Assemblages of herb-dwelling spiders (Araneae) of various steppe types in Ukraine and Central Chernozem of Russia

2012·9.6 MB·English

by  Nina Y Polchaninova

Arachnologische Mitteilungen43:66-78 Nuremberg,July2012
Assemblages of herb-dwelling spiders (Araneae) ofvarious steppe types in
Ukraine and the Central Chernozem region of Russia
NinaY.Polchaninova
doi;10.5431/aramit4312
Abstract; A total of 107 spider species from 15 families were recorded in the herbaceous vegetation ofthe
steppeecosystemsofUkraineandtheCentralChernozemregionofRussia.Araneidae,Thomisidae,Salticidaeand
Theridiidaewerethe mostspecies-rich.The speciescomposition depended on the steppetype;adjacentforest
habitatsinfluencedsteppefaunaintheforest-steppeandnorthernpartofthesteppenaturalzone.Thenumberof
generalist,forestandwetlanddwellingspeciesinthesteppevegetationshowedatendencytodecreasetowards
thesouth.Dominanceofherb-dwelling spiderswasspecifictoeach steppetype;nosinglespecieswasfoundto
predominateinallthesteppe habitats.
Keywords:dominancestructure,speciesdistribution,spidercommunities,steppeecosystems
Steppes are the most transformed ecosystem in assemblages(CHERNOV1975).Habitatpreferenceof
Ukraine.The steppe natural zone comprises 40% of speciesdependsonthenaturalzone(KÜHNELT1943,
the countryandabout 80% ofthis territorywas once Walteri960,Bei-BienkO 1966). Accordingtothe
covered with steppe vegetation. Presently, only 3% so-called principleofzonalchangeofhabitats’(BEI-
ofrelativelyundisturbedvirginsteppeshavesurvived BIENKO 1966),ortheprincipleof‘relativestenotopy’
intact.They are preserved mainly in nature reserves (Schaefer1992),widespreadspeciesmovingnorth-
or on gully slopes and saline lands not suitable for wardscanchangetheirhabitatstodryerwarmeropen
agriculture (KOTENKO 1996). siteswithsparsevegetation,whilegoingsouthwards,
Allsteppereservesareisolatedtovariousdegrees, thesamespeciesinhabitmoisterandshadierhabitats
suchthattheycanbeconsideredasecologicalislands, with dense vegetation cover.These two phenomena
surrounded by agricultural landscapes (MALYSHEV wereillustratedbytheexampleofspidercommunities
1980). Different components of the steppe biota ofthe Urals transect (ESYUNIN 2009), oakforests of
respond to isolation in different ways (KOTENKO the East European Plain (ESYUNIN et al. 1994), and
1996,Hanser&cHuntly2006,Laiolo&Tella Ukrainian steppes (POLCHANINOVA 1990a, 1996).
2006).Spidersmovequiteeasilyfromnaturalhabitats Investigation ofthe spider fauna and ecology in the
to agricultural fields, however many ofthem do not area in question began in the 1980s in the Central
penetratebeyond the field margins. Crops are popu- ChernozemReserveofRussia(PiCHKA1984 ,1984b)
latedmainlybyeurytopicspeciestypicalofmeadows and in the steppe reserves ofUkraine (POLCHANI-
ordisturbedhabitats(SEYFULINA&TCHERNYSHEV NOVA 1988, 1990b). Based on the data obtained, a
2001, SEYFULINA2010). firstattemptwasmadetoanalyzethefaunaandspider
Adjacent habitats have an undeniable impact on communities of the steppe ecosystems of Ukraine
steppe communities (CHERNOV &c PENEV 1993), (Polchaninova 1990b). Further research focused
especially in the forest-steppe zone where forest on all steppe reserves in Ukraine (GURIANOVA &c
and steppe are both zonal plant formations, and Khomenko 1991, Polchaninova 1998, Pol-
an active species interchange can be supposed to chaninova &C PROKOPENKPO 2007a, PROKO-
take place. To the north and to the south the dif- PENKO 2001, Prokopenko et al. 2008) and the
ference in microclimatic conditions between zonal adjacent territoryofRussia (POLCHANINOVA2003,
and intrazonal habitats increases, which results in 2009, Ponomarev 2005, 2010). Thus, it became
higher species specificity oftheir animal and plant possibletoanalysethespeciescompositionanddomi-
nance structure ofspiderassemblages to seewhether
NinaY.POLCHANINOVA,DepartnnentofZoologyandAnimalEcology, theydependonthe steppe type.Thepresentpaperis
KharkivNationalUniversity,4,SvobodySq.61022Kharkiv,Ukraine. partofacomprehensivestudyofspidercommunities
E-Mail:[email protected]
ofsteppeecosystemsofUkraineandEuropeanRussia
submitted:/.12.2011,accepted:4.6.2012;onlineearly:31.7.2012 and concerns herb-dwelling spiders only.
Spidersinsteppein UkraineandCentralChernozemRussia 67
Materialsandmethods
The material was collected from three
localities in the Central Chernozem
region ofRussia, and eight localities in
the Left-bankUkraine (Fig. 1). Central
ChernozemisasegmentoftheEurasian
chernozem (blacksoilcontainingahigh
percentageofhumus)beltthatlieswithin
the EastEuropean Plain (also known as
the RussianPlain).Itisawell-delimited
geographical and administrative region.
The Left-bank Ukraine in the present
contextis a territorythat stretches from
the leftbankofthe Dnieper River east-
ward to the border ofthe state.
We
investigated five types of frag-
ments of virgin steppes in the East
European Plain (classification accord-
ing to Gribova et al. 1980). Meadow,
forb-bunchgrassandbunchgrasssteppes
belong to the categoryofzonal steppes,
whilesandyandchalkysteppesbelongto
theazonalsteppes.InRussiangeobotany,
meadow steppes situated in the forest
steppe natural zone are called ‘northern Fig.1:Mapoflocalities(I-forest-steppezone,II-steppezone:lla-subzone
steppes’ in contrast to genuine steppes, offorb-bunchgrasssteppes,Mb-subzoneofbunchgrasssteppes.For
which form two subzones ofthe steppe numberoflocality,seeTab.1).
natural zone (Fig. 1,Tab. 1).We regard
sandy steppes situated in different steppe subzones Tab. 1.Hereandadditionallyinthegraphsandtables,
separately as northern and southern variants. The thelocalitiesarearrangedfromnorthtosouth.Names
studied localities, investigation period and abbrevia- ofsettlementsandnaturereservesaretranslatedfrom
tionsusedfurtherinthetablesandgraphsaregivenin Russian and Ukrainian respectively.
Tab.1:Listandcharacteristicsofstudysitesincludingtypeofsteppehabitat.
Naturezone/ Nameoflocality Steppetype Yearsof sitearea abbrev. Region
subzone study (ha)
1. Streltsovskaya meadowsteppe 1998 730 meadl KurskArea,Central
steppe 1999 Chernozem Res.
2007 5F36'N 36U2T
m
248 a.s.l.
2. Kazatskaya meadowsteppe 1999 1010 mead2 KurskArea,Central
steppe 2000 Chernozem Res.
2007 5T32'N 36°20T
m
Forest-steppe 230 a.s.l.
zone 3.Yamskaya meadowsteppe 2001 500 mead3 BelgorodArea,Belogorye
steppe 2002 Res.
51TLN3r45T
m
193 a.s.l.
4.Mikhailivska meadowsteppe 1985 160 mead4 SymskaAreaUkrainian
Tsilyna 1986 steppe Res.
1988 50°34'N 44°12T
m
172 a.s.l.
68 N.Polchaninova
Naturezone/ Nameoflocality Steppetype Yearsof sitearea abbrev. Region
subzone study (ha)
5. Striltsivskyi forb-bunchgrass 1986 1000 forbl LuhanskArea,Luhanskyi
step steppe 1988 NaturalRes.
2009 49°17'N 40°00T
m
147 a.s.l.
Steppezone: 6. Khomutivskyi forb-bunchgrass 1982 1000 forb2 DonetskArea,Ukrainian
Northern step steppe 1983 steppe Res.
subzone 2004 4ri7'N38°10'E
m
offorb- 57 a.s.l.
bunchgrass 7. Striltsivskyi chalkysteppe 2009 80 chalk LuhanskArea
steppe zakaznyk 2011 49°18'N 39°50T
m
100 a.s.l.
8.Dnieprovsko- Northernsandy 1999 300 sandN DnipropetrovskArea
Orilskyi Res. steppe 2000 48°30'N 3T45T
m
2003 53 a.s.l.
9.Askania-Nova bunchgrass 1984 11000 bunch KhersonArea,
Res. steppe 1985 46°28^N 33°58T
m
1988 29 a.s.l.
Steppezone: 10.Ivano- Southern sandy 1989 2074 sandSl KhersonArea,
Southern Rybalchanskyi steppe 1990 Chernomorskyi Res.
subzoneof 1991 46°27'N 32°07T
m
bunchgrass 6 a.s.l.
steppe 11. Solenoozernyi Southernsandy 1995 1325 sandS2 KhersonArea,
steppe 1996 Chernomorskyi Res.
1998 46°27'N3F58T
m
1 a.s.l.
Ineachlocality,investigationswerecarriedoutfor immature specimens could not be distinguished, we
two consecutiveyears.The materialwas collectedby considered thegenus as awhole ingeneralquantita-
sweeping with an entomological net (d=30 cm), as tive analysis, and then species relationships within
wellasbyhandcollecting.Quantitativesampleswere thegenuswere estimatedseparatelybasedonmature
takeneverymonthfromMaytoSeptember,fivesam- individuals.
ples of50 sweeps each.The material obtained from Inordertodeterminerelativeabundance,weused
two vegetation seasons was considered as a general theTischlerrating scale,where eudominant n>10%,
sample. For comparative analysis, we chose plots in dominant 5<n<10%, subdominant 2.5<n<5%, rece-
naturereserveswithstrictlyprotectedsteppe,because dent l<n<2.5%, and subrecedent n<l% (TISCHLER
traditionalregime management such as haymowing 1949). We considered eudominants and dominants
dramatically impacts upon species composition and together as a dominant complex. The ecological
structure of spider communities (POLCHANINOVA groups ofspecieswere determinedbased on the data
2004). In order to obtain more information on the on their habitat preference within the forest-steppe
spider species composition oflocal faunas and vari- andsteppezonesoftheEastEuropeanPlain(Tab.2).
oussteppetypes,weconductedadditionalresearchin Wedefinegrasslandspeciesasspeciesoccurringinall
differentyears, and also used the data from available habitatswithpredominanceofherbaceousvegetation,
publications (PiCHKA 1984a, 1984b, GURYANOVA in our case meadows, steppes, and open slopes of
&KHOMENKO 1991,Prokopenko2001).Intotal, gullies. We consider permanent residents ofone or
morethan 12,000spiderspecimenswerecollected.A more steppe types to be typical species. Sometimes
list ofspecies is given in the Appendix. Species are theymayoccurin other habitats as rare finds.
arrangedbyfamilies accordingto PLATNICK (2012). Similarity of spider assemblages of the studied
Withthemethodofnetsweeping,alargenumber habitatswasdeterminedbyclusteranalysisperformed
ofjuvenilespiderswerecollected.Inmanycases,they inStatistica7(STATSOFTINC.2004).Twoyearquan-
could be identified to species level as theybelonged titativesamples (seeabove)werepooledtogetherand
to a single species ofthe genus in agiven locality, or apercentageofeachspeciesofthetotalsampleateach
differed in period ofmaturity. For the generawhose sitewascalculated.WeusedtheWardsalgorithmasa
Spidersinsteppein UkraineandCentralChernozemRussia 69
dustermethodandEuclidiandistanceas asimilarity steppe.The spider fauna ofmeadow steppe situated
measure. in forest-steppe zone differs significantly from that
ofsteppe habitats in steppe zone.Within the steppe
Results zone,spidersaremoreevenlyspread,andthefaunaof
Speciescompositionandspeciesrichness eachsteppetypehasonly7-12%ofspeciesnotfound
A total of107 spider species from 15 families were in other steppe types.
recorded in herbaceous vegetation at 11 study sites. Species richness of herb-dwelling spiders was
Four families were the most species-rich: Araneidae maximal in both fragments of the forb steppe and
(20species, 18.7%oftotalspeciesinstudysites),Sal- minimal in both fragments of the southern sandy
ticidae (19 species, 17.8%),Thomisidae (18 species, steppe (Appendix). In the largest steppe fragment
16.8%), and Theridiidae (13 species, 12.1%). Only (bunchgrass steppe in Askania-Nova) the species
Araneidaeweredistributedquiteevenly,rangingfrom richness was slightlylower than that in the smallest
22%ofthespeciesinforbsteppesto27%inthesandy one (chalkysteppe inMilove District).
ones. Linyphiidae and Clubionidae occurred mainly
inmeadowsteppes (Appendix).Thomisidae reached Ecologicalgroupsofspecies
their maximum of species richness in bunchgrass Meadowsteppes arecharacterizedbyalarge number
steppes (23%) and fell to a minimum in meadow offorestandwetland species (Tab.2). Some ofthem
steppes (13%). Salticidae are known to increase in appear in the steppe in wet years, and nine species
terms of the number of species towards the south are permanent residents. The number ofwetland
(Nenilin 1984, Mikhailov 1997). In our collec- and forest species gradually decrease towards the
tion,wefoundunexpectedlyfewsalticidspeciesinthe south. In different steppe types, generalist species
bunchgrasssteppe(4species,10%);intheothersteppe comprised 15-20%.The numberofsteppe species at
types this increase could be seen not in absolute but all sites was significantly lower than that ofspecies
onlyin relative numbers because ofimpoverishment widelydistributedingrasslands. In meadowsteppes,
ofthe speciescompositioninthe southernsteppes (9 we found no steppe species.
species, 15% in meadow steppes, 11 species, 17% in Therewere no specialist species inthe herb layer
forbsteppes,9species,20%innorthensandysteppes, ofsteppe vegetation. Eight species whichwere con-
7 species,21% in the southern ones). sideredtobetypicalwerealsofoundinotherhabitats;
A third ofthe species found in meadow steppe however, they reached their maximal abundance
have not been recorded southwards in genuine onlyin a certain steppe type.These species are Run-
Tab.2:Ecologicalgroupsofspidersofvarioussteppetypes.
Numberofspeciesinsteppetype
Ecologicalgroups meadow forb-bunch- chalky sandyN bunch-grass sandyS
steppe grass
generalist 10 11 9 8 6 5
wetland + forest 17 7 2 4 3 1
forest 7 3 1
wetland 7 2 2
grassland + forest 5 10 9 9 5 6
edges
grassland 9 12 10 11 9 8
steppe 8 6 2 6 2
unspecified 6 9 2 4 7 5
1
1
typical 2 1
1 1
3
70 N.Polchaninova
100%
Others
^ 75% 03 Miturgidae
q; Oxyopidae
1 0
Salticidae
Theridiidae
c 50% 1^ Philodromidae
_o H Thomisidae
S
o Dictynidae
QO. ^ Araneidae
Q. 25%
0%
sites
Fig.2:Familyabundance(proportionoftotal numbersoffamiliesateachsite) inspiderassemblagesofherba-
ceousvegetationofsteppehabitats.Sitesarearrangedfrom northtosouth.
Among five steppe species of the southern
steppesubzone,threespeciesoccurredinsandy
and chalky steppes of the northern subzone
whiletwo species did notextendbeyondzonal
We
boundaries. did notfindtendencies in dis-
tribution ofthe other species; 22 species were
found as singletons,thereforewe cannotjudge
their habitat preference.
Familyabundance
The herbaceous vegetation ofsteppe habitats
was dominated by Araneidae at all sites (Fig.
2). The second largest family Thomisidae
preferredforb and chalkysteppes.The relative
LinkageDistance numberofDictynidaewentdownfrommeadow
Fig.3:Dendrogramofbiocoenoticsimilarityofherbdwellingspider steppestobunchgrassandsouthernsandyones.
assemblagesofvarioussitesbasedonthe%ofeachspeciesin Philodromidae, likewise, were least abundant
thetwoyearsamplesineachsiteusingWard'smethodascluster
algorithmand Euclidiandistanceassimilaritymeasure. in the three southern sites while they reached
a maximum in forb steppes and in the‘mead4’
ciniagrammica, Pellenus seriatus and Yllenus vittatus site.ThedistributionofSalticidaedidnotshowalati-
in southern sandy steppe, Heliophanuslineiventris in tudinal trend.An unusuallylarge individual number
southern sandyand bunchgrass steppe, and Uloborus ofthis familywas found in one ofthe sandy steppe
A
walckenariusin sandyand chalky steppes. sites. characteristicfeatureofthespiderassemblage
Thirteengeneralistandgrasslanddwellingspecies ofthe bunchgrass steppewas the high abundance of
occurred in almost all investigated steppe fragments Oxyopidae.
(Appendix). Four species found in steppes of the
forest-steppe and northern steppe subzone changed Biocenoticsimilarities
A
their habitat preference to forests orwetlands in the dendrogram ofbiocenotic similarity ofspider as-
southern subzone.Tenwetland species spread to the semblagesofvarioussitesbringsthemtogetherbased
souththroughsteppefragments nofurtherthan sites on a zonal-subzonal principle (Fig. 3). First, two
‘forbl’and‘sandyN’;southwardstheyalsomovedinto clusters ofthe sites ofthe forest-steppe and steppe
intrazonal habitats. Five steppe species were widely zone are separated, and then the sites of northern
spread in the steppe zone, but absent in meadow and southern subzones are divided into two groups.
steppes; three species occurred in meadow steppes As expected, assemblages oftwo neighbouring frag-
onlyinthesite‘mead3’.Itisthedriesteasternsitewith ments of southern steppes were similar, however,
a character ofvegetation closer to genuine steppes. theirsimilaritywaslessmanifestthanthatofthetwo
Spidersinsteppein UkraineandCentralChernozemRussia 71
adjacent meadow steppe plots and even ofthe forb grasslands,butinthesouthitisparticularlyabundant.
and chalkyones. Phylloneta impressa, another dry grassland species,
preferred sandy and chalky steppes. Dictyna arundi-
Dominancestructure nacea,bycontrast,waseudominantinmeadowsteppes
In the study areas, 13 species occurred in all steppe and occurred in high numbers in the forb ones.This
habitats.Inaddition,twospecieswerefoundsporadi- speciesischaracteristicofgrasslandsandforestedges
callyinbothnorthernandgenuinesteppes.However in the forest-steppe and northern part ofthe steppe
none ofthem dominated in all steppe types. zone.Thegroupofspecieswithanarrowerdominance
Neosconaadiantawas distinguishedbythewidest spectrumwasrepresentedhyNeottiurabimaculataand
habitat spectrum (Tab. 3). It is common in various Araneusquadratusinmeadowsteppes,andbyThomisus
Tab.3:Dominantspiderspeciesindifferentsteppehabitats.
eudominantn>10%,^^-dominant,5<n<10%,W-subdominant2,5<n<5%,
recedent<n<2,5%,#-subrecedentn<1%ofspiderscollected ineach locality
Localities
.apecies
1 2 3 4 5 6 7 8 9 10 11
•
•
Neottiurabimaculata • a
• • •
Araneusquadratus • •
•
Cheiracanthiumpunctorium a a • •
• • • • •
• •
Evarchaspp.
• • • •
• •
• •
Dictynaarundinacea a a
• • • •
• • • •
Tibellusspp. • •
•
Xysticusspp.
•
• • • •
Agalenatearedi • • • • •
• • •
• •
Mangoraacalypha • • • • • •
• • • • •
• • • •
Neosconaadianta • •
• • • •
• • •
• • • •
Heliophanusspp.
• •
• •
Phyllonetaimpressa • • • • • • •
•
•
Cheiracanthiumpennyi • • • • • •
• • • •
Philaeuschrysops •
•
• • •
•
Thomisusonustus • •
•
Uloboruswalckenaerius • a a
•
•
Oxyopesheterophthalmus • • •
72 N.Polchaninova
Despite the local differences in species
compositionanddominancestructureofspider
assemblages,anumberofcharacteristicfeatures
canbe distinguished foreverysteppe type.All
the spider complexes ofsteppe herbage had a
polydominant structure.
In the four sites of meadow steppes, the
group of dominants consisted of Dictyna
arundinacea (16.0-24.3%), Tibellus oblongus
(8.7-14.2%) and Xysticus cristatus (6.0-9.3%).
Araneusquadratuswdcs,amemberofthiscomplex
in three sites,NeottiurabimaculatUyLarinioides
patagiatuSy Cheiracanthiumpunctorium,Evarcha
arcuatay andNeosconaadiantawerelocaldomi-
nants in notmore thanone site each.Marpissa
pomatiay Hypsosinga sanguinea, and Singa
hamata were subdominants in the meadow
steppes. Southwards in genuine steppes they
did not occur, or were found only as a few
specimens.Howeverwecannotconsiderthem
Fig.4:Relativeabundance(%ofindividualsoutofthetotal number tobetypicalspeciesofmeadowsteppesbecause
ofspiderscollectedateachsite)of(A) Tibellusoblongusand T. theywere also numerous inwetlands (personal
macellusand (B)XysticuscristatusandX.striatipesin herbaceous
vegetationofdifferentsteppehabitats.Sitesarearrangedfrom observation). The specificity ofthe spider as-
northtosouth. semblage ofthe driest eastern site ‘mead4’ is
worthmentioning.TheabundanceofNeoscona
onustus and Philaeus chrysops in genuine steppes. Six adiantaandtheappearanceofEvarchamichailoviand
species were dominants at only one site {Uloborus Xysticusstriatipes make it similar to the assemblages
walckenaerius, Cheiracanthiumpennyi^ C.punctorium, offorb steppes.
Philodromus histrio^ Pellenesseriatus, and Yllenus vit- Xysticus striatipes (9.3-15.7%), Tibellus spp.
tatus). (10.7-12.6%), andNeosconaadianta (8.0-9.7%) were
Offurther interest are the latitudinal changes of common dominants offorb steppes,withAgalenatea
abundance of common species of the same genus. redii and Mangora acalypha co-occurringwith them
In the pairs Tibellus oblongus-macellus and Xysticus in the northern site, and Philaeus chrysopSy Evarcha
cristatus-striatipesy the first species ofeach pair was michailoviyHeliophanusflavipesand Thomisusonustus
A
abundantintheforest-steppeandinthenorthofthe in the southernone. range ofspecies typicalofdry
steppe zone (Fig. 3).The second species ofeachpair grasslands appear in these steppes {Heterotheridion
appeared in genuine steppes, and in case ofXysticuSy nigrovariegatumy Theridion innoccuumy Heriaeus ob-
replaced the first one. Further to the south they all longuSyTibellusmacelluSyXysticusmarmoratus,Synageles
decreased in number or disappeared in steppe plots, hilarulusy and Philaeus chrysops). However, we found
although they remained quite common in adjacent no species specific for the forb steppes only.
forest stands (POLCHANINOVA 1990a). Heliophanus Thedominancecomplexofchalkysteppeincluded
flavipesoccurredinaUsteppetypes.Inthethreesouth- Xysticus striatipes {2S.S%)yAgalenatea redii (11.6%),
ern sites, H. lineiventrisco-occurredwith H.flavipes Neoscona adianta (10.0%), Tibellus spp. (6.4%), PhT
andtogethertheymadeupasignificantproportionof laeuschrysops(6.7%),andDictynaarundinacea(5.8%).
the individuals found in these sites (Tab. 3).Evarcha Inthe northern sandysteppeitconsistedofNeoscona
arcuatawas common in meadow steppes and in the adianta(14.9%),Xysticusstriatipes(11.0%),Mangora
northern sandy steppe. E. michailovi occurred in acalypha (8.9%), Dictyna arundinacea (5.8%), Phi-
forb steppes andpenetratedinto one site ofmeadow lodromus histrio (7.7%), and Uloborus walckenaerius
steppes. Southward in the subzone of bunchgrass (5.3%).Intheareainvestigated,thelattertwospecies
steppes, neither ofthese specieswas found. areassociatedwithvegetationofsandysoil.Theywere
also common in calcareous lands (POLCHANINOVA
SpidersinsteppeinUkraineandCentralChernozemRussia 73
2010). Simitidionsimileand Cyclosaoculatapreferred bothhabitatandvegetationlayermovingfromherbs
the same habitats but they had larger ecological to the ground (POLCHANINOVA 1990a).Agroup of
flexibility, inhabiting meadows, shrubbysteppes and southern species which spread to the north through
&
even open deciduous forests (POLCHANINOVA the dry open sites made up only 4% ofthe species
Prokopenko 2007b). Gibbaranea bituberculata and found in the steppe habitats. Our data confirm a
G. ullrichicametothesteppeplotsfromneighbouring decrease ofgeneralist, forest and wetland dwelling
pine forests. species to the south and predominance ofgrassland
The spider assemblage ofthe bunchgrass steppe species in genuine steppes. The same tendencies
wascharacterizedbylargenumbersofthreeeudomi- in distribution ofecological groups ofspiders were
nants Neoscona adianta (29.8%), Oxyopes heteroph- shownbyESYUNIN (2009) in a transectthrough the
thalmus (26.2%), and Thomisus onustus (14.0%), the Urals.
occurrenceofonesubdominantXysticusstriatipes,and The relative abundance of spider families in
single finds ofotherspecies. Southern sandysteppes steppe vegetation changes depending on the steppe
were dominated atboth sitesbyHeliophanusflavipes^ type.Thisisconfirmedbydataonsteppelocalitiesin
H. lineiventris(15.6-20.0%),Neosconaadianta(11.0- other regions.Thus, in mountain steppes ofBashki-
12.2%),andPhyllonetaimpressa(5.6-6.3%).Inoneof ria (Southern Urals), Tetragnathidae, Thomisidae,
them(Ivano-Rybalchanskysite)thedominancecom- and Araneidae were dominants in numbers, while
plexwassupplementedwith Oxyopesheterophthalmus^ Thomisidae comprised the main partofthe biomass
Cheiracanthiumpennyi^Thomisusonustus^andRuncinia (Efimik 1989). According to our data, Araneidae
grammica, and in the otherone (Solenoozerny) with was the most abundant familywhileTetragnathidae
Pellenes seriatus and Yllenus vittatus. The presence occasionally occurred in steppe as rare finds. In the
ofthe latter three species and the abundance ofH. east ofthe Russian Plain, in stony and forb steppes
lineiventris is characteristic of the southern sandy ofSamarskaya Luka,Thomisidae reached the high-
steppes. est abundance followed byAraneidae and Salticidae
(KRASNOBAEV 2003). In eastern Hungary (Great
Discussion Hungarian Plain), drysandygrassland communities
Spiderspeciesrichnessoftheinvestigatedsteppefrag- that are close to sandy steppes were also dominated
mentsdependedonthesteppetype.Thelownumber by Araneidae and Thomisidae while Salticidae and
ofspecies on the largest site ofbunchgrass steppes Dictynidaecomprisedasecond-rankedgroup(HOR-
canbe explainedbythe monotonous plainreliefand VATH et al. 2009).
absenceofneighbouringintrazonalhabitats.Presum- NosinglespeciesdominatedinaUthesteppetypes
ably, invertebrate communities within small sites and evenwithin one type,a dominance complexwas
increase in species richness due to species exchange composedofdifferentspecies.Meadowsteppeswere
with adjacent habitats (MAGURA & KODOBOCZ dominatedbywetlandspeciesandgenuinesteppesby
2007). This statement is true for the forest-steppe wide-spreadgrassland species.In azonalsteppes,the
and northernpartofthe steppezonewhere an active complexincludedspeciesinhabitingsparsevegetation
interchange takes place (CHERNOV 1975). In our ofsandyandchalkysoils.Dominantspiderspeciesof
investigations,werecordedthepresenceofforestand steppe and steppe-like habitats oftheUrals alsovary
wetland species in meadow steppes. However in the depending on zonal and local conditions (ESYUNIN
south ofthe steppe zone, despite a rich local fauna 2009).
in general and the existence ofneighbouring forest Inconclusionthespiderassemblagesofthenorth-
stands and lakes, the species composition ofspiders ernmostmeadowsteppesandthesouthernmostsandy
ofsteppe fragments remained the poorest. Onlyone steppes differeddramatically, as expected,because of
wetland specieswas found there. zonal-climatic differences. From a zonal standpoint,
About 12%ofthe species oftheinvestigatedarea thefirsthavemuchincommonwithmeadows(PON-
&
occurred in all steppe types. Another 12% changed OMAREV POLCHANINOVA 2006), and the latter
their habitat preference spreading from north to withbrackish meadows and salt-marsheswith dense
south and moved from steppe sites to shaded and vegetation(POLCHANINOVA1998).Inourstudy,the
moist ones. These data illustrate well the ‘principle most specificwere herb dwelling spider assemblages
ofzonal change ofhabitats’by BEI-BIENKO (1966). ofthebunchgrass stepperepresentedinUkrainebya
Someofthem,suchasNeottiurabimaculata^changed single site in the middle ofplowed Dnieperlowland.
74 N.Polchaninovo
Thesecondspecificgroupswereassemblagesofsandy GribovaS.A,T.I.Isachenko&Y.M.Lavrenko (eds)
steppesinthenorthernsteppesubzonebecausethese (1980):[VegetationoftheEuropeanpartoftheUSSR].
fragmentsareisolatedanddifferinmicroclimaticcon- Nauka,Leningrad.432pp. [in Russian].
ditionsandvegetationstructurefromothergrasslands. Horvath R., T. Magura, C. Szinetar & B. Toth-
Steppes of a certain type are formed under both MERESZ (2009): Spiders are not less diverse in small
climate and soil conditions of the zone and local garnadssilsaonldast:edAgrfaiseslldansdtsu,dybu(tEalesstsHduivnegrasreyi,nNoyvierrsgerga).ze-d
topography (GrIBOVA et al. 1980). Accordingly, the Agriculture, Ecosystems and Environment 130: 16-22
structure ofthe animal communities ofthe steppes -doi: 10.1016/j.agee.2008.11.011
is a reflection ofthese conditions (MORDKOVITCH HASERS.E.&N.J.HUNTLY(2006):Thebiogeographyof
1982). Assemblages of herb-dwelling spiders are smallmammalsoffragmentedsagebrush-steppelands-
characteristicofeachsteppetypeanddifferinspecies capes. -Journal ofMammology 87: 1165-1174- doi:
composition and dominance structure. 10.1644/05-MAMM-A-385R2.1
KotenkoT.I. (1996): [Steppes ofUkraine: their impor-
Acknowledgements tance,modernstate,scientificvalue,protectionpriority].
The authorisgratefultoJ. Gergerichforrevisingthe first -Vestnikzooologii 1996(1/2): 10-26
draft ofthe English manuscript and to A. Atemasov for KRASNOBAEVYR(2003): [Structureofherpetobiontand
the map design. hortobiont spider populations in Samarskaya Luka].
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