HERPETOLOGICALJOURNAL, Vol. 11,pp. 129-136(2001)

SPECIESDISTINCTION AND RELATIONSHIPSOF THE WESTERN IBERIAN PODARCISLIZARDS (REPTILIA, ) BASED ON MORPHOLOGY AND MITOCHONDRIAL DNA SEQUENCES D. J. HnnnISI ANDP. SA-Sousa, t Centro de Estudos de Ciancia (cECA/uP), campus Agrdrio de vairdo, portugal 2Centro de Ecologia Aplicada, Universidade de Evora, 7002-544, Evora, portugal

Wall () arethe dominant group acrossmost of southernEurope. Their is complex becausemost speciesexhibit substantialintraspecific morphological polymorphisms.We have estimatedthe phylogeny of the particularlydiverse western Iberian forms using partial cytochrome oxidase and cytochrome b mitochondrial DNA sequencedata and have compared this against morphological variation. Of the two currently recognized speciesin the area- P odarcis hispanica andP. bocagei -neither is monophyletic, and extremely high geneticdiversity betweennewly identifiedforms (up to l5%cytochrome b divergences) indicatesthatbotharespeciescomplexes.Podarcisb bocageiisgeneticallydistinctfromP.(b) carbonelli which appearsto be a separatespecies using both mtDNA andprotein electrophoretic data. The insular form previously assigned to P b. berlengensis, and sometimes argued to deservespecies status is not genetically distinct from P. (b ) carbonelli using the mtDNA sequences P hispanica can be separatedinto at least four highly divergent groups, two in western Iberia, one in easternIberia and one in North Africa.

Key words: phylogeny, cytochromeb, cytochromeoxidase, morphology, Iberian lizards

INTRODUCTION (S6-Sousa,2000a). P. hispanicatype I has the follow- ing On the Iberian Peninsulathree speciesof insectivo- characteristics:flattened head and body; either reticulated rous wall lizards have been recognized- the Iberian or shiped dark dorsalpatterns; and whitish- pearly (for wall , Podarcis hispanica Steindachner 1870; coloured belly details see Pdrez-Mellado, 1981a,b; Bocage's wall lizard, Podarcis bocagei Seoane1884; Gal6n, 1986; Pdrez-Mellado and Galindo, 1986 Guillaume, 1987 and Podarcis muralis Laurenti 1768. Podarcis bocagei ; ; Sii-Sousa,199 5 a). P. hispanica (SW and P. hispanicalive in sympatry in large areasof NW Ape 2 Iberian form) has been found Iberia (Gal6n,1986;Pdrez-Mellado & Galindo,1986; in Andalusia,in Extremadura,in the Madrid re- gion, and in Sii-Sousa,1995a). Both exhibit pronouncedsexual di- the western and southern parts of Portugal (Sii-Sousa,2000a).In morphism, adult males being larger than females and Portugal, P. hispanica type 2 prefer (<400 having more intense colour patterns (Gal6n, 1986; seemsto lowlands m) with a Mediterra- nean climate (Sd-Sousa, Pdrez-Mellado& Galindo, 1986). 2000a). This form has the following characteristics: Podarcis hispanica is a medium sized (SVL 65-70 head and body moderately robust; green and/or patterns, mm), morphologically variable rock-dwelling lizard, light brown and yellow- orange belly (see Klemmer, which is found in SW France(Langedoc-Rousillon and 1957; Salvador, 1986; Guillaume, 1987; Cdvennes),the Iberian Peninsula(except the northern- GonzLlez de la Vega, 1989; Srl- Sousa,1995a). most corner) and NW Africa (Galdn, 1986;Guillaume, Given severalrecords of P. hispanica vaucheri in SW Iberia (Boulenger, 1987, 1997). Despite taxonomic controversyabout P. 1905; Klemmer, 1957;Salvador, hispanicasubspecies, two forms have beenrecognized 7974, 1986;Busack, 1986; Guillaume, 1987) one might in Portugal (S6-Sousa1995a,2000a). First, there is a hypothesizethat morphologically P. hispanica NW Iberian form (P. hispanica type 1) that resembles type 2 and P. hispanica vaucheri are the same. However, retain the "lusitanica" form of Guillaume (1987).P. hispanica we the P. hispanica type 2 de- 'Submeseta nomination until further type I is found in Galicia, in the Norte' studies confirm whether similarity in phenotype plateau, in the northern half of Portugal and on the corresponds to a common 'Sistema genofype across Central' mountain range (S6-Sousa,2000a). the entire range. So far all allopatric greenishmorphotypes wall lizard In Portugal,P. hispanicatype I seemsto inhabitmainly of found in the south- ern part of Iberia (e.g. Portugal, highlands(>400 m) where eitherAtlantic or continental southern Andalusiaand Levant) have climatic conditions may prevail. It has been found in been considered as P. h. hispanica (P6rez-Mellado,1998). the northern part of Portugal, north of the Tagus river Also P. hispanicatype 2 is of- ten mistakenfor P. b. bocagei,because oftheir greenor light-brown chromatic pattems, although the Correspondence:D.J. Harris, Centro de Estudosde Ci€ncia Animal (CECA/UP), CampusAgrario de Vair6o, 4480 Vila show severaldifferences and have allopatric distribu- do Conde, Portugal.E-mail:[email protected] tions (S6-Sousa,199 5 a, 1998,2000 a). 130 D. J. HARRIS AND P. SA-SOUSA

Podarcisb. bocageiis a medium-sized(adult snout- There,the type of climate- but alsothe balancebe- vent length,SVL:65-70 mm), ground-dwellinglizard, tweenthe numberof frost daysper yearand the degree the malesof which show greendorsal patterns, while of aridity- appearimportant to explainthe distribution femalesare brown with a pair of greenstripes (Gal6n, of P. b. carbonelli(S6-Sousa, in prep.). 2000; Pdrez-Mellado, l98la,b, 1986; S6-Sousa, Morphologicalcharacters and the existenceof 1995a).This speciesis an Iberian-Atlanticendemic oc- parapatriczones of contactbetween different type s (i.e. curring in W Asturias,Cantabria, Galicia and north of withoutinterbreeding) suggested that P. bocageiand P. Portugal(Gal6n, 1986,1997; S6-Sousa, 1998). On a hispanicamight in fact be speciescomplexes (56- coarsescale, the distribution of P. b. bocageican be Sousa,2000a and in prep.).To resolverelationships largely explainedby macroenvironmentalvariables betweenand within these groups, and to determine andtype of climate(S6-Sousa, 2000b). whetherrecognized clades based on morphological It has been suggestedthat P. b. carbonelli P6rez- charactersare geneticallydistinct, a phylogenetic Mellado1981 merits species distinction (S6-Sousa, in analysiswas conductedusing DNA sequencesderived prep.;S6-Sousa et a|.2000).P. b. carbonel/t(SVL:50- from two mitochondrialgenes, cytochrome 6 andcyto- 55 mm) is a small, green ground-dwellinglizard, chromeoxidase 1. Populationsacross the rangeof the initially thoughtto be restrictedto the W SistemaCen- acceptedsubspecies of P. bocageiwere sampled. For tral range(P6rez-Mella do, l98 l a,b, l 986).However, it the cytochromeb datasets previously published se- hasbeen found in othermountain systems as well as quencesof P. hispanica from easternSpain and alongthe Atlantic lowlands,particularly in Portugal Morocco(Castilla et al.,1998;Hanis & Arnold,1999) (Magraner,1986; S6-Sousa,1995b, 1999, 2000b). wereincluded in the analyses.

TABLE L List of lizardsexamined in themtDNA phylogeny.* indicatespreviously published data was included in theanalysis. Sequencesare deposited in Genbank(accession numbers AF372051 to AF372089).Map codesare shown in Fig. l.

Species mtDNA Code Locality COl/ Cyt 6 Map Code

Gqllotia galloti Gran Canaria U- Lacerta dugesii dugesii Madeira l/* Lacerta perspicillata Mallorca Po darcis his panic a' li olepis' Castell6n,Spain Podarcis hispanica "t rpe l" P.h.r Vila Real, Pt. Ur B P.h.2 Montesinho, Pt. t/- A P.h.3 Montesinho, Pt. l/- A Podarc i s hispanic a "Moroccan" P.h.ml High Atlas, Morocco -11 T P.h.m2 High Atlas, Morocco Ur T P.h.m3 High Atlas, Morocco r/r T Podqrcis hispanica "We 2" P.h.vl Leiria, Pt. 1t- C P.h.v2 Portalegre,Pt. Ul D P.h.v3 Beja, Pt. r/l E P.h.v4 Marvao, Pt. l/- F P.h.v5 Agueda,Pt. U- G Podarcis bocagei bocagei P.b.b1 Montesinho,Pt. U- A P.b.b2 Montesinho, Pt. t/- A P.b.b3 Vila Pouca d Aguiar, Pt. l/- K P.b.b4 Serrado Ger6s,Pt. U- H P.b.b5 Vairdo, Pt. t/- J P.b.b6 Vairdo, Pt. U1 J P.b.b7 Vair6o, Pt. t/l J P.b.b8 Braga,Pt. t/- I P.b.b9 Viana do Castelo,Pt. t/- L P.b.b10 Viana do Castelo,Pt. t/- L P.b.b11 Viana do Castelo,Pt. Ur L P odarcis carbonelli carbonelli P.c.cl Serra da Estrela, Pt. U1 M P.c.c2 Torreira, Aveiro, Pt. l/l N P.c.c3 Monte Cldrigo, Pt. Ul a P.c.c4 Peniche,Pt. t/- P P o darc is c arb onelli b er lengensis P.c.b. Berlengaisle, off Peniche 1/- o PHYLOGENYOF IBERIAN PODARCISLIZARDS l3l

amplifiedregions of approximately350 bp and 550 bp respectively.Thermocycling consisted of 30 cyclesof 93"Cfor 30 secs,55"C for I min and 72oCfor I min, followedby a singlecycle at 72"Cfor 5 min. Successful PCR bands were purified using a QIAEX II kit (Quiagen)and sequencedon an Applied Biosystems Model373ADNA SequencingSystem, using a PRISM Ready Reaction DyeDeoxy Terminator Cycle Sequencingkit. Centrisepspin columns(Princeton SeparationsInc.) wereused for excessdye extraction.

PHYLOGENETICANALYSES Sequenceswere aligned using Clustal W (Thompson et al., 1994). There were no insertions or deletions. They were then imported into PAUP* (Swofford, 2001) for phylogenetic analyses. When estimating phylogenetic relationshipsamong sequences,one as- sumesa model of evolution regardlessof the optimality criteria employed. Determining which model to use given one's data is a statistical problem (Goldman, 1993). We used the approach outlined by Huelsenbeck and Crandall (1997) to test alternative models of evolu- tion, employing PAUP* and Modeltest (Posada & Crandall, 1998). A starting tree was obtained using neighbour-joining. With this tree, likelihood scores were calculated for various models of evolution and compared using a chi-square test, with degreesof free- dom equal to the difference in free parametersbetween the models being tested. The null hypothesestested in this way included:(l) nucleotidefrequencies are equal; (2) transition rates are equal to hansversion iates; (3) transition rates are equal,; (4) transversion rates are equal; (5) rate homogeneity within the data set; and (5) no significant proportion of invariable sites (Table 2). Once a model of evolution was chosen,it was used to estimate a tree using maximum likelihood. Ten repli- cate heuristic searches were made with random sequenceaddition. Confidence in resulting nodes was assessedusing the bootstrap technique (Felsenstein, 1985)with 1000replicates. Genes were analysedsepa- rately and in combination. FIG. l. Map showing the sampling localities used for the mtDNA analysis. The ranges of P. hispanica I (stippled), Je. BIOMETRICS b. bocagei (grey) and P. ft.) carbonelli (black) are shown. P. hispanica 2 is found in the region south o/P. hispanica l, but Eleven biometric variables were obtained from 12 the complete extent of its range is unknown. females and24 males from each of 20 populations (ex- act localities available on request), using 0.05 mm MATERIALS AND METHODS callipers (see procedure in P6rez-Mellado & Gos6, 1988):(l) snout-ventlength; (2) head length; (3) head LABoRAToRY PROCEDURES width; (4) inter-orbital width; (5) frontal width; (6) in- Localities of the lizards from which DNA was ex- ter-nasalwidth; (7) head depth; (8) orbital depth; (9) tracted and/or biometric characterswere scoredare frontal depth; (10) nasal depth; and (l l) hind limb givenin Tablel. Tissuesamples consisted of tail tips length (Fig. 2). Sexes were analysed separately. storedin 100%ethanol. Voucher specimens are kept at Squared Mahalanobis distance between centroids was the University of Evora.Total genomicDNA was ex- used since it takes into account the correlations among tracted from tail tissue using standard methods biometric variables and is independent of the relative (Sambrooket al.,1989). PolymeraseChain Reaction scales ofthe various variables (Legendre & Legendre (PCR) primersused in both the amplificationand the 1998),UPGMA clusteringwas applied to the distance sequencingwere cytochromeDl andb2 (Kocheret al., matrix to assess the lizard phenetic relationships 1989)and COle andCOlf (Palumbi,1998). These (Rohlfi 1993; Sokal & Rohlf, 1995). t32 D. J. HARRIS AND P. SA-SOUSA

TABLE 2. Tests of hypothesesrelating to the model of evolution appropriatefor phylogeny reconstruction (Huelsenbeck & Crandall, 1997). P-valueswere obtained with Modeltest (Posada& Crandall, 1998). For each hypothesis the data set with cy'tochromeoxidase (top), cytochrome6 (middle) and then with the combinedregions (below) is tested.Due to the performance of multiple tests,the significancelevel of rejection of the null hypothesiswas adjustedvia the Bonferroni correctionto o:0.01 .

Null hypothesis Models compared -lnLo -lnl-, df P

Equalnucleotide frequencres Ho:JC69, H,: F81 203s 2009 0.000 1558 I 509 0.000 2843 2822 0.000 Equalti andtv rates Ho:F81, H,:HKY85 2009 1907 0.000 I 509 t439 0.000 2822 2684 0.000 Equalti rates Ho: HKY85, H,: TrN 1907 189 0.000 1439 1425 0.000 2684 2684 0.581 Equaltv rates Ho:TrN, H,: TIM 1896 r 896 0.275 1425 1425 0322 Ho:HKY85,H,: K8luf 2684 2684 0.380 Equalrates Ho: TrN, H,: TrN+G I 896 1817 0.000 amongsites 1425 1357 0.000 Ho:HKY85,Hr: HKY85+G 2684 2591 0.000 Proportionof Ho:TrN+G, H,:TrN+G+i l8l7 t8t7 0.999 invariablesites 1357 l 355 0.057 Ho:HKY85+G, H,: HKY85+G+i 2591 2589 0.032

RESULTS viduals of P. hispanica or P. bocagei were included. For this data set the most appropriatemodel of evolu- Twenty-six individuals from l5 populations of P. tion was the HKY model (transition/transversionratio hispanica or P. bocagei were sequencedfor the cyto- of 4.99) with a discreteapproximation of a gamma dis- chrome oxidase gene. The closely related L. dugesii tribution of variablesites (Table 2). Using this model, a (Hanis et a1.,7998)was sequencedas an outgroup,and ten replicateheuristic search found a singlemost likely the sequencefrom the more distantly related Gallotia tree,with a likelihood of -2586 (Fig. 3C). This ML- galloti was also included in the analyses.The most ap- based hypothesis of relationships was compared propriate model of evolution for this data set was the againsttwo altern ati ve s, with P. b oc a ge i monophyletic, Tamura Nei model (TrN) model with a discrete ap- andP. hispanlcamonophyletic respectively, and other gamma proximation of a distribution of variable sites relationshipsunrestrained. Using the same model of (basefrequencies A: 0.31,C: 0.16,G: 0.24,T:0.29, evolution, ten replicateheuristic searchesfound short- equal transversionratios and A/G 13.8, C/T 8.3, gamma shape parameter 0.18 - Table 2). Using this model, a ten replicate heuristic searchfound a single most likely tree, with a log likelihood of -1811 (Fig. 3A). Twelve individuals from nine populations were also sequencedfor the cytochrome D gene. This was combinedwith four previouslypublished sequences for P. hispanica (Hanis & Amold, 1999), and two addi- tional outgroups, Lacerta dugesii and L. perspicillata. For this data set the most appropriate model of evolu- tion was again the TrN model with a discrete approximation of a gamma distribution of variable sites (basefrequencies A: 0.26, C: 0,29, G: 0.13, T: 0.32, equal transversionratios and A/G 3.5, ClT 9.7, garnma shapeparameter 0.26 - Table 2). Using this model, a ten replicate heuristic search found a single most likely tree, with a log likelihood of -1348 (Fig. 3B). Since both genesare mitochondrial and thereforeinherited as FIG 2. The I I biometric variablesused in the analysis. 24 a single locus, a combined analysis was also carried males and 12 females from each of 19 populations were out. L. dugesii was used as the outgroup, and 12 indi- measured,including all four mainland morphotypes. PHYLOGENYOF IBERIAN PODARCISLIZARDS t33

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FIG. 3. A, maximumlikelihood tee derivedfrom cytochromeoxidase sequences. The treewas rooted using Gallotia galloti and Lacertadugesii as outgroups.Numbers above nodes indicate bootstrap support (1000 replicates).B, maximumlikelihood tree derivedfrom cytochromeb sequences.The tree was rooted using previously published sequences of L. perspicillata andL. dugesii sequencesas outgroups.Numbers above nodes indicate bootstrap support (1000 replicates). C, maximumlikelihood tree derived from combinedcytochrome 6 andcytochrome oxidase sequencos. The treewas rooted using I. dugesiias an outgroup.Numbers abovenodes indicate bootstrap support. 134 D. J. HARRIS AND P. SA-SOUSA

TABLE 3. Maximum-LikelihoodTests (Shimodaira& analyses and bootstrap support is strong, especially in Hasegawa,1999) of alternativetree topologies for Podarcis the combined analysis - 99% for P. (b.) carbonelli, werethe maximumlikelihood tree lizards.Trees compared l00Yofor P. b. bocagei. The degreeofgenetic differen- basedon the combinedDNA sequencedata (Fig. 2C), and (b.) thosebased on altemativehypotheses where either Podarcis tiation between P. b. bocagei and P. carbonelli is bocageior Podarcishispanica are monophyletic. *P is the high: 9-9.6% betweenthe COI sequencesand 13.5- probabilityof obtaininga more extremet-value under the 15.5%between the cytochromeb sequences.The mean null hypothesisof no differencebetween trees. Both these cytochrome b genetic distance for congeneric reptile decreasedfit relativeto the hypothesesshow significantly speciesis 13.6Yo(Harris, in press),and lower levels of maximumlikelihood tree. COI divergence in the iguanian lizards of the genus Tropidurus have been used to recommend speciescan- Tree Log A Log *P didates (Frost e/ al., 1998). Further, within these two likelihood likelihood groupsgenetic distances are very low - 0-0.06%within Max. likelihood P. (b.) bocagei and 0.004-0,09% within P. (b ) free -2586 carbonelli. Maintenance of the presenttaxonomic sys- tem is further complicated by the rejection of P. Monophyletic bocagei monophyly using both morphological and P. bocagei -2s98 T2 0.029 molecular data. We recommend raising P. (b ) Monophyletic carbonelli to species status, following Sa Sotsa et al. P. hispanica -2599 t3 0.021 (2000) using protein electrophoreticdata. P. b. berlengensrsshows no genetic differentiation from mainland P. carbonel/l using the COI sequence Pft. 1SL data, and so should be referred to as P. c. berlengensis. PrrISR This has previously been suggestedbased on the low Pfi tsC P/,tSe genetic distancefound betweenthese groups, D:0.08 P/ttSt P.hzpr (56 Sousaet a1.,2000). P. c. berlengensrsdoes show P.h.2sl some distinct morphological features primarily associ- P.hzw P.h.zeu atedwith an increasedmean body size(Vicente, 1985). P.h,zEI A case has been shown for Gqllotia simonyi P.blGl similar P.b+up simonyi and G. s. machadoi, where an extinct subspe- P.b1p0 P.b.lpF cies from a small island showed no difference in P.c3CB mtDNA sequencesfrom the mainland form, despite P.c3PB P.c.5T0 morphological differences (Carranza et al., 1999). lt is, P.c.5CU however, markedly different from the example of P.c.5ttC r.c.5cR Podarcis atrata from the Columbretes Islands. where inter-island cytochrome 6 divergence is high (Castilla FIG. 4. UPGMA cluster analysisof the biometric variables. et al., 1998). Much has been made of the expected de- Included areP. hispanicatype | (P.h. l) and type 2 (P.h.2), crease in genetic diversity of organisms on small P. bocagei (P.b.) andP. (b.) carbonelli (P.c.). Separationof islands.and the associatedincreased risks ofextinction. the four forms is effective in the females, but less so in the Given the large numbers of insular subspecies of males. esttrees of -ln 2599and -ln 2598respectively. These Podarcis lizards (nearly 300; Bdhme, 1986), it is im- werecompared against the ML treewith the likelihood portantto determinewhich of thesephenomena is more variancetest of Shimodairaand Hasegawa (1999) using common. 1000RELL bootstraps.Both were significantlyless Subspecies-leveltaxonomy within P. hispanica has likely (Table3). been controversial.Some authors accept one subspe- The UPGMA treesderived from the morphological cies in Iberia, P. h. hispanica and one in North Africa, datashow that eachmajor cluster corresponds to oneof P. h.vaucheri(e.g. Pdrez-Mellado, 1986, 1998). Others the four formsof wall lizard(Fig. 4). Clearseparation is argue that P. h. vaucheri is also found in the southern foundin females,while somepopulations of malesbe- Iberian Peninsula, and more separateforms within the longing to one group cluster with other forms. P. Iberian Peninsulaare to be recognized,though as yet bocageibocagei clusters with P. hispanicatype2; both with undeterminedtaxonomic status(e.g. Guillaume, groupedin the next stepwith P. hispanical; and fi- 1987,1997).Electrophoretic analyses have given con- - nally,P. bocageicarbonelli is themost dissimilar. flicting results Busack (1986) found a low genetic distance(D:0.07) betweenAndalusian and Moroccan DISCUSSION populationsof P. hispanica,while Capula (1997) sug- Thereciprocal monophyly of the designatedsubspe- geststhey are well differentiated (D:0.237), and could ciesP. b. bocageiand P. (b.) carbonel//,is strongly representsibling species. supportedby the mtDNA analysis.Specimens from Our analysessupport the conclusions of S6-Sousa acrossthe rangesform monophyleticgroups in all (2000a) that P. hispanica in Portugal is composed of PHYLOGENY OF IBERIAN PODARCISLIZARDS 135 two genetically distinct clades. However the southern Frost,D. R., Crafts,H. M., Fitzgerald,L. A. & Titus, T. A. form (either P. hispanica Wpe 2 or P. hispanica (1998). Geographic variation, species recognition, vaucheri) is also distinct from the population sampled and molecular evolution of cytochrome oxidase I in from Morocco. Whether there are multiple cryptic Afri- the Tropidurus spinulosus complex (Iguania: can species, as has been suggested - based on Tropiduridae).Copeia 1998, 839-85I immunological data (Joger & Bischoff, 1989) cannot Gal6n, P. (1986). Morfologia y distribuci6n del gdnero be assessedfrom the present data. It is clear, however, Podarcis Wagler, 1830 (Sauria, Lacertidae) en el that P. hispanica taxonomy needsto be reassessed- noroestede la Peninsulalb6rica. Rev. Esp. Hero. 1.87- our data indicate that P. hispanica is made up of multi- t42. ple genetically distinct clades that do not form a Gal6n, P. (1997). Podarcis bocagei (Seoane, 1884). monophyletic group relative to P. bocagei and P. Lagartija de Bocage, lagartixa-de-Bocage,lagartixa carbonelli. Additional data from nuclear loci will be galega. In Distribuci6n y biogeograJia de los anfibios neededto confirm this finding basedon mtDNA, and to y en Espafia y Portugal. pp. 237-239 J. M determine whether introgression occurs. Only exten- Pleguezuelos (Ed.). Granada, Monogrdfica Tierras sive sampling across the remainder of the range, del Sur, Univ. Granada and Asoc Herpetol. Espaftola. especially central and eastern Spain and North Africa Gal6n, P. (2000). Females that imitate males: dorsal will allow a more appropriate assessmentof the status coloration varies with reproductive stage in female ofthese clades. Podarcis bocagei(Lacertidae). Copeia2000, 819-825. Goldman,N. (1993). Simple diagnosticstatistical tests of ACKNOWLEDGEMENTS modelsof DNA substitution.J. Mol. Evol.37,650-651 Gonziiez de la Vega,J. P. (1989). Anfibios y reptilesde la D. J. Harris and P. S6-Sousawere supported by provincia de Huelva. Huelva: ImprentaJimdnez, S.L. grants from Fundagdo para a Ci€ncia e Tecnologia Guillaume, C.-P. (1987). Les petits lacertiddsdu Bassin (FCT) Praxis XXI -BD/5268195, PRAXIS/PIBIAI Mediterranden Occidental (genera P odarcis et 1| l7 4/ 1998 and POCTI/3 4 547 /BSE 12000.We are very Archaeolacerta essentiellement). Doctoral thesis. gratefulto P. Celio Alves and J. Teixeira for assistance Montpellier, Univ. Sciences et Techniques du in collecting samples. Thanks to J. W. Arntzen, N. Languedoc. Ferrand and two reviewers for helpful comments on an Guillaume, C.-P. (1997). Podarcis hispanica earlier draft of this manuscript. (Steindachner, 1870). In Atlas of amphibians and reptilesin Europe,pp.278-279. J. P. Gascet al. (Eds.) REFERENCES Paris, Mus. Nat. Hist. Nat. Paris and Soe. Europ. Bcihme,W. (1986).Handbuch der Reptilien und Amfibien Herpetol.(SEH). Europas,3, Weisbaden:Akademische Verlagsgesells- Harris, D. J. (in press). 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