Cytogenetical characterization of six orb-weaver species and review of cytogenetical data for Araneidae

2011·5.7 MB·English

by  Douglas Araujo

—
2011.TheJournalofArachnology39:337-344
Cytogenetical characterization ofsix orb-weaver species and review ofcytogenetical data for Araneidae
DouglasAraujo',VivianeF.Mattos^AndreMarsolaGiroti’^MarciaG.Kraeski-,LeonardoS.Carvalho^andAntonioD.
Brescovit^: 'Universidade Estadualde Mato Grosso do Sul-UEMS, Unidade Universitariade Ivinhema, Mato
Grossodo Sul, Brazil. E-mail: [email protected]; -Universidade Estadual de Mato Grossodo Sul-UEMS, Unidade
Universitariade Mundo Novo, Mato Grosso do Sul, Brazil; ^Universidade Estadual Paulista-UNESP, Institute de
Biociencias,DepartamentodeBiologia,RioClaro,SaoPaulo,Brazil;‘‘UniversidadeFederaldoPiaui,CampusAmilcar
FerreiraSobral, Piaui, Brazil; ^LaboratoriodeArtropodes, Institute Butantan, Sao Paulo, Brazil
Abstract. ThefamilyAraneidaeisthethirdlargestamongspidersandthethirdmoststudiedfromacytogeneticalpointof
view.Inspiteofthis,only2%ofallaraneidshavebeenkaryotyped.Themajorityofaraneidsanalyzedpossess2n = 24
chromosomesinmales;however,thestudyofadditionalspeciescouldrevealunusualkaryotypecharacteristics.Thus,the
aimofthisworkistoanalyzechromosomally,forthefirsttime,sixspeciesbelongingtothreearaneidgenerafromBrazil.
The specimensofAlpaida leucogramnui (White 1841), Alpaida tnmeata(Keyserling 1865), Alpaida veniliae (Keyserling
1865),Parawixiakochi(Taezanowski 1873),ParawLxkivelutina(Taezanowski 1878)and Wagnerianasp.werecollectedin
Barque Nacional de IlhaGrande and in the municipality ofRio Claro. The gonads were treated with colchicine and
hypotonicsolutionbeforefixationwithCarnoyIsolution.Theresultswere2nS=24(IIII+XiXt)inA.leucogramnuiand
P. velutina,and2nS = 22(10n+XiX2)inA. truncata,A. veniliae,P. kochiand Wagnerianasp.Whenthechromosomal
morphologieswereestablished, weobserved telocentricchromosomes in allspecimens saveone femalespecimen ofP.
velutina.Theunivalentsexchromosomeswereeasilyrecognizedondiplotenes.Theunpairedmetacentricelementfoundin
onefemalespecimenofP.velutinawith2n=25probablyarisesbycentricfusion/fission.Araneidaeisamegadiversefamily
composedof—3000speciesdistributedmainlyinthetropics;thustheanalysisofmorespeciesmayprovidenewinsights
aboutorb-weaverchromosomeevolution.
Keywords: Sexchromosomesystem,meiosis,mitosis,chromosome,spider
Despite the fact that the spider family Araneidae Clerck 2011).Thegenus Wagnerianacomprises41 speciesdistributed
1757isthethirdlargest,comprising3006speciesin 168genera mainly in Central and South America (Levi 1991; Platnick
(Platnick2011),only65speciesfrom20generahaveeverbeen 2011).
cytogenetically characterized (approximately 2% ofthe total Cytogeneticalstudiesofspidersarescarce,thushypotheses
diversity). Although relatively few araneid species have been regardingchromosomalevolutionaredifficulttoformulateor
studied,theyrepresentthethirdmoststudiedfamilyfromthe test.Theaimofthisstudyistocharacterizeforthefirsttime
cytogenetical point ofview. However, there are many gaps thespeciesA. leucogramma,A. truncata,A. veniliae,P. kochi,
remainingin ourknowledge ofthisdiverse group. Ofthe65 P. velutina, and Wagneriana sp. in relation to the diploid
speciesstudied,atleast50showadiploidcomplementof2nS number,chromosomalmorphology, andtypeofsexchromo-
= 24, with 22 autosomal elements and an X1X2 sex some system. Our study represents the first cytogenetical
chromosome system. The remaining species’ diploid number characterizationofNeotropical araneidspecies.
vsaerxiecshfrroommo2s«oc?me=s1y3sttoem2niSs =th4e9.mMoosrteocvoerm,mowhnilewitthheinX1tXhe2 METHODS
family,thetypeXoccurredwithasmallerfrequency,andthe Spiders were collected in the Parque Nacional de Ilha
X1X2X3andXYtypeswereeachobservedinonlyonespecies. Grande,Parana,Brazil,andinthemunicipalityofRioClaro,
Thechromosomalmorphologyformostspeciesisclassifiedas SaoPaulo,Brazil(Table2),between2008and2009.Afterthe
acro/telocentric(summarizedinTable 1). gonads were dissected, the specimens were deposited in the
The araneid genera Alpaida O. Pickard-Cambridge 1889, arachnological collection of the Instituto Butantan (IBSP,
Parawixia F.O. Pickard-Cambridge 1904, and Wagneriana curator1. Knysak)in thestateofSao Paulo, Brazil.
F.O. Pickard-Cambridge 1904, represent genera that have Chromosomal preparations and standard staining were
never been cytogenetically evaluated. The genus Alpaida is performedaccordingtoAraujoetal. (2008). Themitoticand
limited to Central and South America and comprises 140 meiotic cells were photographed using a digital-imaging
species (Platnick 2011). Alpaida leucogramma (White 1841) capture system coupled to a light microscope, and the
and Alpaida veniliae (Keyserling 1865) are distributed from chromosomal morphology wasclassified according to Levan
Panama to Argentina (Platnick 2011). Alpaida truncata etal. (1964).
(Keyserling 1865) is found from Mexico to Argentina (Levi RESULTS
1988;Platnick2011).ThegenusParawixiaincludes31 species,
whichoccuralmostexclusivelyinCentralandSouthAmerica. Alpaida. Theanalysesofspermatogonial metaphasesand
Parawixiakochi(Taezanowski 1873) is found fromTrinidad spermatocytes I of A. leucogramma showed 2n^ = 24 =
rtoecoBrrdaezdilfraonmdCPoalroamwbiixaiatoveAlrugtiennatin(aTa(eLzeavniow1s9k9i2; 1P8l7a8t)niciks 2le2n-1g-tXh1X(2Fitge.lolcAe)ntraicndch1r1omaoustoomseosmatlhabtivdaelcernetasseplgursadtuawlolyseixn
337
THEJOURNALOFARACHNOLOGY
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ARAUJOETAL.—CYTOGENETICCHARACTERIZATIONOFORB-WEAVERS 339
340 THEJOURNALOFARACHNOLOGY
1895
Simon
Araneidae
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ARAUJOETAL.—CYTOGENETICCHARACTERIZATIONOFORB-WEAVERS 341
—
Table 2. Listofspeciescytogeneticallyanalyzedinthepresentstudy,withthenumberofspecimens(malesandfemales)studied,numberof
cellsanalyzed(Cells),collectinglocationanddepositnumberofthespecimensinthearachnologicalcollectionoftheInstitutoButantan(IBSP).
ThenumberofspecimenofP. velutinawith2n = 25isinboldletters.
Species d $ Cells Collectionsite IBSP
Alpaidaleucogramma 1 30 CampusUNESP(22°23'33.97"S, 53028
47°32'37.39"W),RioClaro,SaoPaulo,
Brazil
Alpaidatruncata 2 - 107 SaoFranciscoIsland(24°00'32.40"S, 153928, 151186
54°09'52.18"W),Parana,Brazil
Alpaidaveniliae 1 1 27 Xambrelakemargins(23°52'48.64"S, 145318, 145319
54°0T2.25''W),Altonia,Parana,Brazil
Parawixiakochi 1 - 62 SaoFranciscoIsland(24°00'32.40"S, 123695
54°09'52.18"W),Parana,Brazil
Parawixiavelutina 3 3 348(224-2/7=24 SaoFranciscoIsland(24°00'32.40''S, 123717, 154838, 154847,
and26; 124-2n=25) 54°09'52.!8"W),Parana,Brazil 154844, 154842,
151176
Wagnerianasp. I 287 SaoFranciscoIsland(24°00'32.40"S, 152927
54°09'52.18"W),Parana,Brazil
univalents (llII+XiXo) (Fig. 2A). The sex chromosomes Xj bivalents, respectively (Fig.2D, E). Both species have two
and X2 are the smallest ofthe complement (Figs. lA, 2A). sexualunivalentsthatalwaysappearsidebysideandcorrespond
Diplotenes ofA. truncata (Fig. 2B) andA. veniliae(Fig. 2C) to the Xi and X2 chromosomes (Fig. 2D, E). Therefore, the
asnhdowetdwomeisoetxiucalfournmiuvlaaleentcsomp(rIiOsIiIn+gX11X02)auftoorsobmoatlhbisvpaelceinetss, dmiepilootiidcnfuomrbmeulraofof2«Pc.?=ko2c2h.iIinsPI.OvIeIl+utXi1nXa2t,hecomrerieostpiocnfdoirnmgultaoias
XainiudtioacsanotdimneXgs2aansdedixptlhuoeniidXva1nlXeun2mtbsseexralcwhoafryos2mnoaSspop=meeas2r.2,tIoncobAme.ptosrisudenedc-abyto-afsti2hd0ee co1oo1g1mo1p+nlieXaml1eXnm2te,tiawnpadhisacascteoisnngfoiafrmdteiwpdolobifydemctaohlemepsplroeefmseePnn.ctevoeflouf2tni2nd/at?=(F=2i4g..2T6IhBi)is.n
(Fig. 2B). In ail Alpaida studied, the sex chromosomes are Mitotic metaphases of one female specimen of P. velutina
positive heteropycnotic in a number of diplotene nuclei showedanastonishingcharacteristic;thediploidnumberis2«9
(Fig.2A-C). The majority of autosomal bivalents showed =25,withalargeunpairedmetacentricchromosome(Fig. 1C),
only one terminal chiasm, but bivalents with an interstitial contrastingwith the telocentric elements ofthe karyotype. In
chiasmalsooccurred(Fig. 2A-C).Afemalepachytenecellof both analyzed Parawixia species, the autosomal bivalents
A. veniliae revealed 12 chromosomal bivalents (Fig. 3A), showed only one interstitial or terminal chiasm. Differential
indicating the occurrence of 2«9 = 24, comprising 20 pycnosis was not observed in any chromosome of the
cauhtroosmoomseosmaalndmotrhpehXojloXg1yX2wXas2 sneoxtceshtraobmliosshoedmeinsAy.stetmr.uncTahtea cmeotmapplheamseentIIinnuclPeaursawwiaxsiafosupnedcieisn P(.Fikgo.ch2iD,,shEo)w.inOgnl«y=on1e0
andA. venilia—e,duetothelackofmitoticormetaphaseIIcells. autosomesand—nosexchromosomes(Fig. 3B).
Pamwixia. The observation ofspermatogonial diplotenes Wagnenana. The analyses of male diplotene cells in
of P. kochi and P. velutina showed 10 and 11 autosomal Wagneriana sp. revealed 10 autosomal bivalents and two
it;
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B 2 3 4 5 6 7 8 9 10 11 12 13
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c 1 2 3 4 5 6 7 8 9 10 11 12 13
Figures lA-C.—Karyotypes ofaraneid species. A. Alpaida leucogramma, with 2nd = 24 =22+XiX2 telocentric elements; B. Parawixia
tveelluotciennat,ri2chs9a=nd26on=e2u2np+aXir1Xed1Xme2Xta2cetneltorciecntcrhicrocmhorsoommoes,omperso;baCb.lyParreaswuiltxioafvaelcuetnitnrai,chfeutseiroonz.ygSoctaelesp=ec1i0mepnm,.2n9=25=2I+X1X1X2X2,with24
342 THEJOURNALOFARACHNOLOGY
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BW,aFgCin.geurArileapsnaai2dAsa-p.Ft,.niI—nOcIMaIat+alXea1Xndid2p.lAAolrtperanoiewdssaofivenandriialcinaaeteie,dirsnetpseeprcesitceitsti.ivaeAll.yc,AhlilpaOasIimIda+aXalineXduic.aorgDrr.oawmPmhaaer,aadw1si1xiainaudtikocoasctloeimia,tlerIbmOiiIvnIaa+lleXnc1tXhsi2pa.lsuEms.a.tPwSaocmawsleiexx=cihar1vo0epmlmou.tsionam,esII(1III+IXI+1XX12X.2F).;
sneuxmubalerunoifva2lnednt=s (2I2.OITIh+eX1aXu2t)os(Foimga.l2Fb)i,valseungtgsesptriensgenateddipolnoliyd a(2nnd A=. 2l2e)ucaongdraPm.mave{l2unti=na24{)2,na=s we2l4l)acsoubledtwbeeenusP.efuklochais
one terminal or interstitial chiasma. The sex chromosomes a cytotaxonomical character, which, together with genital
werepositiveheteropycnoticinrelationtotheautosomesand characteristics, can be used to identify these closely related,
always appeared closely associated (Fig. 2F). It was not sympatricspecies.
possibleto identifythechromosomal morphologydue to the The 2n^ = 22 diploid number has now been observedfor
lackofmitoticmetaphasesormetaphase II. the first time in Araneidae (see Table 1) for the species A.
DISCUSSION truncata, A. veniliae, P. kochi and Wagneriana sp. It is
interesting to note that Levi (1992), in a taxonomic review,
Theacquisitionandanalysisofcytogeneticdatamayshow indicatedthatthegeneraParawixia,Alpaida,and Wagneriana
more difficulties than the acquisition of morphological or shared (together with Acanthepeira Marx 1883, Wixia O.
molecular data; i.e., the specimens must be kept alive until Pickard-Cambridge 1882, Eriophora Simon 1864, Verrucosa
dissection and must present cells in division (Araujo et al. McCook 1888 and others) some external morphological
2005). However, molecular sequencing, despite becoming putative synapomorphies (see Levi 1992 for details). The
moreaccessible, is notavailable formany researchers out of presence ofspecies with 2n = 22 exclusively in these genera
the majorresearch centers. Thus, chromosomal data can be amongaraneidscould beconsidered furtherevidenceoftheir
used 1)asacomplementtomolecular/morphologicaldata,2) close relationship. Unfortunately, there is no cytogenetical
when molecular data are not available, or 3) when data on other closely related genera such as Acanthepeira,
morphologicalcharacteristicsarenoteffectiveforidentifying Wixia, Eriophora, Verrucosa and Edricus O. Pickard-Cam-
species (i.e., cryptic species). Moreover, the chromosome bridge 1890.
numberhasalreadybeenusedintaxonomicaldescriptionsof Ofthethreearaneidgenerastudiedhere, onlyAlpaidawas
spider species (see Maddison 1996). The differences in male included in the phylogenetic hypothesis of Scharff &
diploid number between A. truncata, A. veniliae {2n = 22), Coddington (1997), whose results do not corroborate the
hypothesis of Levi (1992), since the genera Acanthepeira,
Eriophora and Verrucosa appeared far from Alpaida, that
1 * j a1r8i8s4e,swEintahcinrocsloadmea63M,elwlhoi-cLheiatlasooi1n9c3l2u,desEuBsetratlraanSaiKmeoynser1l8i9n5,g
WixiaandAcacesiaSimon 1895(Scharff&Coddington1997).
» Of these genera, the only one that has a description of
* Jt cytogenetical data is Eustala, which possesses the most
common chromosome number in araneids, 2nS = 24 (see
A B Table 1). Thus, a more extensive cytogenetical study of
Araneidaerepresentativesisnecessaryinordertounderstand
A.FAilgpuariedsa3vAen-iBl.iae-Efaermallyeapnadchlyatteenmeewiiotthic12stbaigveasleinntsa,rianndeiicdatsipnigde2r«s.9 cthhercohmroosmoomseomnaulmbeevorlu{t2inon=p2r2o)ceisnsPtahratawriexsiulateadndinAtlhpeaisdmaa.ller
= 24. B. ParawixiakocliimalemetaphaseII,/! = 10chromosomes. Araneids, in general, have exclusively acro/telocentric
Scale = 10pm. chromosomes (Table 1). Exceptions are 1) two Argiope
ARAUJOETAL.—CYTOGENETICCHARACTERIZATIONOFORB-WEAVERS 343
ji Audouin 1826 specieswith 2nS = 24biarmed orholocentric alsoveryfrequentintheTheridiidaeSundeval 1833b,another
i chromosomes(Amalin1988;Amalinetal. 1992;Carandang& araneoidfamily(Araujoetal. 2010). Wealsoshowthateven
Barrion 1994a) and 2) one Larinioides Caporiacco 1934 and withinagenus(i.e.,ParawixiaandAlpaida),somespeciesare
three Neoscona Simon 1864 species that have karyotypes 2nS = 24, while others are 2nS = 22, suggesting that the
1;( composed of 2nS == 14 chromosomes, with 10 metacentric rearrangements involved in the conversion from 24 to 22
!> elements(Hackman 1948; Suzuki 1951b;Amalinetal. 1993). chromosomes or vice versa are relatively common among
AccordingtoSuzuki(1951b),thesecondcaseprobablywasa araneoidspiders.
'^ skuafrfyeorteydpceenwtirtihc2funsSio=ns2t4oafcoror/mtealokcaenrtyroitcypcehrwoimtohsIonmSes=th1a4t, orbT-hwuesa,vethrisssftorudythperefsiresnttsticmyet.ogTenheeticraelsuldtastaweorfeNesoltirghotpliycailn
with 10 metacentrics originated by the fusions of 20 aero/ disagreementwiththoserecordedforOldWorldspecies,and
I
telocentricelementsand4non-fusionedacro/telocentrics. furtherstudiesarerequiredtoevaluatewhethertheobserved
In the Parawixia velutina specimen with 2/j? = 25, the patternpersistsforotherNeotropical species. Thecytogenet-
I metacentricelement probably arose through a Robertsonian ical characterization ofrepresentatives ofthe genera closely
fusion (centric fusion), wherethecentromeric regions oftwo related to those analyzed in the present study {Parawixia,
acro/telocentric chromosomes fuse to form a single meta/ Alpaidaand Wagneriana), especially Araneines sensu Scharff
submetacentric chromosome, generating a heterozygosis & Coddington (1997), and Acanthepeira, Wixia, Eriophora,
condition. Frequently, chromosomal changes produce repro- VerrucosaandEdricus,assuggestedbyLevi(1992),isstrongly
ductivebarriers when theycause problems duringmeiosis in recommendedastheyseemto beessential forunderstanding
heterozygotes, leading to reduced fertility, owing to the chromosomeevolutioninaraneids.
formation of a trivalent during meiosis and unbalanced ACKNOWLEDGMENTS
segregation.However,duringmeiosissegregationbothunbal-
ancedandbalancedgametescouldbeformed(Sumner2003). The authors wish to thank Cristina Anne Rheims for her
InthecaseoftheheterozygotespecimenofP. velutina2«9= Englishreviewofthemanuscript.Thisresearchwassupported
25, the meiosis can proceed in two ways: 1) generate by Conselho Nacional de Desenvolvimento Cientifico e
unbalancedgametesthatareeliminatedduringgametogenesis, Tecnologico (CNPq-471821/2008-0- DA; 302148/2007-8 -
do not fertilize or produce unviable embryos; or 2) the ADB). Collecting permits were granted by the Instituto
fusioned metacentric pairs with their two homologous Brasileiro do Meio Ambiente e dos Recursos Naturais
telocentricstoformatrivalentandproperlygeneratebalanced Renovaveis - IBAMA and the Instituto Chico Mendes de
gametes,somecontainingthemetacentricandotherscontain- Conservagaoda Biodiversidade(ICMBio)(#15157-1).
ingthetwotelocentrics. Unfortunately,becausetheheterozy- LITERATURECITED
gote individual was a female, it was not possible to observe
meioticpairing. Amalin, D.M. 1988. Morphology,lifehistoryandcytologyofthree
hasCebneternicrefpusoirotnedisinamvoesrtygcroomupmsonofeovroglauntiisonmasry(Wchhiatnege197a3n)d. toLhraebg-uwnPeahai,lviitpnhpeginPsehpsii.ldieprPpshiin(leAis.prpainneease:UAnrivaenresiidtaye,) cLaoussinBganaorsa,neiCdoilslmegei,n
TshpiescimteynpeofofP.revealrurtainngaewmietnht,2n9a-s=pre25s,enwtlaysvoebrsiefrievdedininotnhee Amaclyitno,loDgy.Mo.f,AAr.gAi.opBearcrciitoennu&latLa.M(D.olReusecdhaal.l)19(9A2r.anMeoarep:hAroalnoegiydaaen)d.
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Evarchahoyi(Peckham&Peckham 1883)thatpresents2nS= Amalin, D.M., A.A. Barrion & M. Jayoma. 1993. Comparative
25 (with one unpaired metacentric autosome), contrasting karyomorphology oftwo Neoscona species (Araneae:Araneidae).
withthe2nS = 26(withoutmetacentricautosome)verifiedin PhilippineEntomologist9:1-6.
otherspecimensinthesamestudy(Maddison 1982). Inmany Araujo, D., A.D. Brescovit, C.A. Rheims & D.M. Celia. 2005.
cases, however, the homozygous condition rapidly reached Chromosomaldataoftwopholcids(Araneae,Haplogynae):anew
fixation,withtheoriginaltelo/acrocentricelementsdisappear- diploid number and the first cytogenetical record for the New
if2on«rg=tfhreo1m4prtedhveeisocpurosiplbuyeldcaitbtiyeodnHLa(arWcihkniimtoaeind1e9s(713a9)n4,8d)a,NsesSoeusezcmuoskniato(s1pb9ee5ci1tebhs)ewcaiantsdhe ArasWEupxojitrodr,leerdmDgec.el,ndauedeCgs.r.SeAJce.oyuotrfoRndhcaeehlsirmo(osfAm,roAasrnAaoe.camDehe.,nnoHluBarompegblsyeocorg3vy3vin:ata5rei9a&,1b-iS5lc9Diy6t.t.yModi.nidsCapeeel)ci.iae.Jsoou2fr0n0ta8h.le
Amalinetal. (1993). ofZoologicalSystematicsandEvolutionaryResearch46:89-95.
Ontheotherhand, GasteracanthahasseliiC.L. Koch 1837 Araujo, D., U.M. Maia & A.D. Brescovit. 2010. The first
and Gasteracantha kuhli C.L. Koch 1837 possess 2« = 16 characterizationofthepoisonousblackwidowspiderLatrodectiis
exclusivelyacro/telocentricchromosomes(Datta&Chatterjee, gr. curacaviensis from Brazil, with chromosomal review of the
1983, 1988).AccordingtoDatta&Chatterjee(1988),tandem/ familyTheridiidae(Arachnida,Araneae). Micron41:165-168.
centric fusion followed by pericentric inversion from a Berry,E.H. 1906.The“accessorychromosome”inEpeira.Biological
iknavroylovteydpeinwittheh o2r«icg?in=of24theacrkoa/rtyeoltocyepnetrwiicthch2rnom=os1o6m;easndi,s BolBeul-lGeotwinda1,1:B1.9N3.-210915.8.Astudyofthechromosomesduringmeiosis
owing to the pericentric inversions, there are no metacentric in twenty-two species of Indian spiders. Proceedings of the
cchhrroommoossoommeesnuinmbtehrerkeadruycotitoynp.e with 2n = 16, despite the CarcZaohnordloaomgnoigcs,aolmRe.SsoBc.ioef&tyArAogf.iAoB.peengBlaaurlzroi1no1an:.6(9W-a11l90c98k4..a).,Kaanryoortby-pweeavoifngthsepiedgegr
In this study we found four species with 2nS = 22, (Araneae,Araneidae). PhilippineEntomologist9:443^47.
suggesting that this diploid number could be more common Carandang, R.B. & A.A. Barrion. 1994b. Karyotype of egg
in araneids than previously thought. This diploid number is chromosomesofanorbweavingspiderArgiopeshillongensisSinha
344 THEJOURNALOFARACHNOLOGY
(Araneae:Argiopidae).Pp.32-33.InIntegratedPestManagement: Patau, K. 1948. X-segregation and heterochromasy in the spider
Learning from Experience. Pest Management Council of the Araneareaumiiri.Heredity2:77-100.
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ChromosomeInformationService43:11-13. Manuscriptreceived1 October2010,revised10April2011.