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1 This article was downloaded by: [b-on: Biblioteca do conhecimento online UP] On: 11 June 2012, At: 02:44 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK African Journal of Herpetology Publication details, including instructions for authors and subscription information: Genetic variability and relationships within the skinks Eumeces algeriensis and Eumeces schneideri using mitochondrial markers Ana Perera a, Filipa Sampaio a, Sara Costa a, Daniele Salvi a & D. James Harris a b a CIBIO-UP, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, , Vairão, Portugal b Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, , Porto, Portugal Available online: 03 Jun 2011 To cite this article: Ana Perera, Filipa Sampaio, Sara Costa, Daniele Salvi & D. James Harris (2012): Genetic variability and relationships within the skinks Eumeces algeriensis and Eumeces schneideri using mitochondrial markers, African Journal of Herpetology, 61:1, To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

2 African Journal of Herpetology, Vol. 61, No. 1, April 2012, 6980 Short communication Genetic variability and relationships within the skinks Eumeces algeriensis and Eumeces schneideri using mitochondrial markers ANA PERERA 1 *, FILIPA SAMPAIO 1,SARA COSTA 1, DANIELE SALVI 1 &D.JAMES HARRIS 1,2 1 CIBIO-UP, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, , Vairão, Portugal; 2 Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, , Porto, Portugal Abstract.*The skinks Eumeces algeriensis and Eumeces schneideri are two of the most widespread species of the E. schneideri group. Despite this, data on their intra-specific variation are limited. In this study we analyse the genetic variability of these two species across their distribution range using two mitochondrial fragments, 12S rrna and 16S rrna. The results confirm the paraphyly of the group, with E. algeriensis more related to Scincopus than to E. schneideri. Eumeces algeriensis shows relatively low levels of genetic diversity, which is distributed in two main lineages, one across Morocco, and the other restricted to the southern region (Anti-Atlas). These results are discordant with the high diversity found in other species from the same area and suggest a relatively recent expansion of E. algeriensis across most of its current range. Regarding E. schneideri, we find high levels of genetic variability and a complex pattern of genetic diversity, concordant with the phylogeographic patterns found recently in other species from the Middle East region, but not with the current intra-specific taxonomy of this group. Key words.*eumeces algeriensis, Eumeces schneideri, North Africa, Middle East, mtdna, genetic diversity The genus Eumeces, considered one of the basal groups within the Scincidae (Estes et al. 1988; Wiens et al. 2006), includes more than 50 species spread across the Holarctic region. Long considered monophyletic, Griffith et al. (2000) indicated it may be paraphyletic, and since then, various authors have presented alternatives to split Eumeces into different genera (Schmitz et al. 2004; Smith 2005). The E. schneideri group (sensu Taylor 1935) is distributed across the Saharo-Sindian region from North Africa, Cyprus and the Middle East to southwest Asia. Renamed as Novoeumeces by Griffith et al. (2000), most authors rejected this proposal and suggested to maintain the use of Eumeces for the schneideri group, as they contain the type species of the genus (Brandley et al. 2005). Within this group, different authors recognised the existence of a variable number of species (between two and six; Taylor 1935; Lieb 1985; Sindaco & Jeremcenko 2008), with two of them being widely accepted: E. schneideri (Daudin 1802), distributed from eastern Maghreb to West Asia and E. algeriensis Peters 1864, distributed across the western Maghreb. Following Sindaco & Jeremcenko (2008), six subspecies within E. schneideri are *Corresponding author. perera@mail.icav.up.pt ISSN print/issn online # 2012 Herpetological Association of Africa

3 70 PERERA ET AL.*Genetic variation in Eumeces recognised: E. s. aldrovandii from North Africa; E. s. blythianus from Pakistan, Afganistan and Punjab; E. s. pavimentatus from SE Anatolia to Levant; E. s. princeps distributed across East Turkey, Iraq, North and West Iran, Afghanistan, and nearby countries; E. s. zarudnyi from SE Iran and SW Pakistan; and E. s. barani recently described in western Turkey (Kumlutas et al. 2007). Regarding E. algeriensis, two subspecies E. a. algeriensis, widely distributed across Morocco; and E. a. meridionalis, restricted to East Morocco and NW Algeria are recognised (Sindaco & Jeremcenko 2008). However, the taxonomic position of E. a. meridionalis is still under debate (Schleich et al. 1996; Bons & Geniez 1996; Sindaco & Jeremcenko 2008). A recent study by Carranza et al. (2008) including North African specimens from both species highlighted the paraphyly of this group, with Scincopus as sister taxa of E. algeriensis, and E. schneideri closer to Scincus, as already suggested in previous karyological (Caputo et al. 1993, 1994) and morphological studies (Arnold & Leviton 1977; Caputo et al. 1993). Moreover, the study confirmed E. schneideri and E. algeriensis as two different entities with a disjunct distribution in North Africa (Carranza et al. 2008). However, the variability within each unit is still poorly known since, until now, only five individuals for each species have been analysed (Carranza et al. 2008). North Africa and the Middle East are among the most diverse regions in the Western Palaearctic, with a wide variety of geological, geographical and climatic characteristics shaping a diversity of landscapes (McColl 2005). Moreover, both regions were refugia for biota during the Pleistocene climatic fluctuations (De Lattin 1949; Bons 1973; Abed & Yaghan 2000). The phylogeography of various reptiles from the western Maghreb have been studied in recent years, and in general unexpectedly high levels of genetic variation have been reported (Brown et al. 2002; Rato & Harris 2008; Perera & Harris 2010), often supporting the existence of cryptic taxa (Perera et al. 2007; Pinho et al. 2008, Rato et al. 2010). Regarding the Middle East, the few studies existing also indicate that this area is a source of considerable genetic variability for many taxa (Kapli et al. 2008; Gvozdík et al and references within). Given this, the assessment of genetic diversity within E. algeriensis and E. schneideri is essential to understand better the complexity of patterns and processes underlying the diversity in these two regions. In this study, we analysed 22 individuals from E. algeriensis and E. schneideri from Morocco and the Middle East together with sequences from 14 specimens available in GenBank, and individuals of the genus Scincus and Scincopus. The main goal of this study was to analyse the genetic variability within E. algeriensis and E. schneideri. Additionally, we analysed in a phylogenetic framework the relationships between E. algeriensis and E. schneideri and those between them and representatives of the genera Scincus and Scincopus. Fresh samples were collected from lizards captured by hand, identified based on their morphological characteristics, and a small piece of the tail taken and stored in 96% ethanol for genetic analysis. All individuals were released immediately afterwards at the same place of capture. Additional tissue samples were obtained from voucher specimens from the National History Museum of Crete (NHMC; Table 1). In total, 15 new samples of E. algeriensis and 7 of E. schneideri (Fig. 1; Table 1) were analysed genetically using 2 different mitochondrial DNA fragments 12S rrna and 16S rrna. Additionally, 14 sequences (7 E. algeriensis and 7 E. schneideri) from GenBank were included (Table 1). Finally, in order to investigate

4 Table 1. Samples analysed in this study. For each specimen, locality, GenBank accession numbers (provided after acceptance of the manuscript) and origin is provided. GenBank accession numbers Species Code Locality 12S rrna 16S rrna Origin Eumeces algeriensis algeriensis EA1 Taroudant, Morocco JF JF This study Eumeces algeriensis algeriensis EA2 Agadir - Tiznit, Morocco JF JF This study Eumeces algeriensis algeriensis EA4 Oulad Brahim, Morocco JF JF This study Eumeces algeriensis algeriensis EA5 Volubilis, Morocco JF JF This study Eumeces algeriensis algeriensis EA6 Moulouya Valley, Morocco JF JF This study Eumeces algeriensis algeriensis EA7 Moulouya Valley, Morocco JF JF This study Eumeces algeriensis algeriensis EA8 Ouazzane, Morocco JF JF This study Eumeces algeriensis algeriensis EA9 Mechra Ben Abhou, Morocco JF JF This study Eumeces algeriensis algeriensis EA10 Mechra Ben Abhou, Morocco JF JF This study Eumeces algeriensis algeriensis EA11 Agadir Tiznit, Morocco JF JF This study Eumeces algeriensis algeriensis EA12 Tirhmi, Morocco JF JF This study Eumeces algeriensis algeriensis EA13 Bin-El-Ouidanne, Morocco JF JF This study Eumeces algeriensis algeriensis EA14 Gorges near Guelmin, Morocco JF JF This study Eumeces algeriensis algeriensis EA15 Between Casablanca and Rabat, Morocco JF JF This study Eumeces algeriensis algeriensis EA18 Near Oued Ouahar, Morocco JF JF This study Eumeces algeriensis algeriensis GB1 Massa, Morocco EU EU Carranza et al. (2008) Eumeces algeriensis algeriensis GB2 Essaouira, Morocco EU EU Carranza et al. (2008) Eumeces algeriensis algeriensis GB3 Ras el Ma, Morocco EU EU Carranza et al. (2008) Eumeces algeriensis GB4 Morocco EU EU Carranza et al. (2008) Eumeces algeriensis GB13 Morocco EU EU Carranza et al. (2008) Eumeces algeriensis GB14 AY AY Schmitz (2003) Eumeces algeriensis GB15 AY AY Schmitz (2003) Eumeces schneideri pavimentatus 1 ES10 Al Jaboul lake, Syria JF JF This study 2 (NHMC ) Eumeces schneideri pavimentatus 1 ES11 Azraq, Jordan JF JF This study 2 (NHMC ) Eumeces schneideri pavimentatus 1 ES13 Um Quays, Jordan JF JF This study 2 (NHMC ) AFRICAN JOURNAL OF HERPETOLOGY 61(1)

5 Table 1 (Continued ) GenBank accession numbers Species Code Locality 12S rrna 16S rrna Origin Eumeces schneideri pavimentatus 1 ES14 Dana Nature Reserve, Jordan JF JF This study 2 (NHMC ) Eumeces schneideri pavimentatus 1 ES17 Lattakia Beach, Syria JF JF This study 2 (NHMC ) Eumeces schneideri pavimentatus 1 ES18 Petra, Jordan JF JF This study 2 (NHMC ) Eumeces schneideri pavimentatus GB7 Coastal dunes after Karatas, Turkey EU EU Carranza et al. (2008) Eumeces schneideri pavimentatus GB8 Karaotlak in the Euphrates Valley, Turkey EU EU Carranza et al. (2008) Eumeces schneideri princeps ES19 Echegnatzor, Armenia JF JF This study Eumeces schneideri princeps GB5 Igdir, Turkey EU EU Carranza et al. (2008) Eumeces schneideri schneideri GB6 Egypt EU EU Carranza et al. (2008) Eumeces schneideri schneideri GB16 Egypt EU EU Carranza et al. (2008) Eumeces schneideri GB11 AY AY Schmitz (2003) Eumeces schneideri GB12 West Asia AB AB Honda et al. (2000) Scincus albifasciatus SC14 Around Ayoun el Atrous, Mauritania EU EU Carranza et al. (2008) Scincus albifasciatus SC17 Boû Derga, Mauritania EU EU Carranza et al. (2008) Scincus mitranus SC11 El Ain, UAE EU EU Carranza et al. (2008) Scincus mitranus SC12 UAE AY AY Brandley et al. (2005) Scincus scincus SC3 Unknown AY AY Schmitz et al. (2005) Scincus scincus SC4 AY AY Whiting et al. (2003) Scincus conirostris SC6 Jabal Dannah, UAE EU EU Carranza et al. (2008) Scincopus fasciatus SCP2 ca.30km NW Rosso, Mauritania AY AY Brandley et al. (2005) Scincopus fasciatus SCP3 AY AY Schmitz (2003) 1 Individuals whose subspecific taxonomy could not be assessed and were predicted based on geographical location following Eiselt (1940). 2 Samples obtained from specimens deposited in the Natural History Museum of Crete. Voucher codes are detailed in parenthesis. 72 PERERA ET AL.*Genetic variation in Eumeces

6 AFRICAN JOURNAL OF HERPETOLOGY 61(1) Figure 1. Distribution map of the Eumeces algeriensis and E. schneideri samples included in the study. The small maps show the exact location of the samples for E. algeriensis and E. schneideri. Different symbols represent different lineages (see text for more details). Samples with approximate location are marked with (*), while samples of unknown location are not shown (see Table 1 for more details). the paraphyly of Eumeces suggested in previous studies (Carranza et al and references within), sequences from two Scincopus and seven Scincus (Table 1) were selected from GenBank and incorporated into the analysis. Genomic DNA was extracted from the tail tips following standard saline method (Sambrook et al. 1989). Two mitochondrial DNA fragments, 12S rrna and 16S rrna were amplified through polymerase chain reaction (PCR). Primers used were 12sa and 12sb for the 12S rrna (Kocher et al. 1989) and 16sH and 16sL for the 16S rrna (Palumbi 1996). In both cases, the PCR mix was carried out in a 25ml total volume, with the following conditions: an initial cycle of 928C for 2 min, followed by 30 cycles of 928C for 30 s, 508C for 40 s and 728C for 45 s, and a final cycle of 728C for 5 min. Amplified products were sequenced on an ABI 3730XL automated sequencer using commercial sequencing facilities (Macrogen Inc., Seoul, South Korea). All sequences obtained were aligned using the ClustalW software (Thompson et al. 1994) implemented in MEGA4 (Tamura et al. 2007). Resulting fragments of 373 base pairs (bp) for 12S rrna and 503 bp for 16S rrna were concatenated obtaining a final fragment of 876 bp. Chalcides striatus (GenBank accession numbers EU and EU278067) and C. minutus (GenBank accession numbers EU and EU278063) were included as outgroups (Carranza et al., 2008). Poorly aligned positions of the concatenated sequences were detected and eliminated using the Gblocks software (online version 0.91b; Castresana 2000) considering a relaxed

7 74 PERERA ET AL.*Genetic variation in Eumeces selection of blocks (Talavera & Castresana 2007). A final set of sequences of 863 bp length was used in the phylogenetic analyses. Two different phylogenetic approaches, maximum likelihood (ML) and Bayesian inference (BI) were used for the concatenated fragment, implementing the two genes using a partitioned analysis. The best models of nucleotide substitution for each gene were calculated in jmodeltest (Posada 2008) under the Akaike Information Criterion (AIC; Posada 2008). ML was carried out in TREEFINDER (Jobb et al. 2004). We used a partitioned analysis and the bootstrap analysis option with 1000 replicates to assess the nodal support for the ML tree (BP). Bayesian inference was conducted in MrBayes v (Huelsenbeck & Ronquist 2001). Analysis started with a randomly generated tree and was run for generations using a partitioned dataset and the most appropriate evolutionary model for each partition, sampling one tree each 100 generations. Two independent replicates were conducted in parallel (Huelsenbeck & Bollback 2001) using the default heating values. Convergence of the MCMC chains was assessed using the online program AWTY (Wilgenbusch et al. 2004) and the data sampled from generations preceding the stationarity were discarded. The remaining trees were used to assess posterior probabilities (BPP) for nodal support. In addition to the tree-building methods, we analysed the genealogical relationships among E. algeriensis and E. schneideri haplotypes across their ranges constructing a Median-Joining network (Fig. 2) using the program Network v.4.5 (# Fluxus Technology; Bandelt et al. 1999). Poorly aligned positions from subsets of the data (E. algeriensis and E. schneideri) were eliminated using Gblocks following the same procedure as described previously. Uncorrected p-distances were calculated using the software MEGA4 (Tamura et al. 2007). The amount of genetic variation within E. algeriensis and E. schneideri was estimated as haplotype (h) and nucleotide (p) diversity and as average number of nucleotide differences (K) using the software DnaSP v. 5 (Librado & Rozas 2009). The hypothesis of past population expansion was tested by means of statistics based on the distribution of mutation frequencies such as Tajima s D (Tajima 1989) and R 2 (Ramos-Onsins & Rozas 2002), and on haplotype distribution such as F S (Fu 1997). Of the 863 bp analysed, 227 positions were variable and 206 were parsimonyinformative. The best evolutionary models were TIM2 G for 12S rrna and TPM2uf IG for 16S rrna. Trees obtained in ML and BI phylogenetic approaches had similar topology, with the exception of minor branches in the E. schneideri clade (Fig. 3). Our results confirm the paraphyly of Eumeces, with E. algeriensis and Scincopus fasciatus, as sister taxa, and E. schneideri related to them. The relationship between Scincopus and E. algeriensis is well supported (BP/BPP 89/99; Fig. 3). This result was already reported in previous studies (Griffith et al. 2000; Schmitz et al. 2004; Carranza et al. 2008; Giovannotti et al. 2009). Divergence between S. fasciatus and E. algeriensis was estimated to be 10.5% uncorrected p-distance (12S and 16S rrna concatenated), similar to the values found previously for 12S rrna (10.8%; Carranza et al. 2008). Within E. algeriensis from Morocco, genetic diversity is low (h0.589, p0.008, K 6.654), but two divergent lineages geographically congruent and well supported (BP/BPP99) are observed. The first one is widespread across Morocco and includes most of the individuals analysed, while the second is restricted to

8 AFRICAN JOURNAL OF HERPETOLOGY 61(1) Figure 2. Median Joining Network based on the mitochondrial fragment (12S rrna and 16S rrna concatenated). (a) Median Joining Network for E. algeriensis based on a 863 bp fragment; (b) Median Joining Network for E. schneideri based on 861 bp. Grey circles represent different haplotypes with size proportional to the sample frequency, and black circles represent missing haplotypes (extinct or not sampled). the southern region (Anti-Atlas) with the exception of a southern locality (Agadir-Tiznit) where individuals from both clades are found (EA2 and EA11). It is noticeable the low intra-group genetic diversity in both clades (widespread lineage: h0.350, p0.001, K 0.500; southern lineage: h 0.733, p0.003, K 2.267). The Median-Joining network also illustrates this, with only eight different haplotypes distributed in two groups (five from the widespread lineage, and three from the southern one), separated by 18 mutational steps from each other (Fig. 2). These results are unexpected, as other studies in the region report, in general, high levels of variability (Rato & Harris 2008; Rato et al. 2010) and occasionally, the existence of differentiated lineages in the Anti-Atlas region (Perera & Harris 2010). The low

9 76 PERERA ET AL.*Genetic variation in Eumeces Figure 3. Phylogenetic tree based on the 863 bp mtdna fragment (12S rrna and 16S rrna concatenated). Topology of the tree corresponds to the maximum likelihood (ML) analysis. Numbers separated by slash near the nodes represent the bootstrap support (BP) values for the ML analysis, and the Bayesian posterior probability values (BPP), respectively. * and ** refer to BP and BPP values above 95 and 98 respectively, while values with full support for both BP and BPP are represented by a dark rectangle. (1) indicates individuals whose subspecific taxonomy was not clear and that were designated on the basis of their geographical location following Eiselt (1940). variability within E. algeriensis reported here might be tentatively explained as the result of a recent expansion of this species across most of its current range, facilitated by the higher dispersal abilities related to the large body size of these skinks. This hypothesis is supported by both significant large negative values of the Tajima s D (Tajima s D1.831; PB0.05), and F S (F S 1.790; PB0.02) statistics, although the latter was marginally significant. By contrast, the low but not significant value of the R 2 statistic (R ; P0.05), did not support a past sudden expansion scenario. The finding of a locality in the south where both lineages co-occur also supports a fast expansion scenario and might indicate that both lineages were previously confined to the south of the Atlas, and in the relatively recent past only one of them expanded northwards to occupy the current range. With all this information, and taking into account the wide distribution of this Moroccan lineage and the shape of the network (Fig. 3), we can tentatively hypothesize a past geographic-demographic expansion for this lineage, although further data are necessary to fully test this hypothesis. Finally, our sampling did not include any individual from the subspecies E. a. meridionalis. This subspecies has been reported from a few localities across the High Plateaux from East Morocco and NW Algeria (Bons & Geniez 1996;

10 AFRICAN JOURNAL OF HERPETOLOGY 61(1) Sindaco & Jeremcenko 2008), and has been considered by authors as part of E. algeriensis (Caputo et al. 1993), E. schneideri (Eiselt 1940) or a unique species (Schleich et al. 1996). The two subspecies present in Morocco occupy similar habitats and have the same colour pattern and size, differing only on scalation characters (Schleich et al. 1996). Future analysis of specimens of E. a. meridionalis will be fundamental to clarify the relationships within these two groups and to confirm the validity of this form. Regarding E. schneideri, considerable within group diversity is observed (h0.945, p0.017, K ; Fig. 3) with a remarkable genetic differentiation between haplotypes (Fig. 2). The phylogenetic tree reveals three well-supported lineages corresponding to the three subspecies of E. schneideri included in this study, although the relationships among them are not well supported (Fig. 3). One group is constituted by E. s. princeps from northeast Turkey and South Armenia. A second group is formed by E. s. schneideri from Egypt, plus the sample from West Asia and another of unknown origin (Figs. 1 and 3). Finally, the third group is constituted by E. s. pavimentatus, that includes several lineages with some geographical concordance. Eumeces s. pavimentatus from Syria (ES10) and Turkey (GB7 and GB8) show closely related haplotypes. Interestingly, despite the fact that sample GB8 fits within the E. s. princeps distribution according to Kumlutas et al. (2007), based on our mitochondrial DNA sequences, it is E. s. pavimentatus. This result expands the distribution range (Kumlutas et al. 2007) of the subspecies in Turkey. Another group is formed by the samples from Jordan and Syria, and finally, the remaining samples from Jordan formed a group with some geographical congruence, although not well supported (Fig. 3). The Median-Joining network confirmed the high variability within E. schneideri, with 10 different haplotypes in 14 individuals, and a distance of 34 mutational steps between the most divergent haplogroups (3.2% uncorrected p-distance; Fig. 2). The two E. s. princeps samples share the same haplotype, which was well separated from the remaining E. schneideri samples. Interestingly, the genetic differentiation between E. s. pavimentatus and E. s. schneideri haplogroups (1623 mutational steps) was similar to that observed within E. s. pavimentatus haplotypes from different regions of Jordan (1517 mutational steps) or Syria (22 mutational steps) (Fig. 2). Several E. schneideri taxa (species/subspecies depending on the authors) have been described for the Middle East region, mostly based on morphological features (Eiselt 1940; Göçmen et al. 2002; Kumlutas et al. 2007). Our results, although limited in their interpretation we could not assign to the subspecies level some of the individuals provide interesting information regarding the genetic variability of Eumeces in this region. Genetic differentiation within E. schneideri is quite high and not completely congruent with the current intra-specific taxonomy. Indeed, while E. s. princeps forms a well supported and differentiated entity, E. s. pavimentatus has a more complex structure, reflected in its major haplotypic diversity and genetic differentiation. Interestingly, Werner (1988) observed ecological differences between E. schneideri and E. pavimentatus, one being more linked to Mediterranean habitats and the other more associated to desert areas, respectively. Although samples from Jordan and Syria were assigned to E. s. pavimentatus based on geographical location (following Eiselt (1940) and Sindaco & Jeremcenko (2008)), the genetic differentiation observed suggests the occurrence of several undescribed entities from Syria and Jordan. In addition, the distribution of the lineages found within this region has some

11 78 PERERA ET AL.*Genetic variation in Eumeces geographical structure (north, central and southern Middle East) concordant with patterns observed in Hyla frogs (Gvozdík et al. 2010). Studies in this region that are now emerging, confirm the complexity of this area (Gvozdík et al. 2010; Kornilios et al. 2010) and our results also point in this direction. The Middle East is a crossroad of Palearctic, Oriental and Afrotropic ecozones and has been considered a refugia for the herpetofauna during Pleistocenic climatic oscillations (Hewitt 1999; Fritz et al. 2008; Gvozdík et al. 2010). It will be interesting in the future to increase the sampling in this region and surrounding areas, to complete the picture of the variation of Eumeces across this region. In conclusion, our results support once more the paraphyly of Eumeces, suggesting the need of a taxonomic reassessment for this genus. Moreover, this preliminary assessment of the genetic diversity within E. algeriensis and E. schneideri has revealed surprising patterns. The former shows a relatively low genetic variability in North Africa and the existence of two widely divergent lineages, which might suggest a history of a recent expansion. Regarding the latter, complex patterns, not completely congruent with the currently described subspecies, have been retrieved. Our results confirm the need of a deeper assessment in the Middle East region in order to understand the exact distribution of the different subspecies and their genetic differentiation. ACKNOWLEDGEMENTS We are grateful to all the people from CIBIO that collaborated in the sampling in Morocco. This study was partially funded by the FCT project PTDC/BIA-BDE/ 74349/2006. AP is supported by a FCT grant SFRH/BPD/26546/2006, DS by a FCT grant SFRH/BPD/66592/2009 and SC and FS did the study under the FCT program BII/CIBIO/INTBIO/2009. We are especially grateful to Petros Lymberakis from the National History Museum of Crete for the donation of Eumeces samples. REFERENCES ABED, A.M. & R. YAGHAN On the paleoclimate of Jordan during the last glacial maximum. Palaeogeogr. Palaeocl. 160: ARNOLD, E.N. & A.E. LEVITON A revision of the lizard genus Scincus (Reptilia: Scincidae). Bull. Brit. Mus. Nat. Hist. Zool. 3: BANDELT, H., P. FORSTER & A. ROHL Median-joining networks for inferring intraspecific phylogenies. Mol. Biol. Evol. 16: BONS, J Herpétologie marocaine II. Origines, évolution et particularités du peuplement herpétologique du Maroc. B. Soc. Sci. Nat. Maroc 53: BONS, J.& P. GENIEZ Amphibians and Reptiles of Morocco. Asociación Herpetológica Española, Barcelona. BRANDLEY, M.C., A. SCHMITZ & T.W. REEDER Partitioned Bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards. Syst. Biol. 54(3): BROWN, R., N.M. SUAREZ & J. PESTANO The Atlas mountains as a biogeographical divide in North- West Africa: evidence from mtdna evolution in the Agamid lizard Agama impalearis. Mol. Phylogen. Evol. 24(2): CAPUTO, V., G. ODIERNA, G. APREA & T. CAPRIGLIONE Eumeces algeriensis, a full species of the E. schneiderii group (Scincidae): karyological and morphological evidence. Amphibia-Reptilia 14:

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