterofthecave-dwelling spiders.
Keywords:cave-spidersfauna,endemics,troglobites,zoogeography
TheearliestdataonGreekcave-dwellingspiderswere on the southern end ofthe Balkan Peninsula. The
presented by SIMON (1885), KULCZYNSKI (1903), country has borders with Albania, the Republic of
ROEWER (1928, 1959), DRENSKY (1936); Kra- MacedoniaandBulgariato the north,andTurkeyto
TOCHVIL (1937, 1938), HADJISSARANTOS (1940), the east. The Aegean Sea lies to the east and south
and FAGE (1945). More recent publications derive ofmainland Greece,while the Ionian Sealies to the
from the investigations ofBRIGNOLI (1968, 1971a, west. Bothparts ofthe EasternMediterraneanbasin
1971b,1972,1974a,1974b,1974c, 1976,1977, 1978, feature a vast number ofislands, islets and rock is-
1979, 1984), Deeleman-Reinhold (1971, 1977, lands (Fig.1).Two-thirdsoftheterritoryofGreeceis
1983, 1985, 1989, 1993), DEELEMAN-REINHOLD dominatedbylimestone,manyofwhicharekarstified
& DEELEMAN (1988), SENGLET (1971, 2001), (CLENDENON 2009).
Deltshev (1979, 1985, 1999,2000,2008), Beron The territory ofGreece can be divided into 13
& & &
(1985, 1986), Beron Stoev (2004),Thaler geographicalregions(BOSMANS CHATZAKI2005;
Knoflach (1995),Wunderlich (1995), Bosse- Fig. 1).Thereare7geographicalregionsonthemain-
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),CHATZAKIetal2002,BOSMANS Islands are situated on thewesternborderofGreece
dtCHATZAKI(2005),CHATZAKI&ARNEDO(2006), in the Ionian Sea.There are several island groups in
and PLATNICK (2009).The criticalincorporation of theAegean Sea on the eastern side ofGreece: Evoia
allavailableliteraturerecordsandtheaccumulationof and the Sporades,the SaronicIslands (groupedwith
newdataarenowsufficienttoallowacriticalanalysis Attica), the Cyclades, the Eastern Aegean Islands,
ofthe distributionofspiders establishedinthe caves the Dodecanese and Crete (Fig. 1).
ofGreece.
Results
Studyareaandmaterial Speciescomposition
Greeceisacountryinsouth-easternEurope,situated ThespidersestablishedinthecavesofGreece(Main-
landandInsularpart) arerepresentedby109 species,
ChristoDELTSHEV,InstituteofBiodiversityandEcosystem included in 52 genera and 25 families: Ctenizidae
Research,1TsarOsvoboditelBlvd;1000Sofia,Bulgaria - 1, Filistatidae - 1, Sicariidae — 1, Scytodidae - 1,
E-Mail:[email protected] Leptonetidae - 9, Pholcidae - 10, Segestriidae - 3,
submitted:30.12.2010,accepted:9.4.2010;online:10.1.2011 Dysderidae-12,Oonopidae-1,Mimetidae-1,Ere-
24 C.Deltshev
(6), Dysderidae (3), Nesticidae (2), and Linyphiidae
(1). The genera with the largest number ofspecies
are Tegenaria (8), Lepthyphantes (6), Harpactea (5),
andHistopona (5).
The species are distributedin caves belongingto
thegeographicterritoriesofGreeceasfollows: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
degreeofexplorationintheseterritories is notequal:
the territories ofEvoia-Sporades, the Cyclades, the
Dodecanese and Central Greece are lessexplored.
Cave-dwelling spiders can be categorised into four
ecologicalgroups (SKET2008):
• troglobites: species limited to a life cycle in caves.
Often they show a suite ofcharacters, associated
with their adaptation to subterranean life: loss of
LIV1KÖ8f>ELAQOS pigment, loss ofeyes and elongation ofappenda-
ges.
Figure1:Mapofdifferentgeographical regionsinGreece. • eutroglophiles: species which can live their entire
life in caves,but also occurin other environments.
sidae-1,Uloboridae-1,Nesticidae-4,Theridiidae • subtroglophiles: species which utilise caves, but
-4,Anapidae- 1, Linyphiidae- 16,Tetragnathidae mustleave the caves to complete theirlife 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 Thelargestfractionoftroglobitespecieswasencoun-
speciesisnewfortheGreekspiderfauna:Porrhomma tered mainly in the caves ofthe territories ofCrete
convexum(Westring,1861) (markedinthelistwith*), -15 (5 anophthalmic),theIonianIslands-4,Thrace
aspiderwithaHolarcticdistributionandwidespread -2 (2anophthalmic),theAttiki-SaronicIslands2 (2
in European caves. It is also well represented in the anophthalmic),thePeloponnese-2(1 anophthalmic),
caves ofthe Balkan Penisula - and not only estab- andMacedonia,ThessalyandtheCycladeseachby2
lished in the caves ofCroatia, Romania andTurkey species (Table 1).Alltroglobitesareendemicsforthe
(DELTSHEV2008).Thenumberofspeciesishighand territoryofGreece or the Balkan Peninsula.
%
represents about 13 ofthe Greek spiders. This is Very important is the presence ofeutroglophiles
also evident from a comparison with the number of (35 species), because together with troglobites (29
cave-dwellingspidersrecordedfromthe othercoun- species),theycanbeconsideredasdependentfaunistic
tries ofthe Balkan Peninsula: Bulgaria- 99, Croatia elements ofcaves. The largest number ofspecies is
- 63, Serbia - 59, Bosnia and Herzegovina - 52, establishedinthe caves ofCrete (14 species),the Io-
Macedonia-44,Montenegro-44,Slovenia-43,Al- nianIslands(11 species),theAttiki-SaronicIslands(9
bania-10,Turkey-8,andRomania-4(DELTSHEV species),Macedonia(8species),andthePeloponnese
2008). The established number ofspecies, however, (8 species). Here, the endemics are represented by
A
depends not onlyon the size ofthe regions,but also 14 species (35 %). present day example ofactive
onthe degreeofexploration.The mostcharacteristic subterranean colonisation and cave penetration are
are the families: Leptonetidae (8.2 %), Pholcidae thespeciesLepthyphantescentromeroidesandPallidup-
(9.2 %), Dysderidae (11 %), Linyphiidae (14.6 %), hantesspelaeorum,widespreadintheBalkanPeninsula
and Agelenidae (19.2 %). The families with largest (DEELEMAN-REINHOLD 1978). Here, the species
number of anophthalmic species are Leptonetidae Palliduphantesistrianusshould also be included.
1 ;
Cove-dwellingspidersofGreece 25
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Cave-dwellingspidersofGreece 27
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28 C.Deltshev
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(KveCost.ciLhta,. (dKaullmcaztyihcsaki, (fPearnrzugeirn,ea (KCopa.cghLa,.na (KsilovLecstrh.is, B1arc9ihg7ane7oali, Barriiagdnnoaeli, (dComleestriccak, B1hra9iugs7nero9ili, Blraibygrnionltih,i Dpealrtasgahmeiva,ni (pFaoruirectroiyn,a B1pri9ieg7npo9elrii, Bsrchimganlofulsis,i Wcurnedtaeernlsiicsh, "H"<RRR1cLs> HX<>1U<Ib8l> Cshtarriatnodniov, (lWaalpcikednoaesru,s (rTihpoarrieelnls,is (leCnt.igLin.osa (1K8oL7c0h.), (1K8oL6c6h.), (KCo.cLh., (wAalucdkeonaueirin, CKco.ollLcinh.us, (Fabosriucimus, H4o<iO=o-Di (1C7le5r7ck), (caNneistnrinniii,
.s clivicola femellus oblongus glob o falcata
TAXA Maimuna Malthonica Malthonica Malthonica Tegenaria Tegenaria Tegenaria Tegenaria Tegenaria Tegenaria Tegenaria Tegenaria Tegenaria Tegenaria Amaurobiidae Amaurobius gRRs ’’ORsR'R Amaurobius Gnaphosidae Drassodes Nomisia Pterotricha Zelotes Zelotes Zelotes Sparassidae Eusparassus Philodromidae Philodromus Thomisidae Synema IR'Salticidae Evarcha Mendoza
Cave-dwellingspidersofGreece 29
Table2:ZoogeographicalcompositionofcavedwellingspidersinGreece
Complexes Chorotypes Species
Classification Code Number %
Cosmopolitian COS 7 6.4
HOL
Holarctic 5 4.6
Widelydistributed Palearctic PAL 9 8.2
WP
WestPalearctic 5 4.6
European-CentralAsiatic ECA 1 0.9
Total 27 24.7
European E 8 7.3
Middle-SoutheastEuropean MSEE 1 0.9
European South EastEuropean SEE 1 0.9
Total 10 9.2
MCA
Mediterranean-CentralAsiatic M 1 0.9
Mediterranean 5 4.62
Mediterranean NEM
North EastMediteranean 8 7.3
Total 14 12.8
EndemicforcontinentalandislandGreece EGR 55 50.4
EndemicforBalkanPenisnula EBP 3 2.7
Endemics
58 53.2
Total
Thegroupofsubtroglophilescomprises11 species humid Tertiary forests (DEELEMAN-REINHOLD
occurringindarkplacessuchasbuildings,screes,rock 1977).However,duetoalackofdata,itisdifficultto
crevices andcaves.Theycanbeconsideredasregular determinewithcertaintywhichcavespiderendemics
inhabitants ofthe caves.Three species are endemics oftheBalkansareTertiary,andwhichareQuaternary,
(Maimunacretica,Hoplppholcusminous,and Tegenaria elements.
labyrinthi). The complex ofwidely 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 %),followedbyCosmopolitan (19.4
but their presence in caves should not be ignored. %) Holarctic (13.8 %),WestPalearctic (13.8 %), and
On the other hand some ofthem 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 ecologyis gathered. lowlands, buildings, caves, woodlands and high al-
Here, the endemics are representedby 11 species. titude zones of mountains. Here characteristic for
caves are Nesticus cellulanus Porrhomma convexum
,
Zoogeographicalanalysis andTegenariadomestica.Thesespeciesarealsolargely
Accordingtotheircurrentdistribution,theestablished
109 species can be classified into 12 zoogeograpical
categories,grouped into fourcomplexes (widelydis-
tributed, European,Mediterranean,endemics) (Tab.
1&2, Fig. 2).
Bestrepresentedisthecomplexofendemicswith
58 species (53.2 %), which include 29 troglobites,
16 troglophiles, and 12 trogloxenes.The established
numberis highandreflects thelocalcharacterofthe
cavefauna.TheendemicsarebestrepresentedinCrete
-26 species and2 genera (.Minotauria, Rhodera),the
IonianIslands-9species,thePeloponnese-8species,
Macedonia and the Eastern Aegean Islands - each Figure2:Relativerepresentationofthezoogeographical
with4species.Therecentcavespiderfaunaisformed categoriesofcavedwellingspidersinGreece(forabbre-
viationsseeTable 1).
after gradual changes in the fauna of the ancient
30 CDeltshev
widespreadtaxainEuropeancaveswherePorrhomma 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.
%). European species are dominant (8 %) among Pertrov,andP.Stoevforthecollectedmaterialandvaluable
this complex. The Middle Southeast European and discussions.TheresearchwasalsosupportedbytheProject
-Nos.B-1523/05 andBG051P0001-3.3.04/41.
SoutheastEuropeanspeciesarerepresentedbysingle
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861-869
• All troglobitic spiders are endemic, which leads
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