Intraspecific variability of the Carpetane Lizard (Iberolacerta cyreni [Müller & Hellmich, 1937]) (Squamata: Lacertidae), with special reference to the unstudied peripheral populations from the Sierras de Avila (Paramera, Serrota and Villafranca)

2010·8.1 MB·English

by  Oscar J Arribas

Bonn zoological Bulletin Volume 57 Issue 2 pp. 197-210 Bonn, November 2010
Intraspecific variability of the Carpetane Lizard
&
{Iberolacerta cyreni [Miiller Hellmich, 1937]) (Squamata: Lacertidae),
with special reference to the unstudied peripheral populations
from the Sierras de Avila (Paramera, Serrota and Villafranca)
Oscar J. Arribas
Avda. Fco. Cambo 23; E-08003 Barcelona, Spain; E-Mail: [email protected]
Abstract. Canonical Discriminant (CDA), ANOVA and ANOSIM analyses were calculated for all recently known dis-
tribution areas ofIberolacerta cyreni including several small and unstudied peripheral populations. The only differenti-
atedsampleisGuadarrama(thenominatesubspecies),withverylimitedoverlap intheCDA(correctclassification>70%)
and different from nearly all the other samples in ANOSIM. Guadarrama is a recently differentiated but well diagnos-
able (morpho)subspecies (with lowervalues ofdorsalia, ventralia and greatervalues ofcircumanalia). Despite the mtD-
NA differences ofthe Bejar specimens, their morphology is largely equivalent to that ofI. cyreni castiliana (Gredos),
but clearly differ in their female body elongation (near 1 cm) with shorter limbs, a possible strategy to increase clutch
size. Populations from the Sierras de Avila (Villafranca, Serrota and Paramera) are very similar among them. Villafran-
ca (in males) together with Bejar (in females) are the most connected samples in MST, and the root ofthe species dif-
ferentiation from a morphological pointofview, once discarded geographical andclimatic influence on morphology.All
populations exceptGuadarrama shall beconsideredas/. c. castilianabytheirmorphological identity withGredos.These
morphological similarities probably are the reflect ofextensive gene flow among them, responsible ofmaintaining their
morphology largely equivalent.
Key words. Lacertidae, Iberolacerta cyreni, Intraspecific variability, subspecies, Geographical variation, Iberian Penin-
sula.
Resumen. SehacalculadounAnalisis Discriminante Canonico,ANOVAyANOSIM contoda el areade distribucion de
Iberolacerta cyreni, incluyendo varias poblaciones perifericas no estudiadas hasta la fecha. Launica muestra diferencia-
da es Guadarrama (la subespecie nominal), con muy poco solapamiento en el CDA (clasificacion correcta > 70%) que
difiere depracticamente todas las demas muestras en elANOSIM. Guadarrama es una poblacion recientemente diferen-
ciada, pero bien diagnosticable como (morfo)subespecie (valores bajos de dorsalia y ventralia, y altos de circumanalia).
Apesarde las diferencias mitocondriales de Bejar, su morfologia es ampliamente asimilable a/. c. castiliana (Gredos),
siendo destacable el relativo elongamiento corporal de lashembras (casi 1 cm) con miembros proporcionalmente cortos,
unaposible estrategiaparaincrementarel tamano depuesta. Las poblaciones de las Sierras de Avila (Villafranca, Serro-
tayParamera) sonmuy similares entre si.VillafrancajuntoconBejar(enmachosyhembrasrespectivamente)estanmor-
fologicamente en laraizde ladiferenciacion de laespecie (MST), unavezdescartadacualquier influenciaclimaticao de
distancia geografica. Excepto Guadarrama, todas deben considerarse como/. c. castiliana por su identidad morfologica
con Gredos, lo que refleja la probable presencia de un flujo genetico extensivo y reciente entre ellas.
INTRODUCTION
The Spanish Sistema Central consists ofa series ofSier- (over 240 km in length), which is inhabited in several
ras, more orless aligned in aENE-WSW direction, which points by populations ofthe Carpetane Lizard (Iberolac-
&
separate the Duero (to the North) and Tajo (to the South) erta cyreni [Miiller Hellmich, 1937]).
river drainages, or what is the same, the Old and New
Castile plateauxes. It runs from the Portuguese Serra da The Carpetane Lizard is widespread through the main
Estrela (inhabited by a relict population ofIberolacerta parts ofthe Sistema Central and is mainly known from
monticola [Boulenger, 1905]), across the Spanish Sierra Sierras de Bejar, Gredos and Guadarrama (Fig. 1). It was
de Gata (apparentlytoo low anddry forIberolacerta), the raised to species level (Arribas 1996) based on allozymes
&
Sierra deFranciawithIberolacerta martinezricai(Arribas, (Mayer Arribas 1996), karyology (heterochromatiniza-
1996) andthe mainpart ofthe Spanish portion ofits range tion of sex-chromosome and localization of the NORs;
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
198 Oscar J. Arribas
Odierna et al. 1996) and adult and hatchlings pattern and zor, 2.592 m), whereas the otheraxis is constitutedby the
coloration. Two subspecies were defined, the nominal /. Sierras de Villafranca (Moros, 2.065 m), La Serrota (Ser-
c. cyreni from Guadarrama (type locality: Puerto de rota, 2294 m), La Paramera (Zapatero, 2.160 m) and
Navacerrada), and /. cyreni castiliana from Gredos (type slightly separated by lowerareas, Guadarrama (Penalara,
locality: Circo de Gredos, Avila) to which frequently are 2.430 m). The two axes greatly overlap longitudinally
assimilated Bejar specimens. This latter subspecies differs leaving the Villafranca, Serrota and Paramerasjust to the
from the nominate one by a reduced dark pattern, more Northofthe Sierra de Gredos, butattheirorographic shad-
dorsalia, ventralia, slightly larger diameter ofthe masse- ow for rains, and climatically more continentalised. This
teric andhindlimb length, and lowercircumanalia (Arribas explains the botanical similarities between the Paramera-
1996). Serrota-Villafranca axis and the Sierra de Guadarrama
(Luceno and Vargas 1991).
The degree ofgenetic differentiation between /. c. cyreni
(Guadarrama) and /. c. castiliana (Gredos) was analyzed In Guadarrama (where Podarcis muralis also exists), /.
by Mayer & Arribas (2003), who found a mtDNA se- cyreni occurs only at the highest areas, from 1.760 m
quence divergence of 0.6 % in the 12s rRNA (12s) and (Puerto de Cotos, Puerto de Navacerrada) up to the peaks
16s rRNA(16s) mitochondrial genes, which corresponds (2.340 m in Penalara). In Gredos itlives almost from 1.700
to approximately 0.6 MY BR Carranza et al. (2004) sug- to 2.500 m. It was seen in 17. VII. 1986 (own data) in
m
gested that both subspecies diverged approximately Puerto del Pico (at 1.352 close to one ofthe fountains
MY
0.8±0.2 BP, an estimation mainly based on the Cy- ofthe pass) but recent research in this area has been to-
tochrome b (Cyt b) mitochondrial coding gene (the 12s tally unfruitful. It is possible that these lower stations
andthenucleargene C-moswereuninformative atthis lev- favouredby accelerated coldwinds in the mountainpass-
el), a divergence time almost identical to the one calcu- es (Venturi effect) haddisappeared by climatic orbest, by
lated by Crochet etal. (2004) using also the Cytb gene [1.6 habitat degradation due to human over-frequentation dur-
%
genetic divergence, which roughly corresponds to 0.6 ing the last 20 years. In Bejar it has been found between
to 1, with a mean of0.8 MY BP). These two values were 1.837 m (own data) and 2.443 m (see Lizana et al. 1988,
very similarto the above-mentioned ciphers. The inferred 1992, 1993; and Martin 2005 for general data; own data
MY
divergence time increased up to 1.2 (Cytb) or 1.6 BP corrections for the confirmed lower limits).
(Cyt b+12s) when different terminal taxa evolutionary
models and phylogenetic methods were used (Arribas et Apart from the better known Sierras, the presence ofthe
al. 2006; Arnold et al. 2007; respectively). Carpetane Lizard in the small parallel mountain ranges
called "Sierras de Avila" or "Parameras" (composed by
On the other hand, specimens from Sierra de Bejar three Sierras: Villafranca, La Serrota, and La Paramera)
branched at the base ofthe /. cyreni clade in some mtD- was first discovered by the mountaineering group "Valle
NA analyses (Carranza et al. 2004; Arribas & Carranza de Ambles" (Lizana et al. 1993), but no specimens have
2004). It was suggested that the split ofthis populations been studied so far. All aspects ofmorphology, status and
MY
occurred approximately 1.7±0.3 BP (Carranza et al. relationships ofthese small and isolated populations from
2004). However, in analyses using the same mtDNA re- the Sierras deAvila are totally unknown. In these Sierras
gions but differenttaxa andotherevolutionarymodels and the species is extremely localized, especially in La Ser-
phylogenetic methods than above (Arribas et al. 2006; rota and Paramera. In Sierra de Villafranca, the area with
Arnold et al. 2007), the specimens from Bejar formed a a relatively more extended suitable area, I have found it
trichotomywith/. c. castiliana from Sierra de Gredos and from 1.850 m probably up to the highest areas (Pico Mo-
/. c. cyreni from Navacerrada. ros, 2.065 m). In La Serrota it is extremely rare and lo-
m
calized, also cornered in the highest parts, from 2.284
m
Apart from the uninformative C-mos nuclear gene frag- (perhaps 1.935 where excrements, possibly of this
ment analyzed by Carranza et al. (2004) there is only one species, were seen; pers. obs.) to the very summit (Pico
m
other information about differences at the nuclear level, Serrota, 2.294 m); and in La Paramera from 1.700 in
the analysis of allozyme data by Almeida et al. (2002), the northern slopes to the summit (Pico Zapatero, 2.160
which showed a Nei's distance of 0.002 between speci- m) (own data).
mens from Gredos and Guadarrama.
After a three-year prospection ofthese parallel ranges, I
From West to East, the distribution of/. cyreni can be di- gathered data from these localized and barely known pop-
vided into two axes connected by low mountain valleys ulations in order to check the relationships ofall the Car-
(see appendix II), but not clearly interrupted by clear cut petane Lizards throughout its range. My aim is: a) to re-
barriers as river valleys. One axis runs across Sierra de assess differences between/. c. cyreni and/. c. castiliana
Bejar (summit in La Ceja, 2.425 m) and Gredos (Alman- inthe lightofthe existence ofothersmall and isolatedpop-
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
Intraspecific variability in Iberolacerta cyreni 199
illations; b) to ascertain the taxonomic status ofthe Bejar Cabezo, Gredos Oriental Massif, Avila province, Spain).
populations and to check ifthese represent a further sub- 6 males and 7 females. GREDOS: Circo de Gredos (Gre-
species; c) to study both the relationships among the sam- dos Central Massif, Avila province, Spain). 23 males and
ples from the Sierras de Avila (=Parameras) massifs, as 46 females cyreni castiliana]. BEJAR: Sierra de Be-
[/.
well as their similitude and differences with their neigh- jar (Gredos Occidental Massif, also known as Sierra de
bouring and well known populations from Gredos, orthe Candelario, Salamanca province, Spain). 11 males and28
more distantpopulations from Guadarramaand Bejar; and females. VILLAFRANCA: Sierra de Villafranca (Avila
d) as the type series of/. cyreni was destroyed during the province, Spain). 20 males and 14 females. SERROTA:
Second World War (SWW), to choose a Neotype for the La Serrota (Avilaprovince, Spain). 3 males and2 females.
species (see appendix I) in order to fix unequivocally the PARAMERA: Sierra de La Paramera (Avila province,
type locality (although apparently all lost, there were al- Spain). 4 males and 1 female.
so specimens from Gredos in the original type series).
Thesepopulations are discontinuous amongthem (Appen-
dix II) andconstitute discrete geographical OTUs (Fig. 1).
MATERIAL & METHODS
Due to lower sample, it was necessary to cluster the Ser-
Morphology rota and Paramera specimens in the male discriminant
analysis, andthese two plus Villafranca (all ofthem "Sier-
A
total of 106/92 male specimens, and 136/135 female ras deAvila") in the female one. However, reciprocal dis-
specimens of /. cyreni with a complete measurements tances between each one ofthese poorlyrepresented sam-
dataset and snout-vent length greater than 45 mm, were ples to the best represented ones were carefully checked
included intheunivariate (ANOVA) andmultivariate (dis- and commented in the results section. As the three popu-
criminant) analyses, respectively. Given that these popu- lations from Sierras deAvila seemed to be largely equiv-
lations present sexual dimorphism (Arribas 1996, 1999a; alent in the multivariate analyses, aposteriori, all ofthem
Arribas et al. 2006), analyses were carried out for males were treated as a single OTU (SaAVILA) in ANOVA.
and females separately.All material is from OscarArribas
(OA) database.
Characters studied
OTUs names, localities and specimens included in the
morphological multivariate analysis were as follows (Fig. Biometric characters. Snout-vent length (SVL); Forelimb
1): length (FLL); Hindlimb length (HLL); Pileus length (PL);
Pileus width (PW); Parietal length (PaL); Masseteric scale
GUADARRAMA: Sierra de Guadarrama (Madrid and diameter (DM); Tympanic scale diameter (DT); Anal
Segovia provinces, Spain). 25 males and 36 females [/. width (AW) and Anal length (AL). All linear measure-
cyreni cyreni]. MIJARES: Puerto de Mijares (Sierra del ments were made with a digital calliperto the nearest0.01
mm. These measurements weretransformedto the follow-
ing more informative and not dimensional-depending ra-
tios: FLL/SVL (relative forelimb length; "FLL index");
HLL/SVL (relative hindlinb length, "HLL index");
PL/PW (pileus shape, "Pileus index"); DM/PaL (relative
masseteric plate size, "Masseteric index"); DT/PaL (rel-
ative tympanic size, "Tympanic index"); AL/AW (anal
plate surface, "Anal form index") and AS/SVL
(V(AL*AW)*100/SVL, relative anal plate size with re-
spectto the total length, "Anal size index") (Arribas 1996,
2001 ). The results ofthe linear measurements and index-
es yielded largely similar results. All ratios were given
multiplied by 100 to avoid excessive decimal scores.
Scalation characters. Supraciliar Granula (GrS) for the
Fig. 1. Schematic representation ofthe distribution ofIbero- rightand left sides; Gularia(GUL); Collaria(COLL); Dor-
lacerta cyreni in the Spanish Sistema Central. The different lo-
calities (OTUs) citedinthetextarerepresented. 1: Bejar; 2: Gre- salia (DORS); Ventralia (VENT); Femoraliarigth (FEMr)
dos; 3: Villafranca; 4: Serrota; 5: Paramera; 6: Mijares; 7: Gua- and left (FEM1); 4th. digit Lamellae (LAM); and Circum-
darrama. analia (CIRCA).
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
200 Oscar J. Arribas
StatisticalProcedures populations permit to test significance ofthe differences
amongthe concerned groups and to detectwhich ones are
Statistical analyses used in the morphological study in- really different from the others.
ANOVA
cluded both Univariate ( for SVL, scalation char-
acters and indexes, withpost-hoc Tukey-Kramer tests at Mahalanobis' (squared) distance matriceswere compared
P < 0.05 and P < 0.01 to detect differences among sam- by means ofMantel Test (with 1000 permutations) with
ples) as well as Multivariate techniques (Canonical Dis- matrices composed by Euclidean (squared) distances for
criminantAnalysis, CDA). Inthis lateranalysis, eachpop- the climatic characteristics of localities: a) Precipitation
ulation is represented by a centroid (a hypothetical mid- (mm) during the incubation months (July andAugust, as
dle individual). Minimum-length spanningtree (MST) was scalation is invariant during lizard's life); b) Annual pre-
computed from the Mahalanobis' distance matrix to de- cipitation (mm), c) Temperature (°C) (July and August);
tect the nearest neighbours based on their position in the d) Annual average temperature e) Sun radiation (n 10
multidimensional space. MST representation also avoids kJ/(m2*day*micrometerJ(July andAugust), and f) Annu-
UPGMA
distortion of trees by the reciprocal pairwise dis- al Sun radiation. Data were extracted from Ninyerola et
tances recalculation in every step during their construc- al. (2005). Also, these Mahalanobis' distances were com-
UPGMA
tion. frequently clusters samples reflecting sam- pared with (d) the aerial (straight) geographical distances
ple sizes than their true relationships. Distances ofsmall among the sampling localities.
samples or isolated specimens appear greatly exaggerat-
ed with respect to the well represented ones. As a result Multivariate (Discriminant andANOSIM) analyses were
ofthat, the small-sized samples appear ever as the most performed with Community Analysis Package 4.0 (Hen-
UPGMA & MST
external or differentiated in derived trees derson Seaby 2007). trees and Mantel tests were
(Kunkel et al. 1980; Cherry et al. 1982; Arribas 1997). calculatedwithNTSYS 2.1© (Rohlf2000). Univariate sta-
Moreover, theUPGMAtreesbased inveryunevenly sized tistics were processed withNCSS 2001 0package (Hintze
samples also gave very poor Cophenetic Correlation In- 2001).
dexes between the tree-derived ultrametric distances ma-
trix and the original Mahalanobis distance matrix and
therefore we have not used them (Arribas et al. 2006). RESULTS
To test the significance ofthe differences among pre-es- Males
tablished groups for the Discriminant Analysis (based in
a geographical origin), we carried out anAnalysis ofSim- Canonical Discriminant Analysis. The CDA conducted
ilarity (ANOSIM) (Clark 1988, 1993) that tests ifthe as- with 106 male specimens shows three significantaxes that
%
signed groups are meaningful, this is, more similar with- explain an 85 ofthetotal intersample variation. The two
in groups than with samples from different groups. The first axes togetherexplain the main part (70.6 %) and dis-
method uses the Bray-Curtis measure ofsimilarity to con- criminate fairly well the samples, especially the first one,
struct clusters ofspecimens. The null hypothesis is there- the unique with an eigenvalue >1. The first discriminant
%
fore that there are no differences between the members of axis has an Eigenvalue of 1.54 (51.2 of variance ex-
the compared groups (they are randomlyblended). R-sta- plained; Chi-Sq. with 85 df=200.71, P< 0.0001) and dis-
tistic scales from +1 to -1. Values closer +1 correspond tributes the samples with fairly overlap among them, ex-
to aperfect case in which all groups were completely dif- cept Guadarrama, that has only a small coincidence with
ferent (all specimens ofthe same group are more similar the otherones (Fig. 2A). Guadarrama appears in the neg-
among them than to any specimens ofthe other groups). ative part ofthe axis, characterised by the lower values
R = 0 occurs ifthe high and low similarities are perfect- for DORS (0.441553) andVENT (0.560765) and greater
ly mixed and bearno relationship to the group, a common values of CIRCA (-0.572683). Second and third axes
situation ifsome ofthe groups are largely equivalent. A (eigenvalues < 1) present a considerable overlap among
value of-1 indicates that the most similar samples are all the samples and do not discriminate populations.
outside ofthe groups (all groups largely equivalent and
randomly formed). To check forsignificance, pseudorepli- This discriminant analysis appliedto the samples reached
cation tests (1000 randomizations) were run to test ifthe a 72.6% of correct classification among the specimens.
given results canoccurby chance. Ifthe value ofR is sig- The Guadarrama sample (/. cyreni cyreni) reaches a 71.9
%
nificant, there is evidence that the samples within groups ofcorrect classification in respect to all the other sam-
are more similar than would be expected by random ples (/. c. castiliana).
chance. Even more important, pairwise tests among
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
Intraspecific variability in Iberolacerta cyreni 201
DiscriminantPlot-lb cyreni MALES
~3 3 A 0 1 2 i I
Function1
DiscriminantPlot-lb cyreniFEMALES
Fig. 2. Canonical DiscriminantAnalysis (CDA)plots fora) males (above) andb) females (below). Specimens, samplecentroids
andgroupperimeters arerepresented. Guadarrama(invertedtriangles), Gredos (triangles), Bejar(irregularcircles), Mijares (cross),
Villafranca (asterisk) and Serrota-Paramera (sail). In females, the three last samples are grouped as Sierras deAvila (sail). Sample
centroids are represented by a square. See text for axis characteristics and results.
Minimum-length spanning tree (notrepresented) connect- Analysis of Similarity (ANOSIM) (Table 3) shows that
ing the centroid (hypothetical middle specimens) ofeach there is a considerable overlap among samples (R-statis-
sample is fairly congruent with their geographical posi- tic = 0.122088, P < 0.005; 1000 randomizations) as our
tion, connecting in general neighbouring samples. The value (that can range from 1 to -1), although positive, is
most "central" (most connected) population is Villafran- very small. Very significant differences among the (geo-
cathatconnects with Gredos (at Mahalanobis Distance of graphically) assigned groups, appearonly among Guadar-
3.1870), Bejar(3.2199), Paramera (4.8042) andfinally, to rama and Bejar, Gredos andVillafranca (P < 0.01) but do
the most isolated one, Guadarrama (6.8412). Two popu- not reach significance with Mijares and Serrota + Para-
lations show overestimated distances due to their small mera (both with small samples). The otherpopulations are
sample sizes: Mijares (EastGredos) thatconnects with Be- not differentiated among them (P > 0.01).
jar (7.0951), and Serrota with their neighbouring Param-
era (9.1822).
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
202 Oscar J. Arribas
TheAnalysis ofVariance (ANOVA) (Appendix III, Table
1) indicate that Guadarrama differs from all or nearly all
the other populations in VENT and CIRCA (with the
smaller and greater values for these parameters, respec-
tively, in the former population), but also appeared dif-
ferences between Guadarrama and Bejar in Dors (small-
er in the former), and with Gredos in PV (greater in the
former). An interesting and significant difference appears
in DORS among Gredos and Bejar samples (clearly
greater in the later).
There is no significant correlation among Mahalanobis'
distances and any ofthe geographic and climatic param-
eters analyzed (all Mantel Tests P > 0.05).
Females
Canonical Discriminant Analysis: The CDA conducted
with 136 female specimens shows three significant axes
%
that explain a 93.8 ofthe total intersample variation.
Thetwo first axes togetherexplain a largepartofthevari-
ance (85.7 %), and especially along the first one, that ac-
%
counts itself for 62.7 ofthe total variation and is the
unique with an eigenvalue >1 (1.77), discriminating
Guadarrama specimens fromthe otherneighbour samples
only with a small overlap (Fig. 2B). The other samples
show a considerable overlap among them. Guadarrama
discriminates towards the negative part ofthe axis, char-
acterised by the lowervalues ofDORS (0.62) andVENT
(0.66) and greater ones of CIRCA (-0.38). Second and
third axes (eigenvalues < 1) present a considerable over-
lap among the samples and do not discriminate popula-
tions.
The discriminant analysis applied to the samples reached
a 74.26% ofcorrect classification among the specimens.
%
Guadarrama sample (I. c. cyreni) reaches an 87.2 ofcor-
rect classification with respect all the other samples (I. c.
castiliana).
Minimum-length spanningtree (notrepresented) connect-
ing centroids is very similartothemale one. The mostcon-
nected sample is Bejar, which clusters withVillafranca (at
2.9), Gredos (3.5) and Mijares (7.38, but here exaggerat-
edbythe scarce sample ofthe later). Guadarramaconnects
with the scarcely represented (and geographically inter-
mediate) Mijares (East Gredos) (at 6.09), and all the Sier-
ras deAvila samples clustertogether(Villafrancawith Ser-
rota + Paramera at 8.66).
Fig. 3. Iberolacertacyrenicastiliana. a) LaCovatilla Skyre-
sort (Sierra de Bejar), July 2007, Male ; b) El Travieso (Sa de Analysis of Similarity (ANOSIM) (Appendix III, Table
Blyej2a0r)0,4,JuFleym2a0l0e4,(aFteympiaclael;pca)ttEelrnC,alwviittherdoif(fSuimeirrnaatdieonBeajanrd),coJau-- 3) shows that there is a considerable overlap among sam-
lescence in a unique vertebral line); d) Puerto de Mijares (Gre- ples (R-statistic = 0.162588, P < 0.001; 1000 randomiza-
dos Oriental Massif), July 2006, Female. tions), but the results are slightly best than for the male
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
Intraspecific variability in Iberolacerta cyreni 203
analysis. Very significant differences (P < 0.01) appear
among Guadarrama and all the other samples exceptwith
Mijares. The other populations are not differentiated
among them (P > 0.01).
TheAnalysis ofVariance (ANOVA) (Appendix III, Table
2), as in the male analysis, it shows that Guadarrama is
the most different one, especially in DORS, VENT and
CIRCA(the firsttwo characters smaller, andthe third one
greater in the former population). Guadarrama also dif-
fers from Gredos by its lower GUL, from Bejar by its
greaterFLL, HLL, a lower SVL; and from Sierras deAvi-
la by its greater relative anal scale surface.
Also, significant differences appear in SVL between Be-
jar (clearly the great sized female population) and Gre-
dos, and among this latter (with relative greater FLL and
HLL) with Bejar and Villafranca.
As in male analysis, there is no significant correlation
among Mahalanobis'distances andthe geographic andcli-
matic parameters analyzed (all Mantel Tests P > 0.05).
DISCUSSION
From the Discriminant and ANOVA analyses it appears
that the only differentiated sample is Guadarrama. It ap-
pears with very limited overlap with the other samples in
CDA A
the graphs (Figs 2 and B). Diagnostic characters
forthis population (nominate subspecies: /. c. cyreni) are
the lowervalues ofDORS (difference more marked infe-
males), lowervalues ofVENTand greaterCIRCA. More-
over, ANOSIM analyses show that Guadarrama is the
unique OTU that is significantly different from nearly all
the othersamples, except fromthe closepopulation ofMi-
jares (in both sexes) and Serrota + Paramera (butthese ex-
ceptions occuronly in the males andprobably due to their
scarce sample size).
Mijares sample (very small) seems in some aspects ap-
proaching to Guadarrama (specially in DORS and VENT
values) but globally are clearly closer to /. c. castiliana
(specially to Bejar in male and female MST).
Populations West from Guadarrama show a great overlap
in CDAand lack differences inANOSIM, being morpho-
logically fairly equivalent and all ofthem assimilable to
/. c. castiliana. There are only a few scattered very sig-
nificant differences among them (P < 0.01) in ANOVA,
Fig. 4. Iberolacerta cyreni castiliana. a) Pico Zapatero (Si- as for instance among Gredos and Bejar (this latter has
erra de la Paramera), July 2005, Male; b) Puerto de PenaNegra greater DORS and a strikingly greater SVL and propor-
(SierradeVillafranca),July2006, Male; c)Pico Serrota(La Ser- tionallyshorterlimbs that Gredos, butonly in female spec-
rotaMassif),July2005,Female; d) Pico Serrota(LaSerrotaMas- imens). The reason of the longer SVL in Bejar females
sif), July 2006, Male. (from 8 mmto 1 cm greaterthan inthe otherpopulations)
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
204 Oscar J. Arribas
which leaves proportionately shorter limbs, can be acon- be due to isolation-by-distance processes but by histori-
sequence of body elongation that in lacertids appears cal vicariant events (cf. Irwin 2002) as there is no rela-
linked to a greaterclutch size (Brana 1996). This is an in- tionship between morphological differentiation and geo-
teresting question for future study: ifBejar specimens ef- graphic distances. Also there is no relationship amongthe
fectively have greater clutch size that other/, cyrenipop- more obvious climatic parameters (precipitation, temper-
ulations. ature and sun radiation) and these differences.
MST
Both the results (in which Villafranca and Bejar are Concerning the position ofBejar populations, only spec-
the most connected samples) as well as the presence of imens fromthe west-facing slopes ofthe massifhavebeen
related species further West (/. martinezricai and /. mon- studied, and therefore it is possible that in other parts of
ticola), suggest an origin ofthe species towards the west- the massifother haplotypes (the common one with Gre-
ern extreme oftheir current distribution area. From these dos) could be present. The species was cited from "Puer-
westernmost parts, where it also occurs the higher haplo- to de Tomavacas, SA" (for Salamanca, sic.!; a mistake as
type diversity (see below), /. cyreni spread towards the this locality is in Avila) (Lizana et al. 1992). This is the
East. Despite that the Sierras de Avila (Villafranca, Ser- natural pass between Bejar and Gredos, but I have been
rota and Paramera) are slightly more aligned with the unable to find it there. This place is a sub-Mediterranean
Guadarrama axis than with Gredos one, we cannotbe sure environment with Pyrenean Oak open forest inhabiting
from which ofthese two mountain ranges the formerwas populations ofTimon lepidus, Psammodromus algirus and
MST
colonized, as results in males and females are con- even Buthus occitanus, all thermophylous species typical
tradictory. According to the male analysis Guadarrama is from dry conditions. The lower height ofPuerto de Tor-
more related to Villafranca, whereas in the female analy- navacas (1010m) makes me to suspect that/. cyreniis not
ses, it is Mijares (Eastern Gredos) the most related one. there and the record is possibly a mistake. One possibil-
ity is that it was from the higher neighboring mountains.
The results of the mitochondrial analyses of these sam-
ples (Cyt B and 12s) (unpublished, Carranza, pers. com.) An account about the pattern and coloration of/. cyreni
indicates that the interruption ofgene flow is fairlyrecent, is in Arribas (1996) and it is especially detailed for the
as a common haplotype appears in all populations except main Sistema Central massifs in Perez-Mellado et al.
Villafranca and Bejar. All Gredos, Guadarrama and La (1993). Both colour as well as the dark pattern, seem to
Serrota specimens are identical for these two mitochon- be selected in accordance to the substrate characteristics.
drial fragments. Independent changes in one nucleotide Overall, the background colour is brown injuveniles and
withrespectto the common haplotype appearinVillafran- subadult specimens, changing in different percentages to
ca (the unique change is different in two specimens), Para- green in adult specimens (more frequently in males and
mera, Mijares and Bejar specimens, and two changes ac- becoming more vividly linkedto reproductiveprocesses).
cumulate in one Bejar and one Mijares specimen (others In populations inhabitingrocks (plenty ofRhizocarpon gr.
have only one). geographicum lichens) as in Gredos and the upper parts
ofGuadarrama (Pehalara) green adults are more frequent
The current morphological differences of Guadarrama (both males and big females). When living in rocky talus
specimens seem to be relatively recent, and are possibly with sands and bare ground (as in Navacerrada area)
the result of bottleneck effects during the colonization brownish adult specimens are more frequent (Arribas
process, or alternatively ofstrong selective pressures (or 1999b: Figs 9 and 10).
a combination of both causes). Conversely, the absence
of marked differences among the other populations Concerning the reticulate pattern, it also varies in a dif-
(more orless with a similarage) couldbe due to the main- ferent degree among the different populations depending
tenance ofa more continuous gene flow among them, re- on the substrates inhabited. Juveniles and subadults have
sponsible ofmaintaining theirmorphology largely equiv- temporal uniform orreticulate bands that coalesce during
alent (nuclear genes remain unstudied). The current larg- growth with dorsal spots (more frequently in males) giv-
er geographical gap in the distribution of/. cyreni occurs ing reticulated-like patterns. Usually, there is a relation-
precisely between Guadarrama and the remaining popu- ship between the size ofthe granite phanerocrystals (the
lations to the West. granite-rock spotting) and the habitus ofthe lizards liv-
ing on it. Lizards living on rocks that present large crys-
Despite the presumably short isolation time, as comment- tals (as for instance the Bejar ones) are more reticulated
ed above, a considerable selection pressure or a genetic than specimens living in places in which the rocks pres-
bottleneck in the expanding populations might have pro- ent smaller crystals (and thus finely spotted). Coloration
moted and fixed the morphological differences now seen accounts described in Perez Mellado et al. (1993) are fair-
in Guadarrama specimens. These factors do not seem to ly precise, especially for Gredos specimens. Concerning
Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK
[ntraspecific variability in Iberolacerta cyreni 205
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