Zoologica Scripta

Forgotten in the ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago

R. SINDACO,M.METALLINOU,F.PUPIN,M.FASOLA &S.CARRANZA

Submitted: 30 August 2011 Sindaco, R., Metallinou, M., Pupin, F., Fasola, M. & Carranza, S. (2012). Forgotten in the Accepted: 2 February 2012 ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra doi:10.1111/j.1463-6409.2012.00540.x Archipelago. —Zoologica Scripta, 41, 346–362. The Socotra Archipelago, in the north-west Indian Ocean, is considered to be one of the most remote and most biodiversity rich and distinct islands in the world. Often referred to as the ‘Galapagos of the Indian Ocean’, it was designated a UNESCO World Heritage Natural site in 2008. Despite having a very rich and bizarre fauna and flora with a high level of ende- micity at both species and generic levels, the taxonomy of most of the groups is still not clear, and their origin and evolution remain unknown. Reptiles constitute the most relevant verte- brate fauna of the Socotra Archipelago, with 90% of the 30 species and 45% of the 12 genera being found nowhere else in the world. The skinks of the endemic species Trachylepis socotr- ana are the only reptile species in the Archipelago distributed across all four islands (Socotra, Darsa, Samha and Abd Al Kuri). Although the species is very well known from Socotra Island, it was not discovered on Samha until 1999 and on Darsa until 2000, whereas only a few citations and one single Museum specimen exist for the population from Abd Al Kuri. To clarify the systematics, biogeography and evolution of Trachylepis socotrana, we assembled a dataset for Mabuya sensu lato including 904 base pairs (bp) of sequence (392 bp from the 12S and 512 from the 16S rRNA mitochondrial genes) for 115 individuals, including specimens of T. socotrana from all four island populations, numerous representatives of the genus Trachylepis from the Middle East, Africa and Madagascar, plus some individuals from each of the other three genera of Mabuya sensu lato (Chioninia, Eutropis and Mabuya). The results of the phylogenetic analyses indicate that, contrary to what was previously thought, members of the genus Trachylepis have colonized the Socotra Archipelago in two independent events, with the first giving rise to the populations from Socotra, Samha and Darsa and the second to the Trachylepis from Abd Al Kuri Island. According to the calibrations, both colo- nization events occurred within the last fourteen million years, when the Socotra Archipelago had already drifted away from Arabia, thus ruling out vicariance. Both morphological and genetic data show that the Trachylepis from Abd Al Kuri is a distinct taxon, which is herein described as a new species belonging to the T. brevicollis species complex. On the basis of this evidence, the terrestrial herpetofauna from Abd Al Kuri is composed exclusively of endemic species (one of which, the gecko Pristurus abdelkuri, was introduced into some parts of Socotra). Corresponding author: Salvador Carranza, Institute of Evolutionary Biology (CSIC-UPF), Passeig Marı´tim de la Barceloneta, 37-49, E-08003 Barcelona, Spain. E-mail: salvador.carranza@ ibe.upf-csic.es Roberto Sindaco, Museo Civico di Storia Naturale, via San Francesco di Sales 188, I-10022 Carmagnola, Italy. E-mail: [email protected] Margarita Metallinou, Institute of Evolutionary Biology (CSIC-UPF), Passeig Marı´tim de la Barce- loneta 39-47, 08003 Barcelona, Spain. E-mail: [email protected] Fabio Pupin, Museo Tridentino di Scienze Naturali, Via Calepina 14, I-38122 Trento, Italy. E-mail: [email protected] Mauro Fasola, Dipartimento di Biologia Animale, Universita` di Pavia, Via Ferrata 1, I-27100 Pavia, Italy. E-mail: [email protected] Salvador Carranza, Institute of Evolutionary Biology (CSIC-UPF), Passeig Marı´tim de la Barcelon- eta 39-47, 08003 Barcelona, Spain. E-mail: [email protected]

346 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Sindaco et al. d The Trachylepis skinks of the Socotra Archipelago

Introduction 2004; Bosworth et al. 2005). Tectonic activity in the Red The Socotra Archipelago comprises four islands of conti- Sea ⁄ Gulf of Aden region during the Oligocene produced nental origin situated in the north-west Indian Ocean, arching and block faulting along the axis of the embryo near the Gulf of Aden (Fig. 1). It is considered one of the Gulf, thereby giving rise to the Gulf of Aden and the most remote and most biodiversity rich and distinct archi- east–west faults of southern Arabia (Laughton 1966). Sub- pelagos in the world. Termed the ‘Galapagos of the Indian sequent sub-crustal movements also produced forces that Ocean’, it was designated a UNESCO World Heritage both carried the Arabian block north–north-east and Natural site in 2008 (Van Damme 2009). The easternmost rotated it anticlockwise, thus producing a line of parting and largest of these four islands is also called Socotra and, between the African and Arabian continental blocks that at just 3625 km2, comprises about 95% of the total land- passed between Socotra and the Hallaniyat Islands (for- mass of the archipelago. It is also the most ecologically, merly Kuria Muria Islands) in south-west Oman (Fig. 1). geographically and biologically diverse. Abd Al Kuri, the These tectonic movements marked the onset of the rifting second largest (133 km2) and westernmost island of the between Arabia and Somalia in the late Oligocene ⁄ early archipelago, lies about 105 km to the west of Socotra and Miocene (30–17.6 mya). This period, also known as the 100 km east of the Horn of Africa. The other two islands, syn-rift time, was followed by the postrift time, which was namely Darsa (16 km2) and Samha (40 km2), also known characterized by continental break-up and oceanic spread- as ‘The Brothers’, are much smaller and are situated 36 ing at 17.6 mya in the eastern part of the Gulf of Aden and 50 km, respectively, to the west of Socotra. Two and increased the distance between the Socotra Archipel- rocky guano islets (Ka’l Fir’awn and Sabuniyah) situated ago and Arabian mainland (Laughton 1966; Bosworth 20 km north of Abd Al Kuri and 20 km north-east et al. 2005; Autin et al. 2010). of Socotra, respectively, are apparently devoid of As the Socotra Island is separated from Samha and herpetofauna. Darsa by shallow seas, it has been suggested that they The Socotra Archipelago rests on a shelf platform that were connected during the sea-level changes that occurred is attached to the Horn of Africa. Plate tectonic recon- in the Pleistocene. In contrast, Abd Al Kuri is separated structions indicate that, prior to the rifting of the Gulf of from the other islands of the archipelago and from the Aden, the Socotra shelf and its islands were located adja- Horn of Africa by depths of between 200 and 1000 m, cent to the Dhofar region of Southern Oman, an interpre- which, therefore, exceed Pleistocene sea-level changes tation well-supported by comparisons of the Mesozoic and (Cheung & DeVantier 2006). Tertiary successions in these two areas (Laughton 1966; This complex geological history, which includes a long Laughton et al. 1970; Samuel et al. 1997; Fleitmann et al. period of isolation, together with its topography and its

Fig. 1. Map showing the geographical situation of the Socotra Archipelago.

ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 347 The Trachylepis skinks of the Socotra Archipelago d Sindaco et al. different microclimates form the basis of the high biodi- The genus Trachylepis is one of the four units into which versity and endemism of the Socotra Archipelago. For the genus Mabuya, one of the largest genera of the Scinci- example, 37% of Socotra’s 825 plant species, 90% of its dae family, was split based on molecular and distributional 30 reptile species and 95% of its more than 100 land data (Mausfeld et al. 2002). The other three genera are snail species are not found anywhere else in the world Chioninia (Cape Verde Archipelago), Eutropis (Asian spe- (Sindaco et al. 2009; Van Damme 2009; Razzetti et al. cies) and Mabuya (South and Central America and the 2011). Endemism is especially high at the generic level, Caribbean, except for Trachylepis atlantica from the island with 75% of the land snail genera, 41% of the reptile of Fernando de Noronha, Brazil, and the enigmatic genera and 3.5% of the plant genera being unique to the T. tschudii described from the Peruvian Amazonas; see archipelago. Despite the richness and importance of Miralles et al. 2009a). Although these four genera have Socotra’s biodiversity, little is known about the phylog- been accepted in the latest taxonomic works (Mausfeld & eny, evolution and biogeography of most of the islands’ Schmitz 2003; Miralles et al. 2009a,b; Miralles & Carranza species, and many of the groups, therefore, need taxo- 2010; Miralles et al. 2011), the phylogenetic affinities of a nomic revision (Sindaco et al. 2009; Van Damme 2009; fifth unnamed clade formed by a few species from the Razzetti et al. 2011). A clarification of the systematics and Middle East are still unresolved (Carranza & Arnold 2003; a better understanding of the phylogeographic patterns Miralles & Carranza 2010). According to Mausfeld & and genetic population structure in areas of conservation Schmitz (2003), the Middle East clade is part of Trachyl- concern, such as islands, are therefore crucial to delimit epis. However, the monophyly of Trachylepis has never evolutionarily significant units and design efficient conser- been properly tested; therefore, inclusion of the Middle vation measures. East clade in Trachylepis is still unclear. In the present Reptiles constitute the most relevant vertebrate fauna of study, the members of the Middle East clade are consid- the Socotra Archipelago, with 30 terrestrial species (28 ered Trachylepis, and the term Mabuya sensu lato is used to native and two introduced) and a very high level of ende- refer to the monophyletic assemblage formed by Trachyl- mism at both the specific and generic levels (Ro¨sler & epis, Chioninia, Eutropis and Mabuya. Wranik 2004; Sindaco et al. 2009; Razzetti et al. 2011). The genus Trachylepis currently includes more than 70 Although they have been studied on several occasions species distributed throughout Africa and Madagascar, (Arnold 1986; Scha¨tti & Desvoignes 1999; Ro¨sler & Wra- with a few species being found on other Indian Ocean nik 2004), our understanding of most reptile groups is far islands, in the Middle East and one species, namely from complete. One of the most important outstanding Trachylepis socotrana (previously also known as Mabuya issues is an assessment of the level of intraspecific variabil- socotrana and Euprepis socotranus), being present in the Soco- ity and to understand how and from where the islands tra Archipelago. Owing to the scarcity of morphological fea- were colonized, as well as an assessment of the respective tures of taxonomic value (Greer & Nussbaum 2000), the importance of vicariance and dispersal. To date, only the systematics of some widespread and variable species, and snakes of the genera Hemerophis and Ditypophis have been their phylogenetic relationships, are poorly understood. investigated using appropriate dating methods, with the This is particularly true for T. brevicollis, which is wide- results suggesting that these two groups are relicts that spread in East Africa and Arabia and is considered to be a were probably present on Socotra when it separated from composite species by some authors (i.e. Lanza 1983). mainland Arabia, thus making them examples of vicariance The occurrence of Trachylepis in the Socotra Archipel- (Nagy et al. 2003). Phylogenetic data for other groups ago has been known since the report of Euprepes perrotteti such as Hemidactylus (Carranza & Arnold 2006), Pristurus var. (sic) by Blanford (1881) on the basis of a specimen col- (Papenfuss et al. 2009), Trachylepis (Carranza & Arnold lected by Isaac Bayley Balfour. A year later, it was 2003; Gu¨nther et al. 2005; Whiting et al. 2006), Mesalina described as Euprepes (Euprepis) socotranus by Peters (1882) (Joger & Mayer 2002; Kapli et al. 2008), Hakaria simonyi on the basis of a single specimen collected by Emil (Brandley et al. 2005; Austin et al. 2009) and Chamaeleo Riebeck and Georg Schweinfurth. Since its discovery, monachus (Macey et al. 2008) are much scarcer and in most Trachylepis socotrana has been reported in most papers deal- cases do not allow us to draw any conclusions regarding ing with the herpetofauna of Socotra (see Scha¨tti & the number, direction and time of the different coloniza- Desvoignes 1999; Ro¨sler & Wranik 2004). In contrast to tion events. Moreover, the genetic variability at the intra- the situation on Socotra Island, data concerning the pres- specific level has never been assessed for any of the island ence of T. socotrana on the three smaller islands of the archi- reptile endemics, which may influence or change some of pelago are scarcer. Thus, Trachylepis socotrana was not the previous conclusions reached using partial datasets discovered on Samha Island until February 1999 and on (data not shown). Darsa Island until February 2000 (Ro¨sler & Wranik 2000,

348 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Sindaco et al. d The Trachylepis skinks of the Socotra Archipelago

2004). As pointed out by Scha¨tti & Desvoignes (1999), examined here, such as scale counts, presence or absence T. socotrana was reported only once from Abd Al Kuri by of homologous scale fusions and variability in color pat- Steindachner (1903) on the basis of a specimen deposited terns, are routinely used in taxonomic studies of Scincidae. in the Naturhistorisches Museum of Vienna. A record Scale nomenclature, scale counts and measurements used confirmed almost a century later by Herbert Ro¨sler in in the morphological analyses followed mostly Lanza 1999 (Ro¨sler & Wranik 2004). According to Ro¨sler & (1978) and Gu¨nther et al. (2005). Scale arrangements are Wranik (2000, 2004), ‘there are not obvious morphologi- shown in Fig. 2. Measurements of specimens were cal differences between the specimens [of the different recorded to the nearest 0.1 mm with dial calipers. As most islands], but further work is needed to determine the status museums denied permission to open their specimens, these of these populations’. As all records by Ro¨sler & Wranik could not be sexed. A total of 697 pictures were taken (2004) are indicated as ‘visual observation’, it seems that from all the vouchers included in the morphological these authors did not collect any voucher specimen from analyses and uploaded into Morphobank (http://www. the small islands of the Socotra Archipelago. morphobank.org. Project P461). The results of the Various surveys were organized between 2007 and 2010 meristic, mensural and qualitative characters including ref- in the framework of the ongoing ‘Socotra Conservation erence to the pictures examined for the two specimens of and Development Project’ funded by the Cooperazione Trachylepis from Abd Al Kuri can be found in Table 2, and Italiana and under the auspices of the United Nations the same details for all the specimens of T. brevicollis and Development Project (UNDP) to collect ecological data T. socotrana in Appendix S1. on the reptiles of the Socotra Archipelago to improve the sustainable development and conservation of the Socotra Molecular study Archipelago’s biodiversity. As a result of these surveys Samples, DNA extraction and amplification. A total of 116 (Razzetti et al. 2011), a new Hemidactylus species has specimens were used to infer a comprehensive phyloge- recently been described (Sindaco et al. 2009), and system- netic tree of Mabuya sensu lato using partial sequences of atic revisions for all the endemic reptiles are currently the 12S rRNA and 16S rRNA mitochondrial genes. underway. In the present work, we use both morphological Sequences for 103 of the 116 specimens were downloaded and genetic data to revise the taxonomy of the endemic from GenBank and included the outgroup Egernia whitii, Trachylepis socotrana and to unravel its biogeography and all the available specimens of the Afro-Malagasy genus evolution. Trachylepis (including T. aurata, T. vittata and T. septemtae- niata from the Middle East; Mausfeld & Schmitz 2003) Materials and Methods and a good representation of the Neotropical Mabuya, the Origin of socotran tissue samples and specimens Asian Eutropis and the Cape Verdean Chioninia. The Specimens were collected by hand during three fieldtrips remaining 13 specimens were sequenced for this work and to Socotra Island (December 2007–January 2008, Decem- included two specimens of Trachylepis socotrana from Soco- ber 2008–January 2009, March–April 2010) and a short tra Island and one specimen from each of the other three fieldtrip to the three smaller islands of the archipelago smaller islands of the archipelago (Darsa, Samha and Abd (Darsa, Samha and Abd Al Kuri; see Fig. 1) between the Al Kuri), three T. tessellata and one T. brevicollis from 29th of March 2010 and the 1st of April 2010. Specimens Oman, two T. aurata from Turkey and two T. vittata, one were collected, photographed, measured and set free after from Tunisia and one from Turkey. Specimen data and taking a piece of tail; tissue samples for genetic analyses GenBank accession numbers for all 116 specimens used in were preserved in absolute ethanol. Some voucher speci- the analyses are given in Table 1. DNA was extracted mens were fixed in 95% ethanol and then preserved in using the DNeasy Tissue Kit (Qiagen, Valencia, CA, 75% ethanol; these are held in the collections of the Mu- USA). Primers used for both amplification and sequencing seo Civico di Storia Naturale di Carmagnola, Italy reactions were 12Sa and 12Sb (Kocher et al. 1989) for the (MCCI), and in the Museo di Storia Naturale of the Uni- 12S rRNA (12S) gene and 16Sar-50 and 16Sbr-30 (Palum- versity of Pavia, Italy (MSNPV). bi 1996) for the 16S rRNA (16S) gene. PCR conditions were as in Carranza et al. (2008). Morphological study Specimens from the MCCI and MSNPV, Naturhistoris- Phylogenetic analyses. Geneious v5.3 (Drummond et al. ches Museum, Wien (NMW), the Natural History 2010) was used for contig assembly, visualization of Museum, London (BMNH), and the Museo ‘La Specola’, sequences and as a platform for exporting into different Florence (MZUF), were used for the morphological analy- formats. The 12S and 16S DNA sequences of 115 Mabuya ses. The meristic, mensural and qualitative characters sensu lato specimens and the outgroup Egernia whitii were

ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 349 The Trachylepis skinks of the Socotra Archipelago d Sindaco et al.

A T. socotrana

B T. cristinae sp. nov. C T. cristinae sp. nov. D T. socotrana

EF GH T. “brevicollis” IJKL

Fig. 2. Scale arrangement of members of the Trachylepis brevicollis complex. Scale nomenclature: sn = supranasals; in = internasal (frontonasal); pf = prefrontals; fr = frontal; fp = fronto-parietals; pa = parietals; ip = interparietal; nu = nucals; lo = loreals; sc = supraciliaries; po = preocular; pso = presubocular; poso = postsubocular; pt = pretemporal; uat = upper anterior temporal. Scale arrangement: Prefrontals separated (E, G, J, K); prefrontals in short contact (D, F, H, I, L); prefrontals in broad contact (C). Specimens depicted in the figure: (A, D) – Trachylepis socotrana, Socotra; (B, C) – Trachylepis cristinae sp. nov., Abd Al Kuri I. (holotype); (e–l) – T. ‘brevicollis’ (E, F) – Somalia, Afmendu`, (G, H) – Oman, Mughsayl env., (I–K) – Yemen, Mokah, (L) – Ethiopia, Awash National Park. aligned for each gene independently using the online ver- account the shape of the gamma distribution (G) and the sion of MAFFT v.6 (Katoh et al. 2002) with default number of invariant sites (I) for both 12S and 16S parti- parameters (gap opening penalty = 1.53, gap exten- tions. Model parameters were optimized for each partition sion = 0.0). Phylogenetic analyses were performed using in the subsequent analyses. ML analyses were performed maximum likelihood (ML) and Bayesian analyses. JModel- with RAxML v.7.0.3 (Stamatakis 2006) with 100 random Test (Posada 2008) was used to select the most appropri- addition replicates. Reliability of the ML tree was assessed ate nucleotide substitution model under the Akaike by bootstrap analysis (Felsenstein 1985) including 1000 Information Criterion (Akaike 1973). The model selected replications. Bayesian analyses were performed with was the General Time Reversible (GTR), taking into MrBayes v.3.1.2 (Huelsenbeck & Ronquist 2001). Two

350 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Sindaco et al. d The Trachylepis skinks of the Socotra Archipelago

Table 1 Information on the specimens included in the phylogenetic analyses, with the corresponding Genbank accession number and bibliographic reference

Taxa Locality GenBank Accession Numbers 12S ⁄ 16S Code Fig.X Reference*

C. delalandii 1Sa˜o Thiago Island, Cape Verde AY070344 ⁄ AY070361 P2del [3] C. delalandii 2 Cha˜ das caldeiras, Fogo AF280185 ⁄ AY151482 de45de4F [2] C. fogoensis antaoensis Cha˜ de Lagoa, Santo Anta˜o AF280177 ⁄ AY151480 fo60Mfo5SA [2] C. fogoensis nicolauensis Faro de Barril, Sao˜ Nicolau AF280172 ⁄ AY151481 fo17MfoSN [2] C. spinalis maioensis Morrinho, Maio AF280159 ⁄ AY151478 sp53Msp1M [2] C. spinalis salensis Sal Island, Cape Verde AY070327 ⁄ AY070348 P2spisa [3] C. spinalis spinalis Sa˜o Thiago Island, Cape Verde AY070343 ⁄ AY070360 P2spi [3] C. stangeri Calhau, Sa˜o Vicente AF280167 ⁄ AY151479 st44Mst2SV [2] C. vaillanti Feijoal, Fogo AF280198 ⁄ AY151483 va49Mva6F [2] E. carinata 1 Kodaikanal, India AY070336 ⁄ AY070354 P2car [3] E. carinata 2 Madras, India AY159044 ⁄ AY159073 P3Etcar [5] E. longicaudata 1–⁄ AF153572 P1lon [1] E. longicaudata 2 Phong Nha-Ke Bang, Vietnam AY070341 ⁄ AY070359 P2lon [3] E. macrophthalma Java, Indonesia AY159048 ⁄ AY159077 P3Etophta [5] E. macularia 1 Pakistan AY070335 ⁄ AY070353 P2mac [3] E. macularia 2 Mwe Hauk Village, Myanmar AY159049 ⁄ AY159078 P3Etmac [5] E. madaraszi Gammaduwa, SW Sri Lanka AY159051 ⁄ AY159080 P3Etmad [5] E. multicarinata Luzon, Philippines AY159052 ⁄ AY159081 P3Etmul [5] E. multifasciata 1 Tropical Asia AY151424 ⁄ AY151458 muE111014 [4] E. multifasciata 2 Cardamon Mountains, Cambodia AY151425 ⁄ AY151459 muE111015 [4] E. multifasciata 3 Tropical Asia AY151418 ⁄ AY151452 muE111034 [4] E. multifasciata 4–⁄ AF153576 P1mul [1] E. multifasciata 5 Permuteran, Bali, Indonesia AY159053 ⁄ AY159082 P3Etmultifas [5] E. rudis Bogani Nani Wartabone MP, Sulawesi AB028779 ⁄ AB028790 EtrudP3 [5] E. rugifera 1 Indonesia AY159061 ⁄ AY159090 P3Etrug [5] E. rugifera 2 Bali National Park, Bali, Indonesia AY159064 ⁄ AY159093 P3Etrug383 [5] E. sp. Luzon, Philippines AY159066 ⁄ AY159095 P3EtPHI [5] E. tytleri Mt Harriet, Andaman Is., India AY159045 ⁄ AY159074 P3Ettyt [5] M. agilis 1 Exu, NE Brasil AY151428 ⁄ AY151462 agE11108 [4] M. agilis 2 Prado, Bahia, Brazil AY070326 ⁄ AY070347 P2agi [3] M. cf. bistriata Trinidad, Beni, Bolivia – ⁄ AF153563 P1bis [1] M. dorsivittata 1 Brasilia, Central Brasil AY151426 ⁄ AY151460 doE11106 [4] M. dorsivittata 2 Cunha, Sa˜o Paulo, Brazil AY070346 ⁄ AY070363 P2dor [3] M. frenata Mato Grosso do Sul, SW Brasil AY151427 ⁄ AY151461 frE11107 [4] M. guaporicola Mato Grosso do Sul, SW Brasil AY151434 ⁄ AY151468 guE11101 [4] M. heathi Res. Serra das Almas, Ceara´, Brazil AY070330 ⁄ AY070349 P2hea [3] M. mabouya Tobago AY070339 ⁄ AY070357 P2mab [3] M. macrorhyncha Trancoso, Bahia, Brazil AY070333 ⁄ AY070351 P2macro [3] M. nigropunctata 1 Ruerto Inirida, Colombia AY151438 ⁄ AY151484 niE111016 [4] M. nigropunctata 2 Talparo, Trinidad AY151436 ⁄ AY151470 niE11103 [4] M. nigropunctata 3 AY070334 ⁄ AY070352 P2nig [3] T. acutilabris 10 km N Vis Myn, Namibia – ⁄ AF153560 P1acu [1] T. affinis 1 AF153554 ⁄ AF153561 P1aff [1] T. affinis 2 Jully Hotel, North of Kribi, Cameroon AY159109 ⁄ AY159122 P4Epaff108 [6] T. affinis 3 Benakuma, West of Wum, Cameroon AY159106 ⁄ AY159119 P4Epaff743 [6] T. affinis 4 Benakuma, West of Wum, Cameroon AY159105 ⁄ AY159118 P4Epaff742 [6] T. affinis 5 River Bagwor, Fontem, Cameroon AY159107 ⁄ AY159120 P4Epaff746 [6] T. albilabris 1 Uganda AY070331 ⁄ AY070350 P2alb [3] T. albilabris 2 Kika, East of Moloundou, Cameroon AY159102 ⁄ AY159115 P4Epalb [6] T. atlantica 1 Fernando de Noronha, Brasil AY151429 ⁄ AY151463 atE111020 [4] T. atlantica 2 Fernando de Noronha, Brasil AY151430 ⁄ AY151464 atE111021 [4] T. atlantica 3 Fernando de Noronha, Brazil AY070345 ⁄ AY070362 P2atl [3] T. aurata 1 Bodrum, Turkey JQ598768 ⁄ JQ598781 aJR36 NEW T. aurata 2 Kisehir, Turkey AY151435 ⁄ AY151469 aE111012 [4] T. aurata 3 Isiki, Turkey JQ598769 ⁄ JQ598782 aJR35 NEW T. aurata 4 Vil. Gaziantep, Turkey AY159041 ⁄ AY159070 P3Epaur [5] T. binotata Ruacana waterfall, Namibia – ⁄ AF153562 P1bin [1] T. boettgeri Ambatolampy, Madagascar AY070337 ⁄ AY070355 P2boe [3]

ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 351 The Trachylepis skinks of the Socotra Archipelago d Sindaco et al.

Table 1 (Continued)

Taxa Locality GenBank Accession Numbers 12S ⁄ 16S Code Fig.X Reference*

T. brevicollis 1 Salalah, Oman JQ598770 ⁄ JQ598783 be100361 NEW T. brevicollis 2 Tanzania AY159042 ⁄ AY159071 P3Epbre [5] T. brevicollis 3 Kajado, Kenya AY907712 ⁄ AY907714 NewTraBre [7] T. capensis 1 Kouga Mts. E Cape, South Africa AF280139 ⁄ AY151475 cE111015 [4] T. capensis 2 Windhoek, Namibia – ⁄ AF153564 P1cap [1] T. cf. affinis 1 Benakuma, West of Wum, Cameroon AY159104 ⁄ AY159117 P4Epaffcf744 [6] T. cf. affinis 2 Ngoulemakong, NE of Ebolowa, Cameroon AY159103 ⁄ AY159116 P4Epaffcf745 [6] T. cf. dumasi Kirindy, Madagascar AF153555 ⁄ AF153566 P1dum [1] T. cf. perrotetii Tchabal Mbabo, Cameroon AY159099 ⁄ AY159113 P4Eppercf [6] T. comorensis Nosy Tanikely, Madagascar – ⁄ AF153565 P1com [1] T. cristinae sp.n. Abd Al Kuri, Socotra Archipelago, Yemen JQ598771 ⁄ JQ598784 JR15 NEW T. dichroma 1 (Tanzania, presumed origin of parents) AY907711 ⁄ AY907713 NewTraSP [7] T. dichroma 2 Dodoma, Tanzania AY159043 ⁄ AY159072 P3EpSP [5] T. elegans 1 Kirindy, Madagascar (western) – ⁄ AF153568 P1eleW [1] T. elegans 2 Antalaha, Madagascar (eastern) AF153556 ⁄ AF153567 P1eleE [1] T. gravenhorstii Antananarivo-Mandraka, Madagascar AY070340 ⁄ AY070358 P2gra [3] T. hoeschi Swakop, Namibia – ⁄ AF153569 P1hoe [1] T. homolocephala Rine’s Nature Reserve, South Africa – ⁄ AF153570 P1hom [1] T. irregularis Mt. Elgon, Uganda – ⁄ AF153571 P1ire [1] T. maculilabris 1 Momane Swamp, Mozambique JQ598772 ⁄ JQ598785 mE1110110 NEW T. maculilabris 2 Benakuma, West of Wum, Cameroon AY159111 ⁄ AY159124 P4Epmaculi00 [6] T. maculilabris 3 Afan, NE of Ma’an, Cameroon AY159110 ⁄ AY159123 P3Epmaculi99 [6] T. maculilabris 4–⁄ AF153574 P1maculi [1] T. maculilabris 5 Amani, Usambara Mt., Tanzania AY070338 ⁄ AY070356 P2maculi [3] T. maculilabris casuarinae Fogo Island, Mozambique AF280138 ⁄ AY151474 mE111019 [4] T. margaritifera 1 Malema, Mozambique AF280136 ⁄ AY151473 fE1110113 [4] T. margaritifera 2 (Tanzania) – ⁄ AF153575 P1mar [1] T. occidentalis Aus, Namibia – ⁄ AF153577 P1occ [1] T. perrotetii 1 Gambia – ⁄ AF153578 P1per [1] T. perrotetii 2 Crossroads Hina-Moufou, Cameroon AY159100 ⁄ AY159114 P4Epper [6] T. perrotetii 3 Ghana, W. Africa AY151417 ⁄ AY151451 peE111017 [4] T. quinquetaeniata AF153558 ⁄ AF153579 P1qui [1] T. septemtaeniata Syria AY159062 ⁄ AY159091 P3Epsep [5] T. socotrana 1 Darsa Island, Socotra Archipelago, Yemen JQ598773 ⁄ JQ598786 JR12 NEW T. socotrana 2 Samha Island, Socotra Archipelago, Yemen JQ598774 ⁄ JQ598787 JR9 NEW T. socotrana 3 Socotra Island, Yemen JQ598775 ⁄ JQ598786 JR1 NEW T. socotrana 4 Socotra Island, Yemen AF280140 ⁄ AY151476 sE1110132 [4] T. socotrana 5 Socotra Island, Yemen AY159063 ⁄ AY159092 P3Epsoc [5] T. sp. n. Hossere Ngang-Ha, Cameroon AY159108 ⁄ AY159121 P4EpSP [6] T. spilogaster Aranos, Namibia – ⁄ AF153580 P1spi [1] T. striata striata Mtunzini, KwaZulu-Natal, South Africa – ⁄ AF153581 P1str [1] T. striata wahlbergi Grootfontein, Namibia – ⁄ AF153582 P1strw [1] T. sulcata 1 Ongongo waterfall, Kaokoland, Namibia – ⁄ AF153583 P1sul [1] T. sulcata 2 Kamanjab, Namibia AY151420 ⁄ AY151454 suE111037 [4] T. tessellata 1 Mughsayl, Oman JQ598776 ⁄ JQ598788 te100363 NEW T. tessellata 2 Wadi Haql, Oman JQ598777 ⁄ JQ598789 tJR38 NEW T. tessellata 3 Nazwah, Oman JQ598778 ⁄ JQ598789 te100365 NEW T. varia Waterberg, Namibia – ⁄ AF153584 P1varia [1] T. variegata Otjimbingwe, Namibia – ⁄ AF153585 P1var [1] T. vato Mt. Ibity, Madagascar AY159068 ⁄ AY159097 P3Epvat [5] T. vittata 1 Tozeur, Tunisia JQ598779 ⁄ JQ598790 vE1110138 NEW T. vittata 2 Sambayat, Turkey JQ598780 ⁄ JQ598791 vE1110142 NEW T. vittata 3 Frikeh, SE Jisir, Syria AY159069 ⁄ AY159098 P3Epvit [5] T. wrightii Fregate Island, Seychelles AF280124 ⁄ AY151472 wrMawr1 [2] Egernia whitii Tasmania AF280119 ⁄ AY151447 Ewhitii [4]

*References: [1] Mausfeld et al. 2000, [2] Carranza et al. 2001, [3] Mausfeld et al. 2002, [4] Carranza & Arnold 2003, [5] Mausfeld & Schmitz 2003, [6] Mausfeld et al. 2004, [7] Guenther et al. 2005

352 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Sindaco et al. d The Trachylepis skinks of the Socotra Archipelago independent runs of 5 · 106 generations were carried out, for Chalcides could be applied to the inferred phylogeny sampling at intervals of 1000 generations, producing 5000 of Mabuya sensu lato. trees. Convergence and appropriate sampling were con- To calculate the substitution rates of the 12S and 16S firmed by examining the standard deviation of the split gene fragments of Chalcides, a dataset was assembled com- frequencies between the two simultaneous runs and the prising 33 Chalcides and two Eumeces specimens that were Potential Scale Reduction Factor (PSRF) diagnostic. After used as outgroup, and all three partitions (12S, 16S, cytb) discarding the first 500 trees of each run, a majority-rule available from Carranza et al. 2008 (see Appendix S2; consensus tree was generated from the remaining trees. Morphobank Doc3a: 12S rRNA dataset, Doc3b: 16S Topological constraints to test alternative topolo- rRNA dataset, and Doc3c: cytochorome b dataset). First, gies were compared with optimal topologies using the it was tested whether a strict molecular clock or a relaxed approximately unbiased (AU) (Shimodaira 2002) and Shi- one best fit the data, performing an ML ratio test as modaira–Hasegawa (Shimodaira & Hasegawa 1999) tests implemented in MEGA5.0 (Tamura et al. 2011) for each implemented in CONSEL (Shimodaira & Hasegawa 2001). one of the partitions. The hypothesis that the sequences evolve in a clock-like way could not be rejected for the Estimation of divergence times. The progress achieved over 16S and cytb datasets, while it was rejected for the 12S the past years in the development of methods for the esti- dataset (likelihood ratio test statistic ()2logK) = 39.86, 2 mation of lineage divergence times has brought by an 37.76 and 67.03, respectively, which approximates to a v31 ongoing discussion over the performance and accuracy of distribution under the null hypothesis; P = 0.1322, 0.1876 such analyses and the possible pitfalls (see Drummond and < 0.001). Therefore, two independent analyses with et al. 2006; Ho & Philips 2009; Graur & Martin 2004; the three partitions were performed using the software amongst others), while evaluation of the new methodolo- BEAST v.1.6.1 (Drummond & Rambaut 2007) with the gies shows promising results (Battistuzzi et al. 2010). following model and prior specifications (otherwise, by In this study, the lack of internal calibration points in default): GTR+I+G, Relaxed Uncorrelated Lognormal Mabuya sensu lato precluded the direct estimation of the Clock (estimate) for the 12S partition and Strict Clock for time of the cladogenetic events in the phylogeny. Alterna- the 16S and cytb partitions, Yule process of speciation, tively, the substitution rate of the same mitochondrial random starting tree, alpha Uniform (0, 10), yule.birth- region calculated for another scincid group could be used Rate (0, 1000), nucleotide substitution rates Uniform (0, for this purpose. A very well-studied and documented cal- 100) initial value = 1. A single calibration point was ibration point exists for the skinks of the genus Chalcides applied, as mentioned earlier, with a uniform distribution (see Brown & Pestano 1998; Carranza et al. 2008; Brown (lower 1.0, upper 1.2) (for more information on the cali- & Yang 2010): the age of the colonization of the island bration of Chalcides, see Brown & Yang 2010). Two indi- of El Hierro from La Gomera by Chalcides coeruleopuncta- vidual runs were performed for 3 · 107 generations and tus, presumably soon after its appearance (1.0–1.2 MY). Tracer v.1.4 (Rambaut & Drummond 2007) was used to To test whether the Chalcides rate could be applied to the confirm that stationarity was reached, and sufficient effec- Mabuya sensu lato phylogeny, a relative rate test was per- tive sampling size was achieved. The substitution rate for formed to check whether there were statistically signifi- 12S and 16S gene fragments as indicated by the ‘mean- cant differences in the substitution rates of both 12S and Rate’ posterior was 0.01305 substitutions per lineage per 16S mitochondrial fragments between the two lineages site and 0.008 substitutions per lineage per site, respec- compared. For this test, a dataset with 150 skinks was tively. These substitution rates were applied to the Mabuya assembled including 33 Chalcides specimens, with repre- sensu lato dataset to estimate the divergence times within sentatives of all its main clades (Carranza et al. 2008), all the phylogeny. An ML ratio test was also performed for 115 specimens of Mabuya sensu lato used to infer its phy- the 12S and 16S partitions of this dataset, and the ‘strict logenetic relationships and two outgroups, Cordylus war- clock’ was rejected for both (likelihood ratio test statistic reni and Gerrhosaurus validus (see Table 1; Appendix S2; ()2logK) = 447.69 and 386.65, respectively, which approx- 2 Morphobank Doc2a: 12S rRNA dataset and Doc2b: 16S imates to a v114 distribution under the null hypothesis; rRNA dataset). The software RRT (Robinson-Rechavi & P < 0.001 and < 0.001). 12S and 16S rRNA partitions Huchon 2000) was implemented and results were highly were analyzed jointly using BEAST v.1.6.1 (Drummond & non-significant in both cases (P  0.95, 0.74, respec- Rambaut 2007), applying the previously calculated rates tively), indicating that the two groups tested (Chalcides for each gene partition. Models and prior specifications and Mabuya sensu lato) do not differ significantly in the applied were as follows (otherwise, by default): GTR+G+I, evolutionary rates of the 12S and 16S mitochondrial Relaxed Uncorrelated Lognormal Clock (rates 0.01305 genes. As a result of that, the substitution rate calculated and 0.008, respectively), Yule process of speciation, ran-

ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 353 The Trachylepis skinks of the Socotra Archipelago d Sindaco et al. dom starting tree, alpha Uniform (0,10), yule.birthRate there were two independent colonizations of the Socotra (0,1000), nucleotide substitution rates Uniform (0, 100) Archipelago by representatives of the genus Trachylepis.A initial value = 1. Two individual runs were performed for first colonization to Socotra, Samha or Darsa Islands 3 · 107 generations, and the ultrametric tree (Appendix S3) occurred between 7.3 and 13.8 (mean 10.5) million years was inferred after discarding the first 3 · 106 trees of each. ago (mya), and a second, more recent, independent coloni- zation to Abd Al Kuri Island between 1.5 and 4.5 (2.9) Results mya. The uncorrected genetic distances amongst T. socotr- Molecular study ana from Socotra, Darsa and Samha Islands, the specimen The alignment of the Mabuya sensu lato dataset included from Abd Al Kuri Island, T. brevicollis and T. dichroma are a total of 904 base pairs (bp), of which 392 corresponded shown in Appendix S5 and support the genetic distinctive- to the 12S and 512 to the 16S. The alignment matrix is ness of the Trachylepis lineage from Abd Al Kuri. available from Morphobank (Doc1), and the codes used According to our results, T. brevicollis is also polyphy- in the matrix can be deciphered with Table 1. Of the letic, with T. brevicollis from Tanzania and Kenya being total of 392 bp of the aligned 12S sequences, 294 were more closely related to T. dichroma than to the single variable and 169 parsimony informative, while the respec- specimen of T. brevicollis from Oman included in our tive sites for the 512-bp-long 16S fragment were 202 and analyses. To test this topology, a third constraint analy- 195. sis was performed in which T. brevicollis (Fig. 3) was The results of the ML and Bayesian analyses were forced monophyletic. In this case, neither the AU nor almost totally congruent (Appendix S4), and therefore, the SH tests could reject the alternative hypothesis of only the ML tree with the posterior probability values of monophyly of T. brevicollis (AU test: P = 0.379, SH test: the Bayesian analysis is shown in Fig. 3. The genera Ma- P = 0.695). buya, Eutropis and Chioninia appear as independent well- supported clades in the phylogeny, while the genus Morphological study Trachylepis is polyphyletic, with the clade formed by The results of the molecular analyses presented earlier and T. vittata, T. septemtaeniata and T. aurata branching inde- the morphological comparisons support the distinctiveness pendently from all the remaining representatives of of the Trachylepis population from Abd Al Kuri Island. As Trachylepis. To check whether our dataset rejected the a result of that, hereafter, we describe this population as a monophyly of Trachylepis, a constrained tree in which different species. Trachylepis was forced monophyletic was compared with the best ML tree presented in Fig. 3. The results of the Trachylepis cristinae sp. nov. constraint analysis differed between the two tests, and Holotype. Male, YEMEN, Abd Al Kuri Island, Khaysat while the alternative hypothesis was marginally rejected by Salih village env. at about 1211¢N–5214¢E, m 36 a.s.l., the AU test (P = 0.048), the monophyly of Trachylepis was 1.IV.2010, Cristina Grieco, Fabio Pupin, Elisa Riservato not rejected by the more conservative SH test (P = 0.236). and Roberto Sindaco leg. (MCCI-R1585). Trachylepis specimens from the Socotra Archipelago do not form a monophyletic group, making T. socotrana poly- phyletic (see Fig. 3). According to our phylogeny, the Paratype. Female, YEMEN, Abd Al Kuri Island, Oskar specimens from Socotra, Samha and Darsa Islands form a Simony leg. Labelled: ‘Mabuia socotrana (Pet.) Boul ⁄ clade and are very closely related, but the single specimen Abd - el - Kuri. 3f. Su¨dar. Exp. ⁄ 1898 – 22 ⁄ 1 899 – from Abd Al Kuri branches within a well-supported clade Simony’ [= Mabuia socotrana (Peters) Boulenger ⁄ Abd together with the recently described T. dichroma Gu¨nther, el-Kuri Island. 3rd South-Arabian Expedition ⁄ 1898– Whiting and Bauer, 2005 from Tanzania and with all the 22.I.1899 – Oskar Simony leg.]. (NMW 9396). Arabian and African representatives of T. brevicollis. In fact, Mabuya socotrana (Pet.) Blgr. – Steindachner, 1903: 13 the Trachylepis from Abd Al Kuri seems more closely (partim): ‘Abd el-Kuri’. related to Omani T. brevicollis, although with almost no Mabuya soqotranus – Wranik, 1998a: table 1 (partim): support in the ML and Bayesian analyses. A constrained ‘Abd el-Kuri’. tree in which T. socotrana was forced monophyletic was Mabuya socotrana – Wranik, 1998a: 150, table 3 (par- significantly different from the best ML tree presented in tim): ‘Abd el-Kuri’. Fig. 3 by two topology tests (AU test: P = 0.002, SH test: Mabuya socotrana – Wranik, 1998b: 171, fig. 12, table P = 0.013), indicating that our dataset rejects the mono- 1(partim): ‘Abd el-Kuri’. phyly of T. socotrana. According to the phylogenetic tree Mabuya socotrana – Scha¨tti & Desvoignes 1999: 121– presented in Fig. 3 and the inferred divergence times, 122 (partim): ‘Abd el-Kuri’.

354 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Sindaco et al. d The Trachylepis skinks of the Socotra Archipelago

Fig. 3. ML tree of the Mabuya sensu lato inferred using the concatenated 12S and 16S mtDNA gene fragments. Numbers by the nodes indicate bootstrap support for ML analyses, followed by an asterisk indicating high posterior probability for BI analyses. The symbol ‘<’ is used to show bootstrap support or posterior probability lower than 50% and 95%, respectively. Age ranges for some selected nodes are indicated with an arrow with the mean age in brackets. The tree was rooted using Egernia whitii. Information on the samples included is shown in Table 1.

ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 355 The Trachylepis skinks of the Socotra Archipelago d Sindaco et al.

Mabuya socotrana –Ro¨sler & Wranik 2004: 524 (par- Table 2 Measurements and scale counts of the type specimens of tim): ‘Abd Al Kuri Island: 1212¢N5215¢E, and slope Trachylepis cristinae, sp. nov. (all measurements in mm) Jabal Salih, 1210¢N5215¢E, 100–450 m, 17- 19.II.1999¢. MCCI-R1585 NMW-9396 Sex M F

Etymology. The species epithet is a genitive Latin noun SVL 114 110 to honour Cristina Grieco, who found the holotype in TotL 220 175 Abd Al Kuri. HL 20.6 20.6 HW 19.5 18.2 Ratio HL ⁄ HW 1.05 1.13 Diagnosis. A member of the Trachylepis brevicollis complex EST 12.0 10.6 characterized by high number of scale rows at midtrunk EED 11.3 10.0 (38), palmar and tarsal scales never pointed nor spinose, MSR 38 38 subdigital lamellae smooth or only weakly keeled; color VGS 65 62 characteristic (see description); SVL of adults up to SL l ⁄ r7⁄ 78⁄ 8 LF4 l ⁄ r17⁄ 16 17 ⁄ 17 114 mm. LT4 l ⁄ r22⁄ 25 22 ⁄ 23 Dorsal keels 2–3 2–3 Description of the holotype specimen. (Fig. 2b–c; Appendix S6 Supranasals arrangement (C) (C) and S7 a–c; Table 2; Morphobank pictures M85683– Prefrontals arrangement C C M85688). Measurements and scale counts are presented in Supraocul. ent. frontal I–II I–II ⁄ I–II-(III) Table 2. Adult specimen (a male) with robust and slightly Tarsal scales B B Palmar scales B B depressed body; snout-vent length = 114 mm. Head Foot lamellae S S equally longer than broad (ratio 1.05), slightly depressed, Hand lamellae S WK relatively pointed, slightly broader than the neck region at Morphobank images M85683–M85688 M85867–M85872 the level of the ear openings. Lower eyelid with a distinct SVL, snout-vent length; TotL, total length; HL1, head length from tip of snout to transparent oval window. Ear openings not covered by posterior border of ear opening; HL, head length from tip of snout to posterior enlarged scales, the left vertically oval and the right a verti- border of parietals; HW, head width at level of ears; EST, distance between anterior cal slit (an abnormality). Snout slightly convex in profile, margin of eye to tip of snout; EED, distance between posterior margin of eye to with a relatively large nostril sunk into the rear edge of the anterior margin of ear; MSR, number of midbody scale rows; SC, number of nasal; a small postnasal, separated from the nostril by the subcaudals; SL, number of supralabials; LF4 l ⁄ r, number of subdigital lamellae under 4th finger (left ⁄ right); LT4 l ⁄ r, number of subdigital lamellae under 4th toe rim of the nasal. Supranasals with a short median contact (left ⁄ right); Dorsal keels: number of dorsal keels; Supranasals arrangement: suture. Internasal (frontonasal) distinctly pointed ante- arrangement of supranasals; S, separated; (C), in short contact; C, in broad contact; riorly. Prefrontals pentagonal, distinctly pointed poste- Prefrontal arrangement: arrangement of prefrontals; S, separated; (C), in short riorly, with a median contact suture long 1 ⁄ 2 of their length. contact; C, in broad contact; Supraocul. ent. frontal: supraoculars entering the Two undivided loreals, of which the larger, posterior one, frontal scale; Tarsal scales: shape of tarsal scales (foot); F, flat; G, granular; B, blunt tubercles; P, pointed tubercles; S, spinose (mucronated) tubercles; (S), slightly borders the first loreal, the prefrontal, two supralabials (3rd spinose; Palmar scales: shape of palmar scales (hand); F, flat; G, granular; P, and 4th), the first supraciliar, and a preocular; the latter is pointed tubercles; S, spinose (mucronated) tubercles; (S), slightly spinose; Foot followed by a single pre-subocular. Supraciliaries five, the lamellae: shape of lamellae under the 4th toe; Hand lamellae: number of lamellae second partly coalescent with the first supraocular. Pretem- under the 4th finger. Morphobank images: Morphobank codes (project P461) for poral two, both contacted by parietal. Sub-postocular two, the pictures of the T. cristinae specimens. upper contacting lower pretemporal. Frontal nearly as long as the distance between its anterior tip and tip of snout, Miralles 2006: 5). Occipital scale indistinct. Eight suprala- laterally in contact with two supraoculars (1st and 2nd). bials on each side, with the 6th distinctly broader than the Frontoparietal shields in contact between them and with others and in subocular position. Seven infralabials on each 2nd, 3rd (the left also with the 4th) supraoculars. Parietal side (the 1st on the right side longitudinally divided). The shields completely separated by a very elongated interparie- postmental borders seven scales, and almost touches the tal shield, without visible parietal eye. Primitive condition 3rd right infralabial. of overlap pattern between parietal scale and upper anterior Fore and hind limbs clearly overlap when adpressed temporal scales (according to Greer & Nussbaum 2000), against the body; each extremity with five fingers and five that is, the parietal overlaps the upper anterior temporal. toes; fore arms length 33.8 mm; hind legs length One pair of almost smooth nuchal shields (the right almost 48.0 mm. Relative length of fingers 3  4>2>5>1, relative divided), each with three rows of cycloid scales homologue length of toes 4>3>5>2>1; 4th finger length (from inser- to nuchal ones; any secondary nuchals present (sensu tion of 3rd finger, claw included) = 9.5 mm; fourth toe

356 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Sindaco et al. d The Trachylepis skinks of the Socotra Archipelago length (from insertion of 5th toe, claw inclu- short contact with the 3rd (left). Interparietal with visible ded) = 14.5 mm; subdigital lamellae under the fourth toe parietal eye. Nuchals undivided. The postmental plate con- 22 ⁄ 25 (left ⁄ right), flat and smooth. Subdigital lamellae un- tacts in a point the third left infralabial. Subdigital lamellae keeled. Palmar scales blunt, tarsal scales are blunt or weakly keeled. Pattern and color in this one century old pre- obtuse. Raised tubercles never pointed. served specimen are similar to the type, but slightly faded; Dorsal and ventral scales are clearly imbricating. the belly is whitish. Thirty-eight rows of scales at midbody. Sixty-five smooth gulars + ventrals from postmentals to vent. Scales at the Comparison with other species. For comparison with back and upper flanks each bearing two or three distinct Trachylepis socotrana and T. brevicollis see Fig. 2e-l; Appen- keels. Scales and shields of the head, scales of the under- dix S2; and Morphobank project P461. Although not clo- surfaces of head, belly, most of tail (except proximal dorsal sely related to it, in Appendix S8, we list the main tail scales) as well as those of the lower flanks unkeeled. differences between T. cristinae sp. nov. and other Arabian, The scales of the dorsum of the fore and hind limbs are East and Southern African species of Trachylepis. Informa- weakly bi- or tricarinate, or almost smooth. tion on the morphology of these species was obtained Tail with a 81-mm-long regenerated tip with total from Lanza & Carfi (1968); Largen & Spawls (2006); length (106 mm) slightly shorter than SVL, its basal part Arnold (1986); Spawls et al. (2002); Broadley & Howell rectangular (height = 12 mm, width = 18 mm), and the (1991); Gu¨nther et al. (2005). distal half clearly laterally compressed. Trachylepis socotrana, the only other Trachylepis of the Socotra Archipelago, is easily distinguishable from T. cristinae Coloration in life. Dorsal parts brownish with indistinct by its smaller size (SVL up to 97 mm, but usually less black spots, attributed to the dark ground color of the in adult individuals, with a mean of 84.4 ± 7.6; N = 20; scales, blackish with light brown along the posterior mar- Appendix S1 and unpublished field data), by having gin. Upper parts of the head brownish, with blackish prefrontals separated or in short contact (largely in contact nuances, particularly along sutures of the head plates and in T. cristinae), frontal scale entering the supraoculars the tip of the snout. Upper parts of the head brownish, (I)-II-III (only the supraoculars I and II in T. cristinae), with margins of the plates irregularly edged with black, internasal (frontonasal) scale pointed posteriorly (truncated and tip of the snouth (rostral, nasals and supranasals) posteriorly); frontoparietals in contact with the supraocu- black. Sides of the head brownish, with temporals, post- lars III and IV [in contact with supraoculars II-III-(IV) in suboculars and posterior supralabials mostly black, as well T. cristinae] (Fig. 2); most dorsal scales with 3–5 keels (2 as the neck (roughly under a line connecting the ear ope- or 3 in T. cristinae), 30–34 scales around midtrunk (data ning with the eye). Underparts of the head bicolor: black from Appendix S2 and from Boulenger 1903: 85) (38 in anteriorly (chin, infralabials and anterior gular shields) and T. cristinae). Moreover, T. socotrana has a completely different on the sides, reddish (with black spots) posteriorly. Flanks, color pattern, usually uniformly brownish-grey; some adults neck and sides of the original tail with rather distinct have orange head, and some specimens bear scattered dark irregular black vertical bands (13–14 between axilla and spots on the throat. Some specimens from Darsa Island groin), 2–3 scales wide, sometimes fused, alternating with have a striped pattern of five longitudinal blackish stripes narrow indistinct whitish-orange bands; the black bands (one vertebral, two dorsolateral and two lateral), alterna- are more irregular on the sides of the neck. Belly, under- ting with two beige broader paravertebral ones and two parts of limbs, and ventral side of the original tail reddish; evident withish dorsolateral ones (MCCI-R1583). regenerated tail uniformly blackish. Iris pink-beige, with Trachylepis cristinae sp. nov. differs from members of the black mottling; pupil black, highly irregularly shaped. T. brevicollis complex in the following characteristics: 38 Weight in life: 39.5 g. (paratype) (Appendix S7 d–f; scales at midtrunk (between axilla and groin), versus 30–37 Table 2; Morphobank: M85867-M85872). Variation in sca- (usually 32–34; mean 33.4 ± 1.71, N = 33) recorded in the lation and body measurements of the NMW specimen is Arabian populations of T. brevicollis (Tornier 1905: 386; reported in Table 2; it differs from the type specimen by the Anderson 1895: 648; Fritz & Schu¨tte 1988: 44–49; original following variations. Relative length of fingers 4>3>2>5>1. data) and 29–35 (usually 30–33, depending on populations) Tail with a 65-mm-long regenerated tip, its distal half not in Africa (Lanza & Carfi 1968: 214-216, Fritz & Schu¨tte clearly compressed. Ratio between length and width of the 1988: 44–49, original data). The subdigital lamellae head = 1.37. Ear opening oval, about 2.5 times as high as under the toes, which are smooth or only weakly keeled in wide, with one small anterior rounded lobule. The first lor- T. cristinae but always keeled in both Arabian and African eal on the right side is divided vertically. Frontal laterally in specimens, seem to be a good character. The number of contact with 1st and 2nd supraoculars (right side) and in subdigital lamellae is higher in the two known specimens

ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 357 The Trachylepis skinks of the Socotra Archipelago d Sindaco et al. of T. cristinae (4th finger = 16–17; 4th toe = 22–25) and in Discussion Arabian T. brevicollis (4th finger = 13–17, As first suggested by Mausfeld & Schmitz (2003) and mean = 15.24 ± 1.07; 4th toe = 16–23, mean = 19.71 ± Carranza & Arnold (2003) and confirmed by the results 1.55, N = 34) than in African T. brevicollis (4th fin- presented here (see Fig. 3), the phylogeny of Mabuya sensu ger = 12–15, mean = 13.54 ± 1,14; 4th toe = 15–19, mean lato includes five well-differentiated lineages that are clas- = 17.11 ± 1,05; N = 27). The appearance of the palmar sified into four different genera. While Mausfeld & Sch- and tarsal scales, which are smooth in Abd Al Kuri speci- mitz (2003) proposed a monophyletic Trachylepis, which mens, is diagnostic only with respect to African specimens would include both the Afro-Malagasy as well as the Mid- (spinose in almost all African specimens examined; N = 27), dle East species, they also already suggested that the Mid- whereas they are smooth in most of the Arabian specimens dle East species may still represent a distinct radiation and studied (in 100% of specimens from Oman and the Hadra- thus would require a separate genus to be erected. maut; N = 9). As in most Arabian specimens (65%; N = 34), Although Mausfeld & Schmitz (2003) recovered the the supranasals are in close contact in T. cristinae, whereas in monophyly of Trachylepis in their ML tree (Fig 2, Maus- African specimens they are mostly separated (40.1%), in feld & Schmitz 2003), the bootstrap support for this contact over only a small portion of the scale (22.2%) or, assemblage was very low (54%), and the tree only included more rarely, in close contact (37%) in the specimens studied two of four genera of Mabuya sensu lato (Eutropis and (N = 27). Furthermore, the prefrontal scales are in close Trachylepis) and a very limited taxon sampling within contact in T. cristinae, whereas they are usually separated Trachylepis (three species from the Middle East clade and (52.9%) or have a narrow suture (20.6%) in Arabian T. brevicollis seven from the Afro-Malagasy clade). (N = 34); similar frequencies also occur in African spe- The phylogenetic tree presented in Fig. 3 has been cimens, where the prefrontals are separated in 48.1%, in inferred with the most complete dataset of Mabuya sensu slight contact in 25.9% and in close contact in 25.1% of the lato assembled to date (Morphobank Doc1), including a specimens examined (N = 27). Other head scalation charac- complete taxon sampling at the generic level and all the teristics (i.e. supraoculars entering the frontal scale), the representatives of Trachylepis from GenBank for which the shape and size of the ear opening and the number and shape 12S and 16S mtDNA regions were available (see Table 1). of the ear opening lobules, appear to be very variable Although the results suggest that Trachylepis is polyphy- amongst individuals and populations. letic (Fig. 3), the constraint analyses (see results section) The coloration pattern seems to be very distinct as all do not rule out a monophyletic Trachylepis including both African T. brevicollis specimens examined are characterized the Middle East and Afro-Malagasy clades. Pending fur- by a light color, with or without dark blotches, ocelli on ther data, we also preliminary consider the Middle East the back, or white spots on the head and trunk. The dis- species to be part of the Trachylepis radiation. It seems tinctive lateral dark sidebands found in T. cristinae sp. nov. clear from Fig. 3 that the two mtDNA regions used in the have also been observed in two specimens from Gardo, present analyses cannot resolve the phylogenetic relation- Somalia (MZUF 10138, 10154), although all NE African ships at the base of the Mabuya sensu lato tree, thus mean- specimens studied lack the extensive dark coloration on ing that until more data, including nuclear genes, are the head and throat. Arabian specimens also exhibit con- added, it will not be possible to satisfactorily resolve the siderable variation. Thus, according to Arnold (1980: 313) taxonomy and the deep level evolutionary relationships of ‘even in Arabia there are differences from place to place as this complex. well as individual variation; in Dhofar a broad, light, dor- The present study shows that the Socotra Archipelago solateral band is present and females have a series of dark was independently colonized by members of the Trachylepis blotches on the dorsum; males are uniform but have black- brevicollis complex twice (see Fig. 3) and that these pro- ish heads with small light spots on the body’, and in some cesses gave rise to two independent species: T. socotrana, areas, animals are often very dark. The specimens exam- which inhabits the islands of Socotra, Darsa and Samha, ined often show whitish spots, dark blotches and ocelli, and a second species, T. cristinae sp. nov., which is ende- and their throats are light, more or less distinctly striped mic to Abd Al Kuri. According to our phylogenetic analy- longitudinally or even mottled. ses, these two colonization events took place at different The head shape of T. cristinae is also characteristic: the times, approximately 10.5 (7.3–13.8) mya for T. socotrana, snout is longer and more slender than in T. brevicollis in and 2.9 (1.5–4.5) mya for T. cristinae. These events are both sexes when viewed from above, with the sides slightly posterior to the late Oligocene ⁄ early Miocene, the time concave between the eye and nostril; this feature is even that has been suggested for the onset of the drifting of more evident in the jaw when observed from below the Socotra Archipelago from its original position in the (Appendix S7 c, f). Dhofar region, Southern Oman (see introduction).

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The geological origin of the archipelago, together with rubri’ [=Ashik Island, Red Sea, Saudi Arabia]), Tiliqua bur- the calibrated phylogenetic tree presented in Fig. 3, there- toni Blyth, 1856 (type locality: ‘Somali Country’), Mabuya fore suggest that, contrary to other Socotran reptiles chanleri Stejneger, 1893 (type locality: ‘Tana River, East (Nagy et al. 2003; unpublished data), the origin of the two Africa’ [now Kenya]), Mabuia (sic) Rotschildi Mocquard, Trachylepis species is not the result of vicariance. According 1905 (type locality: ‘Endessa (Abyssinie)’), Mabuya pulchra to several authors (Laughton 1966; Laughton et al. 1970; Matschie, 1893 (type locality: ‘Scadi prope Lahadsch’ Samuel et al. 1997; Fleitmann et al. 2004; Bosworth et al. [=‘Scadi’, close to Lahej, north of Aden]), Mabuya somalica 2005; Autin et al. 2010; amongst others), by the time of Calabresi, 1915 (type locality ‘Barde`ra [and] Gorie`i’, the inferred colonization events of the Socotra Archipelago Somalia). As stated by Scha¨tti & Gasperetti (1994): ‘the by Trachylepis, the islands had already drifted close to their uncertainty surrounding these names cannot be elucidated actual position in the Arabian Sea, thus meaning that the without a study of the type material; finally, it cannot be two colonization events must have been the result of dis- ruled out that the type locality of M. brevicollis (‘Abys- persal. Long-distance transmarine colonization events are sinia’) might be in error’. Owing to the morphological not rare in reptiles (Carranza et al. 2000; see de Queiroz plasticity of members of the T. brevicollis complex, a 2005 for a review), and, together with some gecko groups, genetic analysis of specimens coming from the type locali- the skinks of the Mabuya sensu lato complex are amongst ties of all of the described taxa would appear to be neces- the best dispersers. For instance, they have naturally sary to solve the systematics of the group and draw species crossed the Atlantic Ocean on two occasions (Mausfeld boundaries. et al. 2002; Carranza & Arnold 2003; Miralles & Carranza With the description of T. cristinae sp. nov., all five spe- 2010) and colonized multiple archipelagoes across their cies of terrestrial reptiles present on Abd Al Kuri can now large distribution range, including the Cape Verde islands be considered to be endemic, thus highlighting the impor- (Carranza et al. 2001; Miralles et al. 2011) and the islands tance of this small island from a biodiversity point of view. of the Gulf of Guinea (Jesus et al. 2005a,b). Although Abd Al Kuri lies just 66 km to the west of Samha Although the island of Abd Al Kuri is closer to mainland and 105 km from Socotra (Fig. 1), our analyses clearly show Africa than to Arabia, Trachylepis cristinae seems to be phy- that the ancestor of T. cristinae is more closely related to logenetically and morphologically more closely related to T. brevicollis from Southern Arabia (more than 350 km to T. brevicollis from Oman than to any African Trachylepis the north) than to T. socotrana from these neighbouring (Fig. 3). In fact, the two T. brevicollis specimens from Africa islands. This biogeographical pattern is also found in other included in our analyses appear to be more closely related groups, especially Mesalina, with M. kuri from Abd Al Kuri to the recently described species T. dichroma from Tanzania being more closely related to mainland Arabian Mesalina (Gu¨nther et al. 2005) than to the single Omani T. brevicollis than to the neighbouring Mesalina balfouri from Samha and included in Fig. 3. These results suggest that T. brevicollis Socotra (Joger & Mayer 2002), in the two Hemidactylus might be polyphyletic, with Arabian and African popula- (H. forbesi and H. oxyrhinus, unpublished data), and in Pristu- tions being unrelated, a hypothesis that has already been rus abdelkuri (Papenfuss et al. 2009), although in this latter proposed by other authors on the basis of morphological case it is not clear whether Abd Al Kuri was colonized from evidence (Lanza 1983; Fritz & Schu¨tte 1988; Scha¨tti & mainland Arabia or from Africa. At present, there are two Gasperetti 1994). A constrained phylogeny in which annual monsoons that affect the Socotra Archipelago: the T. brevicollis was forced to be monophyletic was not south-west monsoon, which blows from early June to early rejected by either the AU or the SH tests (see results), thus October, and the north-east monsoon, which blows from indicating that, at least from a molecular point of view, the April to May. These two monsoons also affect the direction monophyly of T. brevicollis cannot be rejected and that of the oceanic currents, which change direction depending more data are therefore needed. However, our results sug- on the monsoons. During winter, the flow of the upper gest that the taxonomic status of the Arabian and African ocean is directed westwards from near the Indonesian T. brevicollis needs to be reassessed (Scha¨tti & Gasperetti Archipelago to the Arabian Sea. During summer, the direc- 1994) as current data are insufficient to establish whether tion reverses, with eastwards flow extending from Somalia more than one species is present in Arabia (Fritz & Schu¨tte into the Bay of Bengal (Shankar et al. 2002). According to 1988) and whether animals genetically similar to African the results of the present work and what has been inferred T. brevicollis also occur on the Arabian side of the Red Sea. from other groups such as Mesalina and Hemidactylus (Joger Many names are available for both the Arabian and & Mayer 2002; unpublished results), it is suggested that the African populations: Euprepes brevicollis Wiegmann, 1837 westwards winter monsoon current may have played a very (type locality ‘Abyssinia’ = Ethiopia), Euprepes pyrrhocepha- important role in the transmarine colonization of the Soco- lus Wiegmann, 1837 (type locality: ‘in Aschik, insula maris tra Archipelago by several groups.

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Acknowledgements Continental break up history of a deep magma poor margin Heinz Grillitsch, Richard Gemel and Silke Schweiger (Na- based on seismic reflection data (Northeastern Gulf of Aden turhistorisches Museum, Wien) kindly helped us to exam- margin, offshore Oman). Geophysical Journal International, 180, 501–519. ine the collections of the Museum of Vienna; Annamaria Battistuzzi, F. U., Filipski, A., Hedges, S. B. & Kumar, S. (2010). Nistri and Stefano Vanni have made available the collec- Performance of Relaxed-Clock Methods in Estimating tions of the Museo Zoologico ‘La Specola’ in Florence. Evolutionary Divergence Times and Their Credibility Intervals. Stefano Scali (Museo Civico di Storia naturale di Milano) Molecular Biology and Evolution, 27, 1289–1300. provided pictures of specimens of T. brevicollis held in the Bosworth, W., Huchon, P. & McClay, K. (2005). The Red Sea Milan Museum. This research was accomplished within the and Gulf of Aden basins. Journal of African Earth Sciences, 43, framework of the programmes ‘Socotra Conservation and 334–378. Brandley, M. C., Schmitz, A. & Reeder, T. W. (2005). Partitioned Development’ by United Nations Development Program, Bayesian analyses, partition choice, and the phylogenetic and ‘Capacity Development for Soqotra Archipelago Con- relationships of scincid lizards. Systematic biology, 54, 373. servation’ by the Italian Cooperation. Thanks are due to Broadley, D. G. & Howell, K. (1991). A checklist of the reptiles of PROGES s.r.l. and to the Environment Protection Agency Tanzania, with synoptic keys. Harare and Bulawayo, Zimbabwe: of Socotra for their support during field surveys and for National Museum and Monuments of Zimbabwe. the collecting permits. We would also like to thank the Brown, R. & Pestano, J. (1998). Phylogeography of skinks Minister Abd al-Rahman Fadhl al-Iriyani (Ministry of (Chalcides) in the Canary Islands inferred from mitochondrial DNA sequences. Molecular Ecology, 7, 1183–1191. Water and Environment of Yemen) for his support and Brown, R. P. & Yang, Z. (2010). Bayesian dating of shallow interest in the project. DNA work was funded by grant phylogenies with a relaxed clock. Systematic biology, 59, 119. CGL2009-11663 ⁄ BOS from the Ministerio de Educacio´n Carranza, S. & Arnold, E. (2003). Investigating the origin of y Ciencia, Spain. S.C. and M.M. are members of the Grup transoceanic distributions: mtDNA shows Mabuya lizards de Recerca Emergent of the Generalitat de Catalunya: (Reptilia, Scincidae) crossed the Atlantic twice. Systematics and 2009SGR1462; M.M. is supported by a FPU predoctoral Biodiversity, 1, 275–282. grant from the Ministerio de Ciencia e Innovacio´n, Spain Carranza, S. & Arnold, E. (2006). Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) (AP2008-01844). Research work by S.C. at the BMNH elucidated using mitochondrial DNA sequences. 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New Robinson-Rechavi, M. & Huchon, D. (2000). RRTree: relative- York: United Nations Development Programme. rate tests between groups of sequences on a phylogenetic tree. Wranik, W. (1998b). Contributions to the herpetology of the Bioinformatics, 16, 296. Republic of Yemen. 4. Sokotra Island and southern Yemen Ro¨sler, H. & Wranik, W. (2000). Beitra¨ge zur herpetologie der mainland. Zoologische Abhandlungen Staatliches Museum fu¨r Republik Jemen. 6. Erste u¨bersicht zur herpetofauna der Insel Tierkunde Dresden, 21, 163–179. Darsa (Reptilia: Sauria et Serpentes). Faunistische Abhandlungen Staatliches Museum fu¨r Tierkunde Dresden, 22, 85–94. Supporting Information Ro¨sler, H. & Wranik, W. (2004). A key and annotated checklist Additional Supporting Information may be found in the to the reptiles of the Socotra Archipelago. Fauna of Arabia, 20, online version of this article: 505–534. Appendix S1. Measurements and scale counts of the Samuel, M. A., Harbury, N., Bott, R. & Manan Thabet, A. specimens of Trachylepis brevicollis (all measurements in mm). (1997). Field observations from the Socotran platform: their Appendix S2. Specimens used in estimation of diver- interpretation and correlation to Southern Oman. Marine and petroleum geology, 14, 661–673. gence times and RRT tests. Scha¨tti, B. & Desvoignes, A. (1999). The herpetofauna of Southern Appendix S3: Results of the BEAST analysis including Yemen and the Sokotra Archipelago (pp. 1–178). Gene´ve: two partitions (12S and 16S) performed using the software Instrumenta Biodiversitatis IV, Muse`um d’Histoire Naturelle. BEAST v.1.6.1. Scha¨tti, B. & Gasperetti, J. (1994). A contribution to the Appendix S4: Results of the ML (A) and Bayesian (B) herpetofauna of Southwest Arabia. Fauna of Saudi Arabia, 14, analyses. The alignment of the Mabuya sensu lato dataset 348–423. included a total of 904 base pairs (bp), of which 392 corre- Shankar, D., Vinayachandran, P. N. & Unnikrishnan, A. S. (2002). The monsoon currents in the north Indian Ocean. sponded to the 12S and 512 to the 16S. Progress In Oceanography, 52, 63–120. Appendix S5- Uncorrected p-distances based on gene Shimodaira, H. (2002). An approximately unbiased test of fragment 12S above the diagonal and 16S below the phylogenetic tree selection. Systematic biology, 51, 492. diagonal. Shimodaira, H. & Hasegawa, M. (1999). Multiple comparisons of Appendix S6. Living holotype of Trachylepis cristinae log-likelihoods with applications to phylogenetic inference. (MCCI-R1585). Molecular Biology and Evolution, 16, 1114–1116. Appendix S7. Type specimens of Trachylepis cristinae sp. Shimodaira, H. & Hasegawa, M. (2001). CONSEL: for assessing the confidence of phylogenetic tree selection. Bioinformatics, 17, nov. (a, b, c) – holotype, MCCI-R1585; (d, e, f) – 1246. paratype, NMW 9396. Sindaco, R., Ziliani, U., Razzetti, E., Carugati, C., Grieco, C., Appendix S8. Comparison with other Trachylepis from Pupin, F., Al-Aseily, B. A., Pella, F. & Fasola, M. (2009). A Eastern and Southern Africa, and Arabia. misunderstood new gecko of the genus Hemidactylus from Please note: Wiley-Blackwell are not responsible for the Socotra Island, Yemen (Reptilia: Squamata: Gekkonidae). Acta content or functionality of any supporting materials sup- Herpetologica, 4, 83–98. plied by the authors. Any queries (other than missing Spawls, S., Howell, K., Drewes, R. & Ashe, J. (2002). A field guide to the reptiles of East Africa. San Diego, CA, USA: Princeton material) should be directed to the corresponding author University Press. for the article.

362 ª 2012 The Authors d Zoologica Scripta ª 2012 The Norwegian Academy of Science and Letters, 41, 4, July 2012, pp 346–362 Appendix S1: Measurements and scale counts of the specimens of Trachylepis brevicollis (all measurements in mm). Catalogue: BM = British Museum, MZUF = Museum of Zoology, University of Florence; Locality: OM = Oman, SA = Saudi Arabia, SO = Somalia, YE = Yemen; SVL: snout-vent length; HL: head length from tip of snout to posterior border of parietals; HW: head width at level of ear openings; ratio HL/HW; EST: distance between anterior margin of eye to tip of snout; EED: distance between posterior margin of eye to anterior margin of ear; MSR: number of midbody scale rows; counts for specimens indicated with (*) were published by Lanza & Carfì (1968), together with additional specimens not studied by us; SL l/r: number of supralabials (left / right); LF4 r: number of lamellae under the 4th right finger; LT4 r: number of lamellae under the 4th right toe; Dorsal keels: number of dorsal keels; Supranasals arrangement: S = separated; (C) = in short contact; C = in broad contact; Prefrontal arrangement: S = separated; (C) = in short contact; C = in broad contact; Supraocul. ent. frontal: supraoculars entering the frontal scale, * third plate fragmented; Tarsal scales: shape of tarsal scales (foot); F = flat; G = granular; B = blunt tubercles; P = pointed tubercles; S = spinose (mucronated) tubercles; (S) = slightly spinose; Palmar scales: shape of palmar scales (hand); F = flat; G = granular; P = pointed tubercles; S = spinose (mucronated) tubercles; (S) = slightly spinose; Foot lamellae: shape of lamellae under the 4th toe; Hand lamellae: number of lamellae under the 4th finger; Morphobank images: Morphobank codes (project P461) for the pictures of the T. brevicollis and T. socotrana specimens.

Species Catalogue Locality Sex SVL HL HW ratio HL/HW EST EED MSR SL l/r LF4 r LT4 r Dorsal keels Supranasals arrangement Prefrontals arrangement Supraocul. ent. frontal Tarsal scales Palmar scales Foot lamellae Hand lamellae Morphobank images T. brevicollis BM1975.1385 OM, Dhofar, Salalah ? 120 20.9 18.3 1.14 10.1 9.7 34 5/5 16 20 2 (C) (C) I-II-III / I-II-III F F k sc M86875-M86886 T. brevicollis BM1975.1386 OM, Dhofar, Salalah ? 113 19.5 17.8 1.10 9.5 9.3 34 5/5 15 18 2 (C) S I-II / I-II-III S F wk s M86888-M86900 T. brevicollis BM1978.777 OM, Dhofar, Raysut ? 135 21.4 19.3 1.11 11.3 10.9 34 5/5 16 19 2 S S I-II / I-II-III F F k s M86901-M86914 T. brevicollis BM1977.1177 OM, Dhofar, Jebel Qamair ? 180 23.5 20.7 1.14 12.5 10.2 32 5/5 13 16 2 S S I-II-III / I-II-III F F wk s M86915-M86930 T. brevicollis BM1977.1180 OM, Dhofar, Salalah ? 140 20.8 19.0 1.09 11.2 10.4 37 5/5 15 21 2 S S I-II-III / I-II-III F F k s M86931-M86944 T. brevicollis BM1985.618 OM, Dhofar, Khortawt ? 166 24.4 21.5 1.13 12.8 10.8 34 5/5 16 20 2 (C) S I-II-III / I-II-III F F wk wk M86945-M86957 T. brevicollis BM97.3.11.90.93A YE, Hadramaut M 128 21.7 19.7 1.10 10.9 10.1 33 5/5 17 21 2 C C I-II / broken S F wk s M86988-M87001 T. brevicollis BM97.3.11.90.93B YE, Hadramaut M 121 21.7 20.1 1.08 10.8 9.3 32 5/5 13 17 2 C C I-II-III / I-II-III F F wk wk M87002-M87015 T. brevicollis BM97.3.11.90.93C YE, Hadramaut ? 86 14.9 12.0 1.24 7.8 5.1 34 5/5 15 17 2 S C I-II-III / I-II-III S F k wk M87016-M87167 T. brevicollis MZUF-28902 Arabia ? 135 24.5 20.9 1.17 9.3 9.8 37 5/5 16 22 2 (3) S (C) I-II-III / I-II-III P G - wk M85852-M85860 T. brevicollis MZUF-28903 Arabia ? 138 25.6 21.9 1.17 11.5 9.5 37 5/5 16 23 2 (3) (C) S I-II-III / I-II-III P G - wk M85861-M85866 T. brevicollis MZUF-28651 Arabia ? 109 18.8 18.1 1.04 10.0 10.3 35 5/5 16 20 2 C S I-II / I-II P P k s M85845-M85851 T. brevicollis MZUF-28613 Arabia ? 117 18.8 16.7 1.13 9.8 8.3 30 5/5 15 21 2 (3) C S I-II-III / I-II-III P P k wk M85809-M85814 T. brevicollis MZUF-28612 Arabia ? 121 19.4 19.4 1.00 13.0 9.3 32 5/5 15 20 2 C S I-II-III / I-II P P k wk M85794-M85808 T. brevicollis MZUF-28615 Arabia M 103 19.4 19.6 0.99 13.0 9.2 32 5/5 16 19 2 C S I-II-III / I-II-III P P k k M85830-M85844 T. brevicollis MZUF-28611 Arabia M 98 17.6 15.7 1.12 9.2 8.1 32 5/5 16 21 2 (3) C S I-II-III / I-II-III S S k wk M85785-M85793 T. brevicollis MZUF-28614 Arabia imm 79 13.9 11.2 1.24 7.2 6.3 32 5/5 15 18 ? C (C) I-II-III / I-II-III P P k k M85815-M85829 T. brevicollis BM1982.1162 YE, ab Marawagha ? 126 22.6 18.9 1.20 11.6 10.1 35 5/5 15 20 2 S S I-II-III / I-II-III F F wk wk M87167-M87182 T. brevicollis BM1988.312 YE, Wadi Warazan ? 146 23.6 21.3 1.11 12.2 10.7 34 5/5 15 20 3 C (C) I-II / I-II-III B F wk s M87183-M87198 T. brevicollis BM99.12.13.70 YE, Abian Hill Country F 114 19.4 16.3 1.19 9.7 8.2 32 5/5 17 21 3 C C I-II-III / I-II-III S F k s M87199-M87213 T. brevicollis BM99.12.13.74 YE, Abian Country ? 127 20.7 19.6 1.06 10.7 9.6 32 5/5 15 21 2 C C I-II-III / I-II-III S F k s M87567-M87580 T. brevicollis BM95.5.23.74 YE, Sheikh Osman M 122 21.0 18.2 1.15 9.9 8.9 31 5/5 15 21 3 C C I-II-III / I-II-III S F k s M87581-M87593 T. brevicollis BM95.5.23.75 YE, Sheikh Osman F 128 20.5 16.3 1.26 10.3 8.3 32 5/5 14 19 3 C C I-II-III / I-II-III S S wk wk M87594-M87607 T. brevicollis BM95.5.23.71 YE, Aden F 145 24.1 23.0 1.05 12.9 10.2 32 6/5 16 21 3 C C I-II-III / I-II-III S F wk s M87608-M87619 T. brevicollis BM95.5.23.70 YE, Aden M 140 22.4 20.2 1.11 11.3 10.6 32 5/5 15 22 3 C (C) I-II-III / I-II-III S S k s M89774-M89787 T. brevicollis BM1903.3.6.45 SA, Azrahi Ravine M 120 22.3 20.1 1.11 11.2 10.6 35 5/5 16 20 3 C S I-II-III / I-II-III S F wk s M87621-M87635 T. brevicollis BM1903.3.6.46 SA, Gerba ? 117 19.0 15.0 1.27 10.2 8.9 34 5/5 15 19 3 C S I-II-III / I-II-III F F k wk M87636-M87647 T. brevicollis BM1903.3.6.16 SA, el Kubar ? 134 21.7 19.7 1.10 11.4 7.3 34 5/5 16 20 2-3 C S I-II-III / I-II-III F F wk s M87649-M87662 T. brevicollis BM1903.3.6.15 SA, el Kubar ? 121 20.1 17.5 1.15 10.7 9.0 34 5/5 14 19 2 C S I-II-III / I-II-III B F k wk M87664-M87676 T. brevicollis BM1903.6.26.16 SA, el Kubar ? 119 20.0 17.6 1.14 10.7 8.8 34 ??? 16 19 2-3 S (C) I-II / I-II F F k wk M87678-M87690 T. brevicollis BM1979.975 SA, al Jammum ? 121 19.0 16.0 1.19 9.3 8.8 32 5/4 16 19 2 (C) (C) I-II-III / I-II S S k wk M87691-M87703 T. brevicollis BM1964.144 SA, Wadi Lahsiba ? 96 16.7 14.0 1.19 9.3 6.6 ? 5/4 13 18 2 C S I-II-III / I-II F F k wk M87704-M87715 T. brevicollis BM1980.60 SA, 30 km SE Abha ? 130 21.9 19.8 1.11 11.3 10 35 5/5 13 20 2 C S I-II-III / I-II-III F F wk s M87716-M87730 T. brevicollis BM1980.59 SA, Al Jemun ? 114 19.2 16.3 1.18 9.6 8.1 33 5/5 16 18 2 C C I-II-III / I-II-III S F k wk M87731-M87744 Summary Statistics Number of Individuals (N) 34 34 34 34 34 34 33 33 34 34 Mean 123.7 20.6 18.2 1.13 10.6 9.1 33.3 4.9 15.2 19.7 Maximum 180 25.6 23 1.27 13 10.9 37 5 17 23 Minimum 79 13.9 11.2 0.99 7.2 5.1 30 4 13 16 Standard Error Mean 3.41 0.44 0.45 0.011 0.23 0.23 0.29 0.02 0.18 0.26

Species Catalogue Locality Sex SVL HL HW ratio HL/HW EST EED MSR SL l/r LF4 r LT4 r Dorsal keels Supranasals arrangement Prefrontals arrangement Supraocul. ent. frontal Tarsal scales Palmar scales Foot lamellae Hand lamellae Morphobank images T. brevicollis MZUF-1708 SO, Dinsor ? 122 19.5 22.0 0.89 9.3 9.0 (*) 5/4 15 17 2 (3) S S I-II-(III) / I-II S S k k M85718-M85720 T. brevicollis MZUF-1599 SO, Dinsor ? 122 20.6 20.7 1.00 10.9 10.0 (*) 4/5 13 16 2 C S I-II-III / I-II S S k wk M85715-M85717 T. brevicollis MZUF-1759 SO, Dinsor M 139 22.5 23.7 0.95 11.6 11.0 (*) 5/4 13 18 2 S S (I)-II / I-II S S k k M85734-M85736 T. brevicollis MZUF-1756 SO, Dinsor M 124 20.2 20.9 0.97 10.9 11.0 (*) 5/5 14 16 3 (C) (C) I-II / I-II S S k wk M85728-M85730 T. brevicollis MZUF-1757 SO, Dinsor ? 130 20.9 19.7 1.06 11.3 11.3 (*) 5/5 15 17 2/3 (C) (C) I-II-III / I-II-III S S k wk M85731M85733 T. brevicollis MZUF-1709 SO, Dinsor ? 136 22.1 23.0 0.96 12.0 10.4 (*) 5/5 12 17 2/3 S S I-II-III / I-II S (S) k wk M85721-M85723 T. brevicollis MZUF-1755 SO, Dinsor imm 114 18.8 16.5 1.14 - - (*) 5/5 14 17 2 (C) S I-II-(III) / I-II S S k k M85724-M85727 T. brevicollis MZUF-1751 SO, Dinsor imm 102 16.5 17.1 0.96 - - (*) 5/5 13 18 2 (C) S I-II / I-II S S k k - T. brevicollis MZUF-10131 SO, Gardo ? 118 19.5 22.5 0.87 11.6 10.3 (*) 5/5 14 19 2 (3) (C) S (I)-II / (I)-II-III S S k k M85750-M85753 T. brevicollis MZUF-10129 SO, Gardo ? 125 21.1 22.5 0.94 11.2 10.9 (*) 5/5 15 18 2 (3) (C) S I-II-(III) / I-II-(III) S S k k M85743-M85745 T. brevicollis MZUF-10137 SO, Gardo ? 121 19.9 21.0 0.95 11.2 10.5 (*) 5/5 13 17 2 (3) S S II / II-(III) S S k k M85768-M85772 T. brevicollis MZUF-10132 SO, Gardo ? 115 20.5 21.8 0.94 11.7 11.0 (*) 5/5 15 19 2 (3) S (C) I-II-III / (I)-II-III S S k k M85754-M85756 T. brevicollis MZUF-10138 SO, Gardo ? 125 20.3 20.3 1.00 11.6 10.9 (*) 5/5 13 17 2 (3) S (C) (I)-II / I-II S S k k M85773-M85775 T. brevicollis MZUF-10134 SO, Gardo ? 118 19.8 20.0 0.99 11.4 11.2 (*) 5/6 14 17 2 (3) C S I-II-III / I-II-III S S k k M85761-M85764 T. brevicollis MZUF-10130 SO, Gardo ? 117 19.0 18.2 1.04 10.4 10.1 (*) 5/4 - 18 2 S (C) I-II-(III) / I-II-(III) S S k k M85746-M85749 T. brevicollis MZUF-10133 SO, Gardo ? 114 20.0 21.4 0.93 10.9 10.5 (*) 5/4 13 19 3 (2) C S I-II / I-II-III S S k wk M85757-M85760 T. brevicollis MZUF-10154 SO, Gardo ? 101 17.4 16.9 1.03 9.7 9.6 (*) 5/4 15 17 2 (3) S (C) I-II / I-II-III S S k wk M85776-M85778 T. brevicollis MZUF-10136 SO, Gardo ? 94 17.7 17.4 1.02 9.6 9.2 (*) 5/5 13 18 2 S (C) I-II / II S S k k M85765-M85767 T. brevicollis BM1937.12.5.669 SO, Bohodle ? 122 18.5 18.3 1.01 10.1 8.2 32 5/5 12 16 2 C C II-III / II-III S F k wk M87745-M87758 T. brevicollis BM1937.12.5.675 SO, Haud ? 123 19.9 18.6 1.07 11.6 8.0 33 5/5 15 16 2 C C II-III / II-III S F k wk M87759-M87774 T. brevicollis BM1937.12.5.671 SO, Horu Fodi ? 110 19.0 17.3 1.10 9.7 9.0 33 5/5 12 16 2 C C II-III / II-III B F k wk M87775-M87789 T. brevicollis BM1968.1249 SO, Dinsor M 130 21.1 19.3 1.09 10.6 9.2 33 5/5 12 16 2-3 S S I-II / I-II S F k wk M87790-M87803 T. brevicollis BM1994.526 SO, Argeisa ? 135 22.6 21.5 1.05 11.2 11.1 32 4/4 14 17 2 C C I-II-III / I-II S S wk wk M87804-M87817 Summary Statistics Number of Individuals (N) 23 23 23 23 21 21 5 23 22 23 Mean 119.8 19.8 20 0.99 10.8 10.1 32.6 4.8 13.6 17.2 Maximum 139 22.6 23.7 1.14 12 11.3 33 5.5 15 19 Minimum 94 16.5 16.5 0.87 9.3 8 32 4 12 16 Standard Error Mean 2.31 0.31 0.44 0.014 0.17 0.21 0.24 0.06 0.23 0.2

Species Catalogue Locality Sex SVL HL HW ratio HL/HW EST EED MSR SL l/r LF4 r LT4 r Dorsal keels Supranasals arrangement Prefrontals arrangement Supraocul. ent. frontal Tarsal scales Palmar scales Foot lamellae Hand lamellae Morphobank images T. socotrana NMW9397 YE. Socotra ? 97 15.9 14.5 1.10 8.2 11.7 33 4/4 14 23 3 (5) C S II III / II III - - - - M89788-M89793 T. socotrana NMW9393 (1) YE. Socotra ? 87 12.8 9.4 - 7.4 6.3 32 4/4 14 19 3 C S II* / II III blunt blunt wk s M89794-M89802 T. socotrana NMW9393 (2) YE. Socotra ? 78 13.1 11.2 1.17 6.4 5.2 33 4/4 15 21 3 C S II. III / (I) II III blunt blunt k s M89806-M89814 T. socotrana NMW9395 (1) YE. Socotra ? 72 12.4 10.8 1.15 6.1 5.5 32 4/4 15 19 3 C S II / II (III) blunt blunt wk s M89815-M89821 T. socotrana NMW9395 (2) YE. Socotra ? 75 12.3 11 1.12 6.2 5.5 34 4/4 14 21 3 C S II III / II III wk blunt k s M89788-M89793 T. socotrana NMW9395 (3) YE. Socotra ? 78 13.1 11.6 1.13 6.8 5.6 32 4/4 14 18 3 C S II III / II III blunt blunt wk s M89830-M89836 T. socotrana NMW9398 (1) YE. Socotra ? 90 14.3 13.5 1.06 7.5 6.8 32 4/4 14 20 5 C S II III / II III blunt blunt wk s M89837-M89844 T. socotrana NMW9398 (2) YE. Socotra ? 81 13.3 11.2 1.19 6.8 6.5 32 4/4 14 21 3 C S II III / II III blunt blunt s s M89854-M89890 T. socotrana NMW9394 (1) YE. Socotra ? 85 13.3 11.9 1.12 6.9 6.1 33 4/4 14 21 3 (5) C S (I) II III / II III blunt blunt wk s M89891-M89897 T. socotrana NMW9394 (2) YE. Socotra ? 83 13.3 12 1.11 6.8 6.2 32 4/4 13 19 3 (5) C S II III / II III blunt blunt wk s M89898-M89904 Summary Statistics Number of Individuals (N) 10 10 10 9 10 10 10 10 10 10 Mean 82.6 13.3 11.7 1.12 6.9 6.5 32.5 4 14.1 20.2 Maximum 97 15.9 14.5 1.19 8.2 11.7 34 4 15 23 Minimum 72 12.3 9.4 1.06 6.1 5.2 32 4 13 18 Standard Error Mean 2.36 0.33 0.45 0.012 0.20 0.59 0.22 0.00 0.17 0.46

Appendix S2 Specimens used in estimation of divergence times and RRT tests

Taxa Locality GenBank Accession Code Fig.X Reference Numbers 12S/16S Cordylus warreni NC_005962 Corwar Kumazawa 2004 Gerrhosaurus validus Limpopo, RSA HQ167135/HQ167246 Gerval Stanley et al. 2011

Eumeces schneideri schneideri Egypt EU278004/EU278071 Eumsch1 Carranza et al. 2008 Eumeces schneideri algeriensis Massa, Morocco EU278021/EU278086 Eumalg1 Carranza et al. 2008 Chalcides boulengeri Oued Shili, Tunisia EU277924/EU278045 Chabou1 Carranza et al. 2008 Chalcides sepsoides (Egypt EU277925/EU278046 Chasep1 Carranza et al. 2008 Chalcides sphenopsiformis Massa, Morocco EU277875/EU278031 ChasphMO Carranza et al. 2008 Chalcides mauritanicus Ras El Ma, Morocco EU277971/EU278060 Chamau1 Carranza et al. 2008 Chalcides guentheri Nahal Qetalav, Israel EU278001/- Chague Carranza et al. 2008 Chalcides minutus 1 Debdou, Morocco EU277972/EU278061 Chamin1 Carranza et al. 2008 Chalcides minutus 2 Jebel Bou Iblane, Morocco EU277973/EU278062 Chamin2 Carranza et al. 2008 Chalcides mertensi Ain Soltane, Tunisia EU277975/EU278064 Chamer2 Carranza et al. 2008 Chalcides chalcides chalcides Giglio Island, Italy EU277979/- Chachac3 Carranza et al. 2008 Chalcides chalcides vittatus Tunis, Tunisia EU277984/EU278065 Chachav1 Carranza et al. 2008 Chalcides pseudostriatus Skhirat, Morocco EU277985/EU278066 Chapse1 Carranza et al. 2008 Chalcides striatus Salamanca, Spain EU277995/EU278067 Chastr11 Carranza et al. 2008 Chalcides colosii Mokrisset, Morocco EU277930/EU278049 Chacol1 Carranza et al. 2008 Chalcides parallelus Ras El Ma, Morocco EU277921/EU278043 Chapar Carranza et al. 2008 Chalcides lanzai Azrou, Morocco EU277920/EU278042 Chalan1 Carranza et al. 2008 Chalcides bedriagai bedriagai Jaen, Spain EU277916/EU278041 Chabedb2 Carranza et al. 2008 Chalcides ocellatus ocellatus Tata, Morocco EU277938/EU278052 Chaoco1 Carranza et al. 2008 Chalcides ocellatus tiligugu Algiers, Algeria EU277937/EU278051 Chaoct3 Carranza et al. 2008 Chalcides ocellatus ocellatus Negev Desert, Israel EU277951/EU278055 Chaoco13 Carranza et al. 2008 Chalcides ocellatus ocellatus Cyprus EU277943/EU278053 Chaoco18 Carranza et al. 2008 Chalcides ocellatus tiligugu Ain Draham, Tunisia EU277958/EU278057 Chaoct33 Carranza et al. 2008 Chalcides viridanus Tenerife, Canary Islands, Spain EU277885/EU278036 ChavirTE Carranza et al. 2008 Chalcides sexlineatus sexlineatus Gran Canaria, Canary Islands, Spain EU277880/EU278034 ChassGC Carranza et al. 2008 Chalcides sexlineatus bistriatus Gran Canaria, Canary Islands, Spain AF054530/AF054544 Chasb4GC Brown & Pestano 1998 Chalcides sexlineatus bistriatus Gran Canaria, Canary Islands, Spain EU277879/EU278033 Chasb6GC Carranza et al. 2008 Chalcides coeruleopunctatus La Gomera, Canary Islands, Spain EU277892/EU278038 Chaco1GO Carranza et al. 2008 Chalcides coeruleopunctatus El Hierro, Canary Islands, Spain EU277891/EU278037 Chaco4HI Carranza et al. 2008 Chalcides simonyi Fuerteventura, Canary Islands, Spain EU277872/EU278030 Chasim Carranza et al. 2008 Chalcides mionecton trifasciatus Sidi Ifni, Morocco EU277870/EU278029 Chamiot1 Carranza et al. 2008 Chalcides mionecton mionecton Essaouira, Morocco EU277868/EU278028 Chamiom2 Carranza et al. 2008 Chalcides manueli Sidi Ifni, Morocco EU277857/EU278023 Chaman1 Carranza et al. 2008 Chalcides polylepis Marrakech, Morocco EU277859/EU278024 Chapol3 Carranza et al. 2008 Chalcides polylepis Morocco EU277863/EU278026 Chapol6 Carranza et al. 2008

Appendix S3: Results of the BEAST analysis including two partitions (12S and 16S) performed using the software BEAST v.1.6.1 (Drummond & Rambaut 2007) with the following model and prior specifications (otherwise, by default): GTR+G+I, Relaxed Uncorrelated Lognormal Clock (rates 0.01305 and 0.008, respectively), Yule process of speciation, random starting tree, alpha Uniform (0,10), yule.birthRate (0,1000), nucleotide substitution rates Uniform (1,100) initial value=1. See Materials & Methods for more details

Appendix S4: Results of the ML (A) and Bayesian (B) analyses. The alignment of the Mabuya sensu lato dataset included a total of 904 base pairs (bp), of which 392 corresponded to the 12S and 512 to the 16S. The alignment matrix is available from Morphobank (Doc1) and the codes used in the matrix can be deciphered with Table 1. Of the total of 392 bp of the aligned 12S sequences 294 were variable and 169 parsimony-informative, while the respective sites for the 512 bp long 16S fragment were 202 and 195.

A

B

Appendix S5- Uncorrected p-distances based on gene fragment 12S above the diagonal and 16S below the diagonal.

1 2 3 4 5 6 7 8 9 10 11 1.- T. brevicollis 1 0.0421 0.0368 0.0500 0.0500 0.0368 0.0763 0.0763 0.0763 0.0737 0.0737 2.- T. brevicollis 2 0.0434 0.0211 0.0395 0.0395 0.0421 0.0895 0.0895 0.0895 0.0868 0.0868 3.- T. brevicollis 3 0.0456 0.0217 0.0395 0.0395 0.0421 0.0842 0.0842 0.0842 0.0868 0.0816 4.- T. dichroma 2 0.0282 0.0390 0.0412 0.0000 0.0500 0.0763 0.0763 0.0763 0.0737 0.0737 5.- T. dichroma 1 0.0282 0.0390 0.0412 0.0000 0.0500 0.0763 0.0763 0.0763 0.0737 0.0737 6.- T. cristinae sp.n. 0.0239 0.0456 0.0434 0.0325 0.0325 0.0789 0.0789 0.0789 0.0763 0.0763 7.- T. socotrana 4 0.0716 0.0738 0.0672 0.0694 0.0694 0.0694 0.0000 0.0000 0.0026 0.0053 8.- T. socotrana 3 0.0716 0.0738 0.0672 0.0694 0.0694 0.0694 0.0000 0.0000 0.0026 0.0053 9.- T. socotrana 5 0.0846 0.0868 0.0803 0.0824 0.0824 0.0824 0.0130 0.0130 0.0026 0.0053 10.- T. socotrana 1 0.0716 0.0738 0.0672 0.0694 0.0694 0.0694 0.0000 0.0000 0.0130 0.0079 11.- T. socotrana 2 0.0694 0.0716 0.0651 0.0672 0.0672 0.0672 0.0022 0.0022 0.0152 0.0022

Appendix S8

Comparison with other Trachylepis from Eastern and Southern Africa, and Arabia.

T. cristinae differs from other Arabian, East- and Southern African Trachylepis by the following characters, according to Lanza and Carfì (1968); Largen and Spawls (2006); Arnold (1986); Spawls, Howell et al. (2002); Broadley and Howell (1991); Günther, Whiting et al. (2005); in square brackets are indicated the state of the same characters in T. cristinae.

T. bayoni: frontoprietals fused [not fused]; lower margin of the subocular clearly shorter than its upper margin [not clearly shorter], sometimes excluded by the lip [reaching the lip]; E-African highland endemic. T. boulengeri: 28-32 midbody scales [38]; build slender [robust]; scales with (3) 7-9 (11) keels [2-3].

T. brauni: small (Ltot to 13 cm) [large, Ltot = 22 cm]; subocular clearly shorter than its upper margin (less than a third of upper border) [not clearly shorter]; dorsals usually with 2 keels, or with a poorly defined median keel [2-3 keels]; distinct pale vertebral and dorsolateral stripes [different pattern]; scales on soles of feet keeled and spinose [smooth and not spinose]; a mountain endemic of S-Tanzania. T. dichroma: dorsal and lateral scales with 2 keels [2-3]; different finger 3>4>2>5>1 [3≈4>2>5>1] and toe 3>4>2>5>1 [4>3>5>2>1] formulae; frontal in contact with three supraoculars [only 2]; frontoparietals not in contact [in contact]; subdigital lamellae keeled [smooth]; different colouration. T. hemmingi: dorsals unkeeled in the anterior part of the body [keeled], with 3 barely visible keels posteriorly. T. hildebrantii: auricular lobes conspicuously long and pointed [almost absent]; adult males uniform pale brown above with a series of large dark blotches in a longitudinal row behind the ear [different pattern], body scales sharply 3-keeled, in 32 series at midbody. T. irregularis: subocular with upper margin clearly shorter than its upper margin (less than a third of upper border) [not clearly shorter]; dorsals with 2-5 keels [2-3 keels]; a distinct vertebral double-stripe and dorsolateral stripes [different pattern]; a mountain endemic of Kenya. T. isselii: frontoparietal shields fused [not fused]; mid-body scale rows 30-34 [38], the dorsals tricarinate [2-3 carinated]; a dark lateral band from eye to groin with a pale dorsolateral stripe above and pale lateral stripe beneath [different pattern]. T. maculilabris: dorsal scales with 5-8 keels [2-3]; mid-body scale rows 30-38 (but rarely more than 34) [38]; pattern usually with distinct white, black-speckled lips [different head pattern], but occasionally some individuals have dark vertical flank bars. T. margaritifer: mid-body scale rows 38-52 [38]; adult males uniform above with a series of large dark blotches in a longitudinal row behind the ear, young and female striped [different pattern]. T. megalura: slim build [robust] with little limbs [well developed] and very long tail [not particularly elongated]; mid-body scale rows 22-28 [38]; dorsal scales smooth [keeled]; striped pattern [different pattern]. T. planifrons: mid-body scale rows 26-32 [38]; dorsal scales with 3 (rarely 4-5) keels [2-3 keels]; supranasals generally in broad contact [in narrow contact]; dark lateral band strongly demarcated from the pale lower flanks [different pattern]. T. quinquetaeniata: adult males uniform above with a series of large dark blotches in a longitudinal row behind the ear, young and female striped [different pattern]; mid- body scale rows 32-46 (most commonly 34-40) [38]; dorsal scales with 3 (rarely 4 or 5) distinct keels [2-3 keels]; T. striata: subocular scale usually excluded from the edge of the lip [entering the lip]; a dark lateral band from eye to groin and broad dorsolateral pale stripes [different pattern]; scales on soles of feet keeled and spinose [unkeeled and not spinose]. T. varia: lower margin of the subocular clearly shorter than its upper margin [not clearly shorter]; a dark lateral band from eye to groin bordered by conspicuous pale dorsolateral and lateral stripes [different pattern]; small (SVL ~ 60 mm) [SVL = 114 mm]; scales on soles of feet keeled and spinose [unkeeled and not spinose]. T. wingati: a very prominent pale line with distinct dark margins running from beneath the eye, along the lower flank to the groin [different pattern]; mid-body scale rows 30-32 [38]; dorsal scales with 3 distinct keels [2-3]. T. septemtaeniata: 4 sopraocular scales in front of the subocular [5]; typically a pattern of dark stripes on foreparts and light dorsolateral streaks along body [different pattern]; 32-38 midbody scales [38]; third supraocular shield in contact with the frontal shield [not in contact]. T. tessellata: back scales virtually smooth [keeled]; frontal scale does not contact the 1st supraocular [frontal entering the 1st sopraocular]; 4 sopralabial scales in front of the subocular [5]; colour usually uniform [pattern different]; 29-34 scaled around midbody [38].

According to FitzSimons (1943), Branch (1992) and Laurent (1964), among other southern African species, T. capensis, T. chimbana, T. occidentalis and T. variegata have scales on sole of feet keeled and usually spinose, and subdigital lamellae sharply uni- or tri-carinated, as well T. acutilabris, T. damarana, T. lacertiformis, T. sulcata, T. spilogaster, and T. punctulata that have also the subocular distinctly narrowed below or not reaching the lip. T. homalocephala (including the taxa peringueyi, smithii and depressa) is characterized by a lower number of scales around midbody (28-30). T. 2 binotata has the subocular narrowed below, the lower border from ½ to /3 length of upper, and a broad black band on either side of neck. T. hoeschi and T. laevis have 29- 33 midbody scales; moreover T. laevis has smooth dorsal scales.