Bonn Zoological Bulletin Volume 57 Issue 2 Pp

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 distribution areas of Iberolacerta cyreni including several small and unstudied peripheral populations. The only differenti-

ated sample is Guadarrama (the nominate subspecies), with very limited overlap in the CDA (correct classification > 70%) and different from nearly all the other samples in ANOSIM. Guadarrama is a recently differentiated but well diagnosable (morpho)subspecies (with lower values of dorsalia, ventralia and greater values of circumanalia). Despite the mtD-

NA differences of the Bejar specimens, their morphology is largely equivalent to that of I. 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 of the species differentiation from a morphological point of view, once discarded geographical and climatic influence on morphology. All

populations except Guadarrama shall be considered as /. c. castiliana by their morphological identity with Gredos. These morphological similarities probably are the reflect of extensive gene flow among them, responsible of maintaining their morphology largely equivalent.

Key words. Lacertidae, Iberolacerta cyreni, Intraspecific variability, subspecies, Geographical variation, Iberian Penin- sula.

Resumen. Se ha calculado un Analisis Discriminante Canonico, ANOVA y ANOSIM con toda el area de distribucion de Iberolacerta cyreni, incluyendo varias poblaciones perifericas no estudiadas hasta la fecha. La unica muestra diferencia- da es Guadarrama (la subespecie nominal), con muy poco solapamiento en el CDA (clasificacion correcta > 70%) que difiere de practicamente todas las demas muestras en el ANOSIM. Guadarrama es una poblacion recientemente diferen- ciada, pero bien diagnosticable como (morfo)subespecie (valores bajos de dorsalia y ventralia, y altos de circumanalia).

A pesar de las diferencias mitocondriales de Bejar, su morfologia es ampliamente asimilable a /. c. castiliana (Gredos),

siendo destacable el relativo elongamiento corporal de las hembras (casi 1 cm) con miembros proporcionalmente cortos, una posible estrategia para incrementar el tamano de puesta. Las poblaciones de las Sierras de Avila (Villafranca, Serro- ta y Paramera) son muy similares entre si. Villafranca junto con Bejar (en machos y hembras respectivamente) estan mor- fologicamente en la raiz de la diferenciacion de la especie (MST), una vez descartada cualquier influencia climatica o 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 of a series of Sier- (over 240 km in length), which is inhabited in several ras, more or less aligned in a ENE-WSW direction, which points by populations of the 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 of Iberolacerta parts of the 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 (apparently too low and dry for Iberolacerta), the raised to species level (Arribas 1996) based on allozymes Sierra de Francia with Iberolacerta martinezricai (Arribas, (Mayer & Arribas 1996), karyology (heterochromatiniza- 1996) and the main part of the Spanish portion of its range tion of sex-chromosome and localization of the NORs;

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198 Oscar J. Arribas

Odierna et al. 1996) and adult and hatchlings pattern and zor, 2.592 m), whereas the other axis is constituted by 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 lower areas, 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 Parameras just to the from the nominate one by a reduced dark pattern, more North of the Sierra de Gredos, but at their orographic shad- dorsalia, ventralia, slightly larger diameter of the masse- ow for rains, and climatically more continentalised. This teric and hindlimb length, and lower circumanalia (Arribas explains the botanical similarities between the Paramera- 1996). Serrota-Villafranca axis and the Sierra de Guadarrama (Luceno and Vargas 1991).

The degree of genetic 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 it lives 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 gested that both subspecies diverged approximately Puerto del Pico (at 1.352 m close to one of the fountains 0.8±0.2 MY BP, an estimation mainly based on the Cy- of the 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 and the nuclear gene C-mos were uninformative at this lev- favoured by accelerated cold winds in the mountain pass- el), a divergence time almost identical to the one calcu- es ( Venturi effect) had disappeared by climatic or best, by lated by Crochet et al. (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 of 0.8 MY BP). These two values were 1.837 m (own data) and 2.443 m (see Lizana et al. 1988, very similar to the above-mentioned ciphers. The inferred 1992, 1993; and Martin 2005 for general data; own data divergence time increased up to 1.2 (Cyt b) or 1 .6 MY 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 of the 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 of the /. 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 of this populations been studied so far. All aspects of morphology, status and occurred approximately 1.7±0.3 MY BP (Carranza et al. relationships of these small and isolated populations from 2004). However, in analyses using the same mtDNA re- the Sierras de Avila are totally unknown. In these Sierras gions but different taxa and other evolutionary models 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 trichotomy with /. 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 localized, also cornered in the highest parts, from 2.284 m Apart from the uninformative C-mos nuclear gene frag- (perhaps 1.935 m 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 other information about differences at the nuclear level, Serrota, 2.294 m); and in La Paramera from 1.700 m 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 of these 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 of all 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- in the light of the existence of other small and isolated pop-

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Intraspecific variability in Iberolacerta cyreni 199

illations; b) to ascertain the taxonomic status of the Bejar Cabezo, Gredos Oriental Massif, Avila province, Spain). populations and to check if these 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, or the Candelario, Salamanca province, Spain). 1 1 males and 28 more distant populations from Guadarrama and 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 (Avila province, Spain). 3 males and 2 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). These populations are discontinuous among them (Appen-

dix II) and constitute 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, and these two plus Villafranca (all of them "Sier- A total of 106/92 male specimens, and 136/135 female ras de Avila") in the female one. However, reciprocal dis- specimens of /. cyreni with a complete measurements tances between each one of these poorly represented sam- dataset and snout-vent length greater than 45 mm, were ples to the best represented ones were carefully checked included in the univariate (ANOVA) and multivariate (dis- and commented in the results section. As the three popu- criminant) analyses, respectively. Given that these populations from Sierras de Avila seemed to be largely equiv- lations present sexual dimorphism (Arribas 1996, 1999a; alent in the multivariate analyses, a posteriori, all of them

Arribas et al. 2006), analyses were carried out for males were treated as a single OTU (S a AVILA) in ANOVA. and females separately. All material is from Oscar Arribas (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 calliper to the nearest 0.01 mm. These measurements were transformed to the follow- ing more informative and not dimensional-depending ratios: 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 (relative 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-

spect to the total length, "Anal size index") (Arribas 1 996,

2001 ). The results of the linear measurements and indexes 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 of the distribution of Ibero- right and left sides; Gularia (GUL); Collaria (COLL); Dor- lacerta cyreni in the Spanish Sistema Central. The different lo- salia (DORS); Ventralia (VENT); Femoralia rigth (FEMr) calities (OTUs) cited in the text are represented. 1 : Bejar; 2: Gre- th dos; 3: Villafranca; 4: Serrota; 5: Paramera; 6: Mijares; 7: Gua- and left (FEM1); 4 . digit Lamellae (LAM); and Circum- darrama. analia (CIRCA).

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200 Oscar J. Arribas

Statistical Procedures populations permit to test significance of the differences among the concerned groups and to detect which ones are Statistical analyses used in the morphological study in- really different from the others. cluded both Univariate ( ANOVA for SVL, scalation characters and indexes, with post-hoc Tukey-Kramer tests at Mahalanobis' (squared) distance matrices were compared P < 0.05 and P < 0.01 to detect differences among sam- by means of Mantel Test (with 1000 permutations) with ples) as well as Multivariate techniques (Canonical Dis- matrices composed by Euclidean (squared) distances for criminant Analysis, CDA). In this later analysis, each pop- the climatic characteristics of localities: a) Precipitation ulation is represented by a centroid (a hypothetical mid- (mm) during the incubation months (July and August, as dle individual). Minimum-length spanning tree (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/(m 2 *day*micrometerJ(July and August), and f) Annu- distortion of UPGMA 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- tion. UPGMA frequently clusters samples reflecting sam- pared with (d) the aerial (straight) geographical distances ple sizes than their true relationships. Distances of small among the sampling localities. samples or isolated specimens appear greatly exaggerat- ed with respect to the well represented ones. As a result Multivariate (Discriminant and ANOSIM) analyses were of that, the small-sized samples appear ever as the most performed with Community Analysis Package 4.0 (Hen- external or differentiated in UPGMA derived trees derson & Seaby 2007). MST trees and Mantel tests were

(Kunkel et al. 1980; Cherry et al. 1982; Arribas 1997). calculated with NTSYS 2.1© (Rohlf 2000). Univariate sta- Moreover, the UPGMA trees based in very unevenly sized tistics were processed with NCSS 2001 0 package (Hintze samples also gave very poor Cophenetic Correlation In- 2001). dexes between the tree-derived ultrametric distances matrix and the original Mahalanobis distance matrix and therefore we have not used them (Arribas et al. 2006). RESULTS

To test the significance of the differences among pre-es- Males tablished groups for the Discriminant Analysis (based in a geographical origin), we carried out an Analysis of Sim- Canonical Discriminant Analysis. The CDA conducted ilarity (ANOSIM) (Clark 1988, 1993) that tests if the as- with 106 male specimens shows three significant axes that signed groups are meaningful, this is, more similar with- explain an 85 % of the total intersample variation. The two in groups than with samples from different groups. The first axes together explain the main part (70.6 %) and dis- method uses the Bray-Curtis measure of similarity to con- criminate fairly well the samples, especially the first one, struct clusters of specimens. 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 randomly blended). R-sta- plained; Chi-Sq. with 85 df 200.7 1 P < 0.000 1 and dis- tistic scales from +1 to -1. Values closer +1 correspond tributes the samples with fairly overlap among them, ex- to a perfect case in which all groups were completely dif- cept Guadarrama, that has only a small coincidence with ferent (all specimens of the same group are more similar the other ones (Fig. 2 A). Guadarrama appears in the neg- among them than to any specimens of the other groups). ative part of the axis, characterised by the lower values R = 0 occurs if the high and low similarities are perfect- for DORS (0.441553) and VENT (0.560765) and greater ly mixed and bear no relationship to the group, a common values of CIRCA (-0.572683). Second and third axes situation if some of the 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 of the groups (all groups largely equivalent and randomly formed). To check for significance, pseudorepli- This discriminant analysis applied to the samples reached cation tests (1000 randomizations) were run to test if the a 72.6% of correct classification among the specimens. given results can occur by chance. If the value of R is sig- The Guadarrama sample (/. cyreni cyreni) reaches a 71.9 nificant, there is evidence that the samples within groups % of correct 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

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Intraspecific variability in Iberolacerta cyreni 201

Discriminant Plot- lb cyreni MALES

~3 3 A 0 1 2 i I

Function 1

Discriminant Plot - lb cyreni FEMALES

Fig. 2. Canonical Discriminant Analysis (CDA) plots for a) males (above) and b) females (below). Specimens, sample centroids and group perimeters are represented. Guadarrama (inverted triangles), Gredos (triangles), Bejar (irregular circles), Mijares (cross), Villafranca (asterisk) and Serrota-Paramera (sail). In females, the three last samples are grouped as Sierras de Avila (sail). Sample centroids are represented by a square. See text for axis characteristics and results.

Minimum-length spanning tree (not represented) connect- Analysis of Similarity (ANOSIM) (Table 3) shows that ing the centroid (hypothetical middle specimens) of each 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- ca that connects with Gredos (at Mahalanobis Distance of graphically) assigned groups, appear only among Guadar- 3.1870), Bejar (3.2199), Paramera (4.8042) and finally, to rama and Bejar, Gredos and Villafranca (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 other populations are sample sizes: Mijares (East Gredos) that connects with Be- not differentiated among them (P > 0.01). jar (7.0951), and Serrota with their neighbouring Param- era (9.1822).

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202 Oscar J. Arribas

The Analysis of Variance (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, respectively, in the former population), but also appeared differences between Guadarrama and Bejar in Dors (smaller 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 of the 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 % of the total intersample variation. The two first axes together explain a large part of the variance (85.7 %), and especially along the first one, that accounts itself for 62.7 % of the total variation and is the unique with an eigenvalue >1 (1.77), discriminating Guadarrama specimens from the other neighbour samples only with a small overlap (Fig. 2B). The other samples show a considerable overlap among them. Guadarrama discriminates towards the negative part of the axis, characterised by the lower values of DORS (0.62) and VENT (0.66) and greater ones of CIRCA (-0.38). Second and

third axes (eigenvalues < 1) present a considerable overlap among the samples and do not discriminate populations.

The discriminant analysis applied to the samples reached a 74.26% of correct classification among the specimens.

Guadarrama sample (I. c. cyreni) reaches an 87.2 % of cor-

rect classification with respect all the other samples (I. c. castiliana).

Minimum-length spanning tree (not represented) connect-

ing centroids is very similar to the male one. The most con-

nected sample is Bejar, which clusters with Villafranca (at

2.9), Gredos (3.5) and Mijares (7.38, but here exaggerat- ed by the scarce sample of the later). Guadarrama connects with the scarcely represented (and geographically inter-

mediate) Mijares (East Gredos) (at 6.09), and all the Sier- ras de Avila samples cluster together (Villafranca with Ser- rota + Paramera at 8.66). Fig. 3. Iberolacerta cyreni castiliana. a) La Covatilla Sky re- sort (Sierra de Bejar), July 2007, Male b) El Travieso (S a de ; Analysis of Similarity (ANOSIM) (Appendix III, Table

Bejar), July 2004, Female ; c) El Calvitero (Sierra de Bejar), Ju- 3) shows that there is a considerable overlap among sam- ly 2004, Female (atypical pattern, with diffumination and coa- ples (R-statistic = < 0.001; 1000 randomiza- lescence in a unique vertebral line); d) Puerto de Mijares (Gre- 0.162588, P dos Oriental Massif), July 2006, Female. tions), but the results are slightly best than for the male

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Intraspecific variability in Iberolacerta cyreni 203

analysis. Very significant differences (P < 0.01) appear among Guadarrama and all the other samples except with Mijares. The other populations are not differentiated among them (P > 0.01).

The Analysis of Variance (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 first two characters smaller, and the third one greater in the former population). Guadarrama also dif-

fers from Gredos by its lower GUL, from Bejar by its greater FLL, HLL, a lower SVL; and from Sierras de Avi-

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 and the geographic and cli-

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 appears with very limited overlap with the other samples in the CDA graphs (Figs 2 A and B). Diagnostic characters

for this population (nominate subspecies: /. c. cyreni) are the lower values of DORS (difference more marked in females), lower values of VENT and greater CIRCA. More-

over, ANOSIM analyses show that Guadarrama is the unique OTU that is significantly different from nearly all the other samples, except from the close population of Mi- jares (in both sexes) and Serrota + Paramera (but these ex- ceptions occur only in the males and probably 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 CDA and lack differences in ANOSIM, being morpho- logically fairly equivalent and all of them assimilable to

/. c. castiliana. There are only a few scattered very significant differences among them (P < 0.01) in ANOVA, as for instance Bejar (this latter has Fig. 4. Iberolacerta cyreni castiliana. a) Pico Zapatero (Si- among Gredos and erra de la Paramera), July 2005, Male; b) Puerto de Pena Negra greater DORS and a strikingly greater SVL and propor- (Sierra de Villafranca), July 2006, Male; c) Pico Serrota (La Ser- tionally shorter limbs that Gredos, but only in female spec- rota Massif), July 2005, Female; d) Pico Serrota (La Serrota Mas- imens). The reason of the longer SVL in Bejar females sif), July 2006, Male. (from 8 mm to 1 cm greater than in the other populations)

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204 Oscar J. Arribas

which leaves proportionately shorter limbs, can be a con- 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 greater clutch size (Brana 1996). This is an in- tionship between morphological differentiation and geo- teresting question for future study: if Bejar specimens ef- graphic distances. Also there is no relationship among the fectively have greater clutch size that other/, cyreni pop- more obvious climatic parameters (precipitation, temper- ulations. ature and sun radiation) and these differences.

Both the MST results (in which Villafranca and Bejar are Concerning the position of Bejar populations, only spec- the most connected samples) as well as the presence of imens from the west-facing slopes of the massif have been related species further West (/. martinezricai and /. mon- studied, and therefore it is possible that in other parts of ticola), suggest an origin of the species towards the west- the massif other haplotypes (the common one with Gre- ern extreme of their 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 cannot be sure environment with Pyrenean Oak open forest inhabiting from which of these two mountain ranges the former was populations of Timon lepidus, Psammodromus algirus and colonized, as MST 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 of Puerto de Tor- more related to Villafranca, whereas in the female analy- navacas (1010m) makes me to suspect that /. cyreni is 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 samples (Cyt B and 12s) (unpublished, Carranza, pers. com.) An account about the pattern and coloration of /. cyreni indicates that the interruption of gene flow is fairly recent, 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 in juveniles and with respect to the common haplotype appear in Villafran- 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 linked to reproductive processes). cumulate in one Bejar and one Mijares specimen (others In populations inhabiting rocks (plenty of Rhizocarpon gr. have only one). geographicum lichens) as in Gredos and the upper parts of Guadarrama (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 of strong 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 or less with a similar age) could be due to the main- ferent degree among the different populations depending tenance of a more continuous gene flow among them, re- on the substrates inhabited. Juveniles and subadults have sponsible of maintaining their morphology largely equiv- temporal uniform or reticulate 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 of the granite phanerocrystals (the lations to the West. granite-rock spotting) and the habitus of the 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 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at

[ntraspecific variability in Iberolacerta cyreni 205

Bejar (=Candelario in Perez Mellado et al. [1993]) the de- REFERENCES

scription should be corrected as, although it is true that Almeida AP, Rosa HD, Paulo OS, Crespo EG (2002) Genetic old specimens are fairly reticulated, very especially males, differenciation of populations of Iberian rock-lizards Ibero- females more frequently have two paravertebral rows of lacerta (Iberolacerta) sensu Arribas (1999). Journal of Zoo- distinctive spots (photo 2), as in females of other popu- logical Systematics and Evolutionary Research 40: 57-64

lations and in mid-grow specimens of both sexes in all lo- Arnold EN, Arribas O, Carranza S (2007) Systematics of the Palaearctic and Oriental lizard tribe Lacertini (Squamata: Lac- calities. The statement that "the most common background ertidae: Lacertinae), with descriptions of eight new genera. colour of the back and flanks is greenish or bluish" prob- Zootaxa 1430: 1-86 ably is true for fully adult specimens during the breeding Arribas O (1996) Taxonomic revision of the Iberian 'Archaeo- period, but in July and August only some big males con- lacertae T: A new interpretation of the geographical variation serve greenish tones, appearing even fully adult females of 'Lacerta' monticola Boulenger, 1905 and 'Lacerta' cyreni more or less brownish. Mtiller & Hellmich, 1937 (Squamata: Sauria: Lacertidae). Herpetozoa9 (1/2): 31-56 Arribas OJ (1997a) Morfologia, filogenia y biogeografia de las Concluding: a) /. c. cyreni is a recently differentiated but lagartijas de alta montana de los Pirineos. Ph. D. Thesis. Uni- well diagnosable (morpho)subspecies, a case paralleling versidad Autonoma de Barcelona. (Bellaterra); 353 pp. (8 pp the relationship of /. monticola monticola from Serra da and microfiche. Pub. U.A.B.) Estrela (Portugal) with repect to /. m. cantabrica (from Arribas O (1999 a) Taxonomic revision of the Iberian 'Archae- Galicia and Cantabrian Mts.), in this later case, with no olacertae' 2: diagnosis, morphology, and geographic variation 'Lacerta' aurelioi Arribas, 1994 Sauria: genetic differences, but with singular morphological traits of (Squamata: Lac- ertidae). Herpetozoa 11: 155-180 that distinguishes it from other /. monticola in a multivari- Arribas OJ (1999 b) New data on the Pena de Francia Moun- ate analysis (Arribas & Carranza 2004; Arribas et al. ' tain Lizard 'Lacerta cyreni martinezricai Arribas, 1996. Her- variable as for instance in- 2006). Highly nuclear markers petozoa, Wien 12 (3/4): 119-128 trons or microsatellites may help to clarify definitively the Arribas OJ & Carranza S (2004) Morphological and genetic ev- status of these well diagnosable (morpho)subspecies. idence of the full species status of Iberolacerta cyreni martinezricai (Arribas, 1996). Zootaxa, Aucklan 634: 1-24 Arribas O, Carranza S, Odierna G (2006) Description of a new b) Despite the mtDNA differences of the Bejar specimens endemic species of mountain lizard from Northwestern Spain: (one or two nucleotides), their morphology is largely Iberolacerta galani sp. nov. (Squamata: Lacertidae). Zootaxa equivalent to /. c. castiliana. Lacking data from other Be- 1240: 1-55 jar populations and genetic nuclear markers, I assume that Arribas OJ (2008) Comments about the original description and these morphological similarities reflect the presence of a type specimens of Iberolacerta monticola (Boulenger, 1905). very recent gene flow with other neighbouring popula- Herpetozoa 21 (1/2): 94-95 Brana F (1996) Sexual dimorphism in lacertid lizards: male head tions. The Bejar populations are however outstanding by increase vs female abdomen increase? Oikos 75: 511-523 their female body elongation (up to near 1 cm larger), con- Carraza S, Arnold EN, Amat F (2004) DNA phylogeny of Lac- serving proportionately shorter limbs, which can be a strat- erta (Iberolacerta) and other lacertine lizards (Reptilia: Lac- egy to increase clutch size. ertidae): did competition cause long-term mountain restric- tion? Systematics and Biodiversity, London 2: 57-77 c) The Sierras de Avila populations are very similar; the Cherry LM, Case MS, Kunkel JD, Wyles JS, Wilson AC (1982) Body shape metrics and organismal evolution. Evolution 36: closer among all the populations compared. One of them, 914-933 the Sierra de Villafranca, is together with Bejar, the most Clarke KR (1988) Detecting change in benthic community struc- connected sample, and it is, from a morphological point ture. 1 3 1-142 in Oger R (eds.) Proceedings of invited papers, of view, at the root of the species expansion. Both also 14th international biometric conference, Namour, Belgium present the unique slightly variant haplotypes. All they Clarke KR (1993) Non-parametric multivariate analyses of should be considered as /. c. castiliana by their closer iden- changes in community structure. Aust. J. Ecol. 18: 117-143 Y, tity with Gredos. Crochet PA, Chaline O, Surget-Groba Debain C, Cheylan M (2004) Speciation in mountains: phylogeography and phylogeny of the rock lizards genus Iberolacerta_(Repti\ia: Lacertidae). Acknowledgements. Dr. Sergi Pla (Barcelona, Spain), Jesus Molecular Phylogenetics and Evolution, Orlando; 30: 860-866 Garcia (Huesca, Spain) and Dr. Vicente M. Ortuno (Madrid, Franzen M, Glaw F (2007) Type catalogue of Reptiles in the Zo- Spain) helped in some prospections. Dr. Salvador Carranza ologische Staatssamlung Miinchen. Spixiana 30, 2: 201-274 (Barcelona, Spain) corrected and improved parts of the manu- Henderson PA & Seaby RMH (2007) Community Analysis Pack- script. This study was financed completely by the author and age 4.0 Pisces Conservation Ltd, Lymington, UK. 164 pp. therefore it did not cost any Euro to the public arks. Prospec- (www.pisces-conservation.com) tions of the small Sierras de Avila and Bejar populations were Irwin DE (2002) Phylogeographic breaks without geographic carried out under permissions n° 20051630007003 (2005), barriers to gene flow. Evolution 56: 2383-2394 20061630024599(2006), 2007167004130 (2007), Kunkel JG, Cherry LM, Case SM, Wilson AC (1980) Reply to 20081630020386 (2008), 20092390004760 (2009), issued by the W.R. Atchley's "M-Statistic and Morphometric divergence. competent organisms in charge of the wildlife protection (Jun- Science 208: 1060-1061 ta de Castilla y Leon, Spain)

Bonn zoological Bulletin 57 (2): 197-210 ©ZFMK © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at

206 Oscar J. Arribas

lift!

954

Fig. 5. Iberolacerta cyreni NEOTYPE (here designed). MNCN 11. 39934.

Lizana M, Ciudad MJ, Perez-MelladoV (1988) Distribucion al- Miiller L & Hellmich W (1937) Mitteilungen iiber die Herpeto-

titudinal de la herpetofauna en el Macizo Central de la Sierra fauna der Iberischen Halbinsel. II. zur Kenntnis der Lacerta

de Gredos. Rev. Esp. Herpetol. 3: 55-67 monticola. Zool. Anz. 1 17: 65-73 Lizana M, Ciudad MJ, Gil M, Guerrero F, Perez-Mellado V, Ninyerola M, Pons X, Roure JM (2005) Atlas Climatico Digi- Martin-Sanchez R (1992) Nuevos datos sobre la distribucion tal de la Peninsula Iberica. Metodologia y aplicaciones en bio- de los anfibios y reptiles en el macizo central de la Sierra de climatologia y geobotanica. Universidad Autonoma de Bar- Gredos. Rev. Esp. Herpetol. 6: 61-80 celona, Bellaterra.

Lizana M, Martin-Sanchez R, Morales JJ, Lopez-Gonzalez J, index. htm> Gutierrez J (1993) Nuevas poblaciones de la lagartija serrana Odierna C, Aprea G, Arribas OJ, Capriglione T, Caputo V, 01- (Lacerta monticola cyreni) en las sierras de Avila. Bol. Asoc. mo E (1996) The Karyology of the Iberian Rock Lizards. Her-

Herp. Esp. 4: 5-6 petologica 52 (4): 542-550 Martin J (2005) Lagartija carpetana - Iberolacerta cyreni. En: Perez-Mellado V, Barbadillo LJ, Barahona F, Brown RP, Corti Enciclopedia Virtual de los Vertebrados Espanoles. Carrascal C, Guerrero F, and Lanza, B (1993) A systematic survey of LM, Salvador A (Eds.) Museo Nacional de Ciencias Natura- of the Iberian Rock lizard Lacerta (Archaeolacerta) montico-

les, Madrid, http://www.vertebradosibericos.org/ la. In: Valakos E, Boehme W, Perez-Mellado V and Maragou

Mayer W & Arribas O (1996) Allozyme differentiation and re- P (eds. ) Lacertids of the Mediterranean region. Athens: Hel- lationship among the Iberian-Pyrenean Mountain Lizards lenic Zoological Society 85-105 (Squamata: Sauria: Lacertidae). Herpetozoa 9 (1/2): 57-61 Mayer W & Arribas OJ (2003) Phylogenetic relationships of the European lacertid genera Archaeolacerta and Iberolacerta and their relationships to some other 'Archaeolacertae' (sensu la- to) from Near East, derived from mitochondrial DNA se- Received: 29.VI.2010 quences. Journal of Zoological Systematics & Evolutionary Accepted: 15.X.2010 Research 41: 157-161

tntraspecific variability in Iberolacerta cyreni 207

shape): 2.003. DM/PaL (relative masseteric plate size): 0.491. APPENDIX I DT/PaL (relative tympanic size): 0.352. AL/AW (anal plate surface): 0.6329. AS/SVL (relative anal plate size in respect to to- Neotype designation for Iberolacerta cyreni (Miiller & Hellmich, 1937) tal legth): 5.998.

The type series of Iberolacerta cyreni (Lacerta monticola cyreni Scalation: Number of supraciliary granules: 9 (rigth) and 1 (left). Supralabials: 5 (both sides). Sublabials: 6 (right side) and Miiller & Hellmich, 1937) included 66 specimens (not only from 7 (left side). Submaxillar: 6 (both sides). Gularia: 25. Collar- Guadarrama, the species' type locality, but also some paratypes ia: Dorsalia: 53. Ventralia: 26. (right) from Gredos) formerly deposited in the Zool. Staatssamlung 9. Femoral pores 19 and 18 (left). Lamellae: 25. Circumanal Plates: 8. Rostral in full con- (Herpet. Samml.) Miinchen. In fact, two syntype specimens, male and female, respectively numbered "ZSM (SLM) 2329 a" tact with frontonasal. Supranasal separated from first loreal. One and "ZSM (SLM) 2329 b", were originally considered as types postnasal (in both sides). First Postocular separated from Pari- labelled "Guadarrama, Puerto de Navacerrada. W. Hellmich". etal plate. Alternate wide and narrow scale rings in the tail. Twen- ty-six scales across one of these rings. (Miiller & Hellmich, 1937).

Coloration: (in alcohol). Dorsal tract and pileus brown (proba- Although it seems that all the original type series was destroyed bly also in life), densely spotted of medium-sized black spots during the Second World War (Franzen & Glaw, 2007), and due that in the the nearly transverse to the fact that in this type series there were included some Gre- middle of dorsum form bands and connect the two temporal bands. Temporal bands reticulat- dos specimens (today part of another subspecies; /. c. castiliana), ed fairly dark (black-brown) with traces of clear occelli (bare- and also that there was early confusion about the /. monticola ly visible) inside, also connecting with light reticulated with type locality which lead to the description of a new taxon as Lac- more infratemporal band (barely discernible). No blue axillar occel- erta estrellensis Cyren, 1 928 (Arribas 2008), I design a new type li. Traces of blue points in the outermost ventral scales. specimen (neotype) to fix unequivocally the type locality against Only the ventral scale are clearly spotted. Bel- any contingence (as could be the highly improbable apparition outermost ranges black of any "surviving" original Gredos paratype). ly light bluish or white-bluish.

I designate here as NEOTYPE for the species a specimen from the Museo Nacional de Ciencias Naturales (Madrid) (MNCN n. 39934) (Fig. 5). APPENDIX II A male labelled as follows: Barriers and high mountain passes (among parentheses) between Left hindleg: (white label, Typewriter letter) MNCN (anverse), the different /. cyreni populations. All these intermediate areas 39934 (reverse). are at present apparently devoid of /. cyreni, thus constituting these discrete populations: Right hindleg: (white label, pencil handwritten) "Pto. de OTUs Cotos-Pto de Navacerrada. Srra. de Guadarrama [blank in- ( ) BEJAR GREDOS: no barrier (Puerto de Tornavacas, 1275m). side parenthesis], 21-IV-84, 18,15 h. Sol. Pedriza en pinar con nieve. Ps=8.5 gr." (no collector's data). BEJAR-VILLAFRANCA: Tormes River Valley (no pass).

Left foreleg: (White label, ink handwritten) Neotypus. O. Ar- GREDOS-VILLAFRANCA: Tormes River Valley (Collado de ribas designatio (anverse), "Lacerta monticola cyreni Miiller & Cepegato, 1550m). Hellmich, 1937" (=Iberolacerta cyreni) (reverse). GREDOS-Sa AVILA[Serrota+Paramera]: Alberche River Valley Right foreleg: (Red plastic label, Dymo® lettering) NEOTYPUS. (no passes).

Neotype description (Fig. 5): SERROTA-PARAMERA: no barrier (Puerto de Menga, 1566m). Biometry: Adult male with snout-vent length of 66.85 mm. Tail 126 mm (intact). Forelimb length 23.34 mm. Hindlimb length VILLAFRANCA-SERROTA: no barrier (Puerto de Chia, 34.52 mm. Pileus length 16.4 mm. Pileus width 8.2 mm. Pari- 1663m). etal legth 5.5 mm. Masseteric widest diameter 2.71 mm. Tym- a S AVILA (as a whole) - GUADARRAMA: no barrier (Puerto panic widest diameter 1 .94 mm. Anal plate width 5.04 mm. Anal del Boqueron, 1315m). length 3.19 mm. FLL/SVL (relative forelimb length): 0.349. HLL/SVL (relative hindlimb length): 0.5163. PL/PW (pileus GREDOS-GUADARRAMA: Alberche River Valley (no pass)

208 Oscar J. Arribas

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210 Oscar J. Arribas

Table 3. Analysis of Similarity (ANOSIM) results (with 1000 randomizations). Males above diagonal and females below. The number is the test probability results among each two concerned populations (significant results in bold).

FEM \ MAL BEJAR GREDOS GUAD. MIJARES SERR-PAR VILLAFR.

BEJAR 0.058 0.001 0.644 0.138 0.064

GREDOS 0.021 0.001 0.691 0.867 0.198 GUAD. 0.001 0.001 0.097 0.128 0.001

MIJARES 0.53 0.039 0.327 0.224 0.513 SERR-PAR 0.127 0.025 0.001 0.217 0.86