Diversity of Chrysomelidae (Coleoptera) at a mountain range in the limit of the Eurosiberian region, northwest Spain: species richness and beta diversity

2007·0.89 MB·

by  Andrés Baselga

©EntomologicaFennica.6June2007
Diversity of Chrysomelidae (Coleoptera) at a mountain range
in the limit of the Eurosiberian region, northwest Spain:
species richness and beta diversity
AndrésBaselga&FranciscoNovoa
Baselga, A. & Novoa, F. 2007: Diversity of Chrysomelidae (Coleoptera) at a
mountainrangeinthelimitoftheEurosiberianregion,northwestSpain:species
richnessandbetadiversity.—Entomol.Fennica18:65–73.
ChrysomelidaefromtheSierradeQueixamountains(Galicia,northwestSpain)
weresampled,reporting93species.Theestimatedlocalspeciesrichnessusing
severalnon-parametricestimatorsandaccumulationmodelsvariesbetween104
and142species. TocomparetheChrysomelidaefaunafromSierradeQueixa
withotherGalicianassemblageswehaveassessedbetadiversityamonginvento-
ries and we have tested the differences on the zoogeographic compositions
amongareas.SierradeQueixaisgroupedwithotherGalicianmountainranges
locatedinthetransitionzonebetweenEurosiberianandMediterraneanregions,
anditischaracterisedbyahighproportionofIberianendemicspecies,signifi-
cantly higher than expected. Therefore, Chrysomelidae fauna from Sierra de
QueixarepresentsauniqueassemblageofEurosiberianelementsthatreachedthe
areaduetotheclimaticconditionsandIberianendemicspeciesthatarepresentin
theIberianmountainsduetoitshistoricroleassouthernrefugiaduringglaci-
ations.
A.Baselga.DepartamentodeBiodiversidadyBiologíaEvolutiva,MuseoNaci-
onaldeCienciasNaturales–CSIC,c/JoséGutiérrezAbascal,2,28006Madrid,
Spain;E-mail:[email protected]
F.Novoa.DepartamentodeBioloxíaAnimal,FacultadedeBioloxía,Universi-
dade de Santiago de Compostela, 15706 Santiago de Compostela, Spain; E-
mail:[email protected]
Received4May2005,accepted2May2006
1.Introduction tory(Baselga&Novoa2006)aswellastoassess
betadiversity amonglocalfaunas,includingin-
Knowledge about Chrysomelidae (Coleoptera) ventoriesfromcoastalenvironmentstomountain
fromGalicia(northwestSpain)hasbeennotably ranges (Baselga & Novoa 2005). In this scope,
increasedinthelastyearswiththestudyofsev- westudyheretheleafbeetlefaunafromSierrade
eral local faunas (Baselga & Novoa 2000a, Queixa (Ourense, Spain) (Fig. 1), an area in-
2000b,2002a)andthepublicationoftaxonomic cluded in the Eurosiberian phytogeographic re-
papers(Baselga&Novoa2000c,2002b,2002c, gion(Izco1987,Rivas-Martínez1987)butadja-
2003,2004).Theaccumulationofsuchdatahave cent to the limit of the Mediterranean region,
allowedtoestimatetheregionalspeciesrichness searchingfortheunrecordedtaxathatwerepre-
andthusthecompletenessoftheGalicianinven- dictedtobepresentinGaliciabyseveralestima-
66 Baselga&Novoa (cid:127) ENTOMOL.FENNICAVol.18
Fig.1.LocationofthestudiedareainGalicia,northwestoftheIberianpeninsula.–a.Galicia(shadowed)islo-
catedintheboundarybetweenEurosiberianandMediterraneanphytogeographicregions(discontinuousline,af-
terRivas-Martínez1987).–b.LocationofSierradeQueixa(Que)andotherareasselectedfortheanalysisof
betadiversity.Thesquaredpatterncorrespondstothe10kmUTMgrid(zone29T,100kmsquaresMH,NG,NH,
NJ,PG,PHandPJ)andthecontinuouslineistheboundarybetweenphytogeographicregions(Izco1987,
Rivas-Martínez1987).Eachinventoryiscomposedofseverallocalitiesmarkedwiththesamesymbol.SeeMa-
terialsandMethodsforfurtherexplanation.
tionmethods(Baselga&Novoa2006)andsur- terraneanregion.Therefore,inthisareathereisa
veyinganewareainordertoimprovethedescrip- high turnover, with Eurosiberian vegetation re-
tion of turnover patterns among Galician local placingMediterraneanoneinasmallextent.
faunas(Baselga&Novoa2005).Ourstudywas An extensive sampling in Sierra de Queixa
thefirstextensiveoneinSierradeQueixa. was carried out by authors in 2002 and 2003,
Theaimsofthispaperare(i)todescribethe though some previous captures are also consid-
local species richness of this highly diverse ered in this paper. Twenty localities were sur-
group,(ii)toestimatethecompletenessofthein- veyed (Fig. 1). They were selected in order to
ventory,assessingtheaccumulationcurvegener- rangetheentirealtitudinalgradientandthemain
atedfromoursampling,and(iii)tocomparethis vegetationunits.Atotalof789specimenswere
faunawithotherGalicianones,analysingbetadi- collectedbysweepingandbeatingthevegetation.
versity among inventories and its differences in Samplings were not standardized, and we col-
the proportions of chorotypes contributing to lectedallthespeciesdetectedineachlocalitybut
eachlocalfauna. notallthespecimensofeachspecies.Allthema-
terialisdepositedinA.Baselgacollectioninthe
DepartamentodeBiologíaAnimal,Universidad
2.Materialandmethods deSantiagodeCompostela,Spain.
To assess the completeness of the inventory
Sierra de Queixa forms part of the Galician bothasymptoticmodelsandnon-parametricesti-
MacizoCentral,agroupofmountainrangeswith mators(ICE,Chao2andjackknifeoffirstorder
summits up to 1,700 m located in the Spanish and second) were used (Colwell & Coddington
province of Ourense (Galicia). The range is in- 1994).Weusedonlyincidence-basedestimators
cluded in the Eurosiberian phytogeographic re- sincenotallthespecimenssawnwerecollectedin
gion(Izco1987,Rivas-Martínez1987)butSierra eachsamplingandthusnoabundancemeasures
deQueixaisadjacenttothetemperatevalleysof are available. Among them, we selected those
SilandBibeirivers,bothbelongingtotheMedi- recommendedbeHortaletal.(2006).Theaccu-
ENTOMOL.FENNICAVol.18 (cid:127) ChrysomelidaeinlimitofEurosiberianregion 67
mulationcurvesandthenon-parametricestima- isthetotalnumberofspeciesthatoccurinthesec-
torsweregeneratedwithEstimateS7.0software ondareabutnotinthefirst.Thisindexwasse-
(Colwell 2004), randomising the sample order lectedbecauseitfocusesoncompositionaldiffer-
100times.Databaserecords(Soberónetal.2000, encesindependentofspeciesrichnessgradients,
Hortal et al. 2001, Martín-Piera & Lobo 2003) andthusitisa‘narrowsense’measureofturnover
and individuals were used as a sampling-effort (Koleffetal.2003).Adistancematrixwascon-
surrogate. Our database comprised 270 records structed with b for each pair of faunas, and
sim
for 789 specimens. Each record is composed of thereafterusedinaanonmetricmultidimensional
thefollowingfields:speciesname,locality,date, scaling (NMDS) (Legendre & Legendre 1998),
number of specimens and collector. Any differ- generated with Statistica 6.0 (StatSoft 2001).
ence in any database field value gives rise to a Three dimensions were considered because the
newdatabaserecord,soincrementsofthenumber screeplotshowedasmallchangeofthestressval-
ofrecordsprovidecorrelativeincrementsofthe ues with dimensionality increments after this
sampling effort (Martín-Piera & Lobo 2003). point (Legendre & Legendre 1998) (stress =
Thereafter, the asymptotic Clench function was 0.0552).Thereafter,thematrixforthesethreedi-
fittedtothesmoothedcurve(Soberón&Llorente mensionsversusthenineinventorieswasusedin
1993,Hortaletal.2004,Jiménez-Valverdeetal. a cluster analysis, in order to recover the major
2006): groupsoflocalfaunas.Thetreewasconstructed
with Clustan Graphics 6.03 (Clustan 2003),
S =ae/(1+be) (1) weightingthethreevariablesaftertherespective
(e)
decreaseofstressexplainedbyeachdimension,
whereS isthenumberofspeciesfoundpersam- andthengroupingthefaunasusingEuclideandis-
(e)
plingeffortunite;aandb,theparametersofthe tanceandsinglelinkage.
function.Theselatterparameterswereadjustedto Biogeographictypesusedinthispaperwere
thedataofthecurvebymeansofaSimplexand synthesized following the chorotypes proposed
Quasi Newton method (StatSoft 2001). The as- byVignaTagliantietal.(1992)andthengroup-
ymptote(i.e.thepredictednumberofspecies)is ingthemintofourmajorcategories:Iberianele-
calculatedasa/b. ments(Ibe),forspeciesendemicfromtheIberian
For the comparison of the present inventory Peninsula; Mediterranean elements (Med), for
with other Galician faunas we have considered species widespread in the Mediterranean coun-
thepreviouslypublisheddata(Baselga&Novoa tries; Eurosiberian elements (Eur), for species
2006),takingintoaccountalltheavailableinven- widespreadinEurope,orEuropeandtheSiberian
tories: coastal dunes and associated marshes range;andfinallywiderangeelements(WR),for
(Dun), agricultural landscape near Santiago de specieswidespreadinalloragreatpartofthePa-
Compostela(San), Atlanticmixed forestoflow laearctic region, and reaching parts of both
altitudeinFragasdelEumeNaturalPark(Eum), Eurosiberian and Mediterranean areas. These
medium altitude mountain ranges of Dorsal majordivisionsareestablishedinordertomake
Gallega(Dor)andLarouco(Lar),thetemperate clearthecontrastbetweenEurosiberian(septen-
valley of the Sil river (Sil) and high mountain trional) and Mediterranean (meridional) contri-
ranges of Ancares (Anc) and Eixo-Segundera butionstoGalicianfaunawhichislocatedacross
(Seg). The present inventory is abbreviated as the Eurosiberian-Mediterranean boundary. The
Quefortablesandfigures.Simpson’sindex(b ) othertwocategoriesareneutralregardingthisas-
sim
wasusedasbetadiversitymeasure(Koleffetal. pect, because almost all Iberian elements are
2003): presentinbothsidesofEurosiberian-Mediterra-
neanlimit,aswellasWRelementsreachbothre-
b =min(b,c)/min(b,c)+a (2) gions.
sim
To test the statistical significance of differ-
whereaisthetotalnumberofspeciesthatoccur encesonbiogeographicpatternsamonginvento-
inbothareas,bisthetotalnumberofspeciesthat riesachisquareanalysis(¤2)wasperformed.The
occurinthefirstareabutnotinthesecond,andc contingencytablewasconstructedwiththefour
68 Baselga&Novoa (cid:127) ENTOMOL.FENNICAVol.18
Fig.2.Speciesaccumula-
tioncurvesgeneratedfrom
thefieldstudyinSierrade
Queixawiththefitted
Clenchfunction,thenon-
parametricestimatorsICE,
Chao2andfirst-orderjack-
knifeandthenumberof
singletonsanddouble-
tons.–a.Curvebasedon
theaccumulationofdata-
baserecords.–b.Curve
basedontheaccumula-
tionofindividuals.
biogeographic categories (columns: WR, Eur, 3.Results
Med,Ibe)versusthenineinventories(rows:Dun,
San, Eum, Dor, Lar, Sil, Anc, Seg and Que). Atotalof93speciesofChrysomelidaehavebeen
Probabilityconsideredtorejectthenullhypothe- collectedinSierradeQueixa.Thefulllistofspe-
sis (differences are due to random chance) is cies and exact sampling localities are available
95%.Thedegreesoffreedomare1forthecells,3 fromtheauthorsonrequest.Theleafbeetlefauna
fortherowsubtotals,8forthecolumnsubtotals inSierradeQueixaismostlycomposedofWRel-
and24forthewholecontingencytable,andthe ements(43.0%).TheproportionsofEurandMed
critical ¤2 values 3.8415, 7.8147, 15.5073, elements are equal (16.1%), and the Ibe ende-
36.4150respectively misms reach near a quarter of the inventory
ENTOMOL.FENNICAVol.18 (cid:127) ChrysomelidaeinlimitofEurosiberianregion 69
Table1.Estimatedspeciesrichnessusingdatabase
recordsandindividualsassamplingeffortsurrogate.
Estimate Recordscurve Individualscurve
Clench 142 105
ICE 115 105
Chao2 115 104
Jacknife1 124 111
Jacknife2 135 116
(24.7%).Amongthecitedtaxa,6speciesarespe-
ciallyinteresting:PsylliodescervinoiBaselga&
Novoa is firstly recorded here since its descrip-
tion from Sierra Segundera (Baselga & Novoa
2003), extending its known distribution to
Queixamountainrange,whereasStylosomusmi-
nutissimus(Germar,1823),Pachybrachissuffri-
aniSchauffus,1862,Galerucainterrupta(Illiger,
1802),Luperuslongicornis(Fabricius,1781)and
PsylliodesobscuroaeneusRosenhauer,1856are
new recordsfor Galicia, increasing theregional
inventoryfrom276to281species.
The completeness of the present inventory
wasevaluatedbymeansofaccumulationmodels Fig.3.ComparisonofleafbeetlesfromSierrade
QueixawithotherGalicianlocalfaunas.–a.Non-
and several non-parametric estimators, using
metricmultidimensionalscaling(NMDS)basedon
both database records and individuals as sam-
Simpson’smeasureofbetadiversity(stress=
plingeffortsurrogates(Fig.2).Ourresultsshow
0.0552).–b.Dendrogramfromtheclusteranalysis,
thatatotalrichnessbetween104and142species
constructedafterthematrixforfaunasvs.dimensions
couldbeexpectedinthestudiedarea,depending producedbytheNMDS.Euclideandistanceandsin-
onthecurveassessedandtheestimatorselected glelinkagewereused.
(Table1).Thereforebetween65%and90%ofthe
estimated number of species of Chrysomelidae
livinginSierradeQueixaseemstobedetectedin becausethescreeplotshowedasmallchangeof
ourfieldstudy,butweshouldexpectthatbetween thestressvalueswithdimensionalityincrements
12and49taxawillbeaddedtothisinventoryin afterthispoint(Legendre&Legendre1998).Ma-
thefuture.Moreover,inbothcurvestheslopeat jorgroupsofGalicianfaunaswererecoveredbya
theendofthesurveyisquitelow:0.1195using cluster analysis, using the matrix for faunas vs.
recordsand0.0156usingindividuals.Theseval- dimensionsproducedbytheNMDS.Thedendro-
uesimplythatmorethan8recordsor64individu- gram constructed from this matrix shows three
alsshouldbeaddedtotheinventory,andthusthe majorfaunisticgroups(Fig.3b).Theleafbeetle
completeness of the inventory is quite high fauna from Sierra de Queixa is grouped with
(Jiménez-Valverde & Hortal 2003, Hortal et al. thoseofothermountainrangeswhicharelocated
2004). in the trasition zone between Eurosiberian and
TocomparetheleafbeetlefaunafromSierra Mediterranean regions (Ancares, Larouco and
deQueixawiththoseofotherGalicianareaswe Eixo-Segundera). The other two clusters were
haveassessedbetadiversity amonginventories, alsofoundelsewhere(Baselga&Novoa2005):
consideringthepresencedatareportedinBaselga ononehand,thelowandmediumaltitudeareas
andNovoa(2006).TheNMDS(Fig.3a)wasper- closertotheAtlanticcoast(coastaldunes,Santi-
formed with three dimensions (stress=0.0552) ago,EumeNaturalParkandDorsalGallega)and,
70 Baselga&Novoa (cid:127) ENTOMOL.FENNICAVol.18
Table2.Contingencytableforthebiogeographicpatternsversusthelocalinventories.Significantdifferencesare
notedinboldface.Observedandexpectedvaluesarenotedas(O)and(E),respectively.¤2totalvalueforthe
wholetable=49.19.
Corotype WR WR¤2 Eur Eur¤2 Med Med¤2 Ibe Ibe¤2 Sub- Subt.¤2
total
Dun(O) 72 11 22 8 113
Dun(E) 59.5 2.6 18.2 2.9 18.5 0.7 16.8 4.6 10.8
San(O) 66 16 10 12 104
San(E) 54.7 2.3 16.8 0.0 17.0 2.9 15.5 0.8 6.0
Eum(O) 43 18 9 17 87
Eum(E) 45.8 0.2 14.0 1.1 14.2 1.9 12.9 1.3 4.5
Dor(O) 45 14 13 11 83
Dor(E) 43.7 0.0 13.4 0.0 13.6 0.0 12.4 0.2 0.2
Lar(O) 40 11 12 10 73
Lar(E) 38.4 0.1 11.8 0.1 11.9 0.0 10.9 0.1 0.2
Sil(O) 50 10 30 13 103
Sil(E) 54.2 0.3 16.6 2.6 16.9 10.3 15.3 0.4 13.6
Anc(O) 69 31 20 21 141
Anc(E) 74.2 0.4 22.8 3.0 23.1 0.4 21.0 0.0 3.8
Seg(O) 35 15 12 15 77
Seg(E) 40.5 0.8 12.4 0.5 12.6 0.0 11.5 1.1 2.4
Que(O) 40 15 15 23 93
Que(E) 49.0 1.6 15.0 0.0 15.2 0.0 13.8 6.1 7.7
Subt.(O) 460 8.31 141 10.26 143 16.2 130 14.4 874 49.2
ontheother,thefaunafromtheSilValleythatis tion of the free living phytophagous functional
themostdifferentone. groupwithinarthropods.Thestudyofleafbeetles
Thecomparisonofthebiogeographiccompo- fromSierradeQueixahasrevealedaveryinter-
sitionsofthelocalinventoriesshowssimilarre- esting local fauna, quite singular among the re-
sults(Table2).Thecolumnsubtotal¤2forSierra gionalassemblagesduetothestrikinghighpro-
deQueixaisquitehigh(closetothecriticalvalue portionofIberianendemics(24.7%)(Fig.4,Ta-
butnotsignificantatp<0.05)almostcompletely ble 2) or the citation of several taxa new for
duetothelargeproportionofIberianendemics, Galicia. Theadditionoffivenew speciestothe
which is significantly higher than expected by regional inventory may be considered valuable,
chance. Only two inventories are significantly since completeness of Galician Chrysomelidae
differenttotheothers:Silvalley,dueespeciallyto listwasalreadyveryhigh,between85and95%
its high proportion of Mediterranean elements of the estimated richness (Baselga & Novoa
andcoastaldunesduemainlytoitssmallpercent- 2006).Thissaturationismoredifficulttogetata
ageofIberianendemisms.Thesedisparitiespro- smaller scale because of the great number of
duceahightotal¤2valueforthewholetable,im- samplingsthatwouldberequiredforeachlocal
plying significant differences among the com- area,andthusadegreeofcompletenessbetween
paredfaunas. 65and90%maybeconsideredacceptable,con-
sideringthatweareworkingwithahighlydiverse
group of invertebrates. Therefore, further sam-
4.Discussion pling effortin SierradeQueixawould add new
taxatotheinventory(between12and49taxaaf-
Chrysomelidaeconstituteaninterestinggroupto ter our estimates) but current knowledge of
develop biodiversity estimates due to two main Queixaleafbeetlesisalreadysuitableinorderto
reasons:ontheonehanditisahyperdiversefam- describe the local assemblage and the regional
ilyand,onother,theyprovideagoodrepresenta- biodiversitypatterns.Comparingthedifferentes-
ENTOMOL.FENNICAVol.18 (cid:127) ChrysomelidaeinlimitofEurosiberianregion 71
Fig.4.Percentagesof
chorotypesfoundin
eachlocalfauna(see
MaterialsandMethods
forabbreviationsof
names).Numbers
aboveeachbararethe
observedspeciesrich-
ness.Ibe,Iberianele-
ments;Med,Mediterra-
neanelements;Eur,
Eurosiberianelements;
WR,widerangeele-
ments.
timates (Table 1), the maximumvalue is recov- found. Queixa is grouped together with other
eredadjustingtheClenchfunctiontotheaccumu- mountain ranges located in both sides of the
lationcurveusingdatabaserecordsassampling Eurosiberian-Mediterranean phytogeographic
effortsurrogate(142),whereastheminimumone boundary (Izco 1987, Rivas-Martínez 1987).
isprovidedbytheChao2estimatorintheindivid- This group was previously recovered in similar
uals curve (104). In all cases, estimates of the comparisonperformedwithoutincludingpresent
curvebasedonindividualsarelowerthanthose data(Baselga&Novoa2005)andtheadditionof
yieldedbythecurvebasedonrecords,butthedif- Queixainventorytothisclusterisnotsurprising
ference between the Clench asymptotes of both taking into account its location. The conclusion
curves is the most remarkable discrepancy. Ex- thatfaunisticisolationofSilValley(Figs.3and4)
aminationofthefrequencyofrecordsperspecies isduetoenvironmentalfactors(i.e.temperature,
inthecurvebasedonrecordsshowsanimportant pluviosity),becausenogeographicalbarriersare
number of species with 4 or less records (not interposed, is here newly supported. Sierra de
shown).Thiscausesthecurvetooverestimateto- QueixaandSilValley arestrictly contiguousas
talspeciesnumber.Suchdrawbackislessimpor- wellasSilValleyandEixo-Segundera.Alsothe
tantwhenusingindividuals.Inthiscurve,values clustering of two faunas located in the phyto-
ofClenchasymptoteandnon-parametricestima- geographic Mediterranean region (Larouco and
tors are more similar. More research on the be- Eixo-Segundera)withtheEurosiberianinvento-
haviour of species accumulation curves is ries is again corroborated. Hence, it should be
needed,becauseinourdatasetthenumberofsin- emphasizedthatalthoughtheLaroucoandEixo-
gletons and doubletons is considerable, so non- Segundera ranges are included into the phyto-
parametric estimators should be overestimating geographic Mediterranean region together with
therichnessvalue,andthusitisstrikingthatas- theSilValley,onlythelatterareahasanunambig-
ymptotic model yield grater values, contrary to uous Mediterranean-type Chrysomelidae fauna
theresultsfoundbyHortaletal.(2006). (Fig.5).ThisconclusionwaspointedinBaselga
When the Chrysomelidae fauna from Sierra &Novoa(2005)butitisstatisticallytestedhere
deQueixaiscomparedwithotherGalicianinven- for the first time. The contingency table shows
toriesanalysingbetadiversityandorderinglocal- significantdifferencesinthedistributionsofbio-
itiesbymeansofaNMDS,threemajorgroupsare geographiccategoriesamonglocalfaunas(Table
72 Baselga&Novoa (cid:127) ENTOMOL.FENNICAVol.18
2). In the case of Sil valley Mediterranean ele- fauna, unique among regional assemblages.
mentsaresignificantlyhigherthanexpected. Moreover, the addition of this study to the re-
Thezoogeographicalzonationisdifferentfor gionalknowledgehasallowedustoreassessleaf
each group considered, but as a general pattern beetlebetadiversityintheregion.Thenewdata
Eurosiberian faunas occupy areas that Rivas- supporttheconclusionsreportedinourstudyof
Martínez (1987) includes in the Mediterranean thefaunisticheterogeneity ofGalicianleafbee-
regioninhisclassificationofIberianvegetation,a tles(Baselga&Novoa2005)anditshouldbeas-
patternalsopointedoutbyHermidaetal.(1994) sumed that the described patterns are robust.
forwesternIberiangastropods,Rodríguezetal. They show that zoogeographic distribution of
(1997) for western Iberian earthworms and ChrysomelidaefaunasinnorthwesternSpaindo
Hortal-Muñozetal.(2000)forPortuguesedung not fit exactly the standard phytogeographic
beetles.AsWilliams(1996)comments,thearbi- zonation(Izco1987,Rivas-Martínez1987).Only
trary “border lines” between biogeographic re- Sil Valley has an unambiguous Mediterranean-
gionsarenotrealisticrepresentationsoftheeco- typefauna,whereasmountainrangesofLarouco
logicalreality,becausetheycanbewiderthanthe and Eixo-Segundera, though located in the
grainusedfortheirdelimitation.Infact,itseems phytogeographic Mediterranean region, are
thatmorethanasharpboundary,thetransitional groupedwithEurosiberianleafbeetlefaunas.
zoneiscomposedbyamosaicofleafbeetlefau-
nas depending on the local climatic conditions. Acknowledgements. We are grateful to S. Barallobre, I.
Theseenvironmentalvariablesaremostlydeter- Gañán,J.Novoa,R.NovoaandM.Santiagofortheircol-
laboration on several samplings. We wish to thank two
minedbythecomplicatedregionalgeomorphol-
anonymousrefereesfortheirinterestingsuggestionsthat
ogy. Therefore, mountains located south of the
greatlyimprovedthispaper.Thisresearchwassupported
boundary (and then included in the Mediterra- by Xunta de Galicia with project PGIDT01MAM
neanregion;i.e.Larouco,Eixo-Segundera)host 20001PR.
Eurosiberian faunas, whereas temperate valleys
locatednorthtotheboundaryshelterMediterra-
nean faunas (i.e the Navia valley in Ancares
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