ZooKeys 355: 79–107, doi: 10.3897/zookeys.355.6190
Two newly recognized species of Hemidactylus (Squamata, Gekkonidae) from the Arabian Peninsula and Sinai, Egypt
Jiří Šmíd 1,2,†, Jiří Moravec 1,‡, Lukáš Kratochvíl 3,§, Václav Gvoždík 1,|, Abdul Karim Nasher 4,¶, Salem M. Busais 5,6,#, Thomas Wilms 7,††, Mohammed Y. Shobrak 8,‡‡, Salvador Carranza 9,§§
1 Department of Zoology, National Museum, Cirkusová 1740, Prague, Czech Republic
2 Department of Zoology, Faculty of Science, Charles University in Prague, Viničná 7, Prague, Czech Republic
3 Department of Ecology, Faculty of Science, Charles University in Prague, Viničná 7, Prague, Czech Republic
4 Faculty of Science, University of Sana’a, Sana’a, Yemen
5 Biology Department, Faculty of Education, University of Aden, Aden, Yemen
6 Faculty of Sciences, University of Hail, Hail, Saudi Arabia
7 Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, Frankfurt am Main, Germany
8 Biology department, Faculty of Science, Taif University 888, Taif, Saudi Arabia
9 Institute of Evolutionary Biology (CSIC-Universitat Pompeu Fabra), Passeig Marítim de la Barceloneta 37–49, Barcelona, Spain

Corresponding author: Jiří Moravec (jiri.moravec@nm.cz)

Academic editor: Pavel Stoev

received 3 September 2013 | accepted 15 November 2013 | Published 25 November 2013

(C) 2013 Jiří Šmíd. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Šmíd J, Moravec J, Kratochvíl L, Gvoždík V, Nasher AK, Busais SM, Wilms T, Shobrak MY, Carranza S (2013) Two newly recognized species of Hemidactylus (Squamata, Gekkonidae) from the Arabian Peninsula and Sinai, Egypt. ZooKeys 355: 79–107. doi: 10.3897/zookeys.355.6190


A recent molecular phylogeny of the Arid clade of the genus Hemidactylus revealed that the recently described H. saba and two unnamed Hemidactylus species from Sinai, Saudi Arabia and Yemen form a well-supported monophyletic group within the Arabian radiation of the genus. The name ‘Hemidactylus saba species group’ is suggested for this clade. According to the results of morphological comparisons and the molecular analyses using two mitochondrial (12S and cytb) and four nuclear (cmos, mc1r, rag1, rag2) genes, the name Hemidactylus granosus Heyden, 1827 is resurrected from the synonymy of H. turcicus for the Sinai and Saudi Arabian species. The third species of this group from Yemen is described formally as a new species H. ulii sp. n. The phylogenetic relationships of the members of ‘Hemidactylus saba species group’ are evaluated and the distribution and ecology of individual species are discussed.


Reptilia, Gekkonidae, molecular phylogeny, Arabia, Red Sea, Hemidactylus saba species group , Hemidactylus granosus Heyden, 1827 , Hemidactylus ulii sp. n.


The genus Hemidactylus Oken, 1817, the second most species-rich genus of Gekkonidae (122 currently valid species; Uetz 2013), has been witnessing a species-description boom within the last decade. Eighteen species have been described within the last two years, most of them from the Arabian Peninsula and surroundings areas where 13 new species and a new subspecies have been discovered (Busais and Joger 2011a; Moravec et al. 2011; Torki et al. 2011; Carranza and Arnold 2012). Despite the large number of taxa added recently to the Arid clade of Hemidactylus [sensu Carranza and Arnold (2006)], it has been shown that the real diversity of Hemidactylus in Arabia and northeast Africa is still underestimated, with at least seven species remaining to be described (Busais and Joger 2011b; Moravec et al. 2011; Šmíd et al. 2013). A recent study (Šmíd et al. 2013) revealed that two of these newly recognized but still unnamed species, one from Sinai [labelled in accordance to previous works (Moravec et al. 2011; Šmíd et al. 2013) as Hemidactylus sp. 1] and one from Yemen (Hemidactylus sp. 4), clustered with the recently described Yemeni endemic Hemidactylus saba Busais & Joger, 2011. They form a very well supported clade within the Arabian radiation of the genus (Fig. 1). Although the phylogenetic relationships among these three species were not resolved satisfactorily, it was inferred that they began to diversify approximately 7 million years ago (95% highest posterior density interval 4.3–10), what was followed by a subsequent dispersal of the Sinai species from southern Arabia to the north (Šmíd et al. 2013).

Figure 1.

Phylogeny of the Hemidactylus Arid clade (light grey rectangle) modified after Šmíd et al. (2013). Dark grey rectangle highlights the Arabian radiation of this clade, dashed red line delimits the ‘Hemidactylus saba species group’ dealt with in this study. Black dots indicate ML bootstrap values ≥ 70 and BI posterior probabilities ≥ 0.95.

The discovery of a monophyletic species group consisting of one recently described and two newly recognized species calls upon a more thorough study of the nomenclatural status, evolutionary relationships, taxonomy and distribution of its members based on further genetic and morphological data. The present study focuses on this task.

Material and methods
Material for phylogenetic analyses

In order to resolve the phylogenetic relationships between the two newly recognized Hemidactylus species and Hemidactylus saba based on genetic data, a dataset containing only representatives of these three species was assembled. Apart from the data used by Šmíd et al. (2013), additional sequences of the following specimens were produced (Table 1): the holotype and two paratypes of Hemidactylus saba (the only known existing material), 21 individuals from Sinai and Saudi Arabia belonging to Hemidactylus sp. 1 (Šmíd et al. 2013), and five individuals of the undescribed species from Yemen (Hemidactylus sp. 4; Šmíd et al. 2013), one of which was included in the study by Busais and Joger (2011a) (labelled as ‘OTU 7’ therein). Total genomic DNA was extracted using DNeasy Blood & Tissue Kit (Qiagen). Subsequently, sequences for up to two mitochondrial (12SrRNA [12S] – ca. 400 bp and cytochrome b [cytb] – 307 bp) and four nuclear (cmos – 402 bp, mc1r – 666 bp, rag1 – 1023 bp, rag2 – 408 bp) were produced using primers and PCR conditions described in details elsewhere (Šmíd et al. 2013). Chromatograms of all newly obtained sequences were checked by eye and assembled in Geneious 5.6.5 (Biomatters, http://www.geneious.com/​​). All genes were aligned individually using MAFFT (Katoh and Toh 2008) with the iterative refinement algorithm with 1000 iterations. Poorly aligned positions in the alignment of 12S were eliminated with Gblocks (Castresana 2000) under low stringency options (Talavera and Castresana 2007), producing a final 12S alignment of 386 bp. Alignments of all coding genes were trimmed so that all started by the first codon position and no stop codons were revealed when translated into amino acids with the appropriate genetic codes.

Table 1.

List of material used for the phylogenetic analyses. Holotype of Hemidactylus ulii sp. n. and Hemidactylus saba are in bold. The column ‘Loc. No’ refers to the locality number as shown in Fig. 6.

Species Code Museum number Country Locality Loc. No Lat, Long 12S cytb cmos mc1r rag1 rag2
Hemidactylus granosus Sher10660 SMB 10660 Egypt Ayoun Musa 1 29.875, 32.649 JQ957071 JQ957216 JQ957148 JQ957282 - JQ957409
Hemidactylus granosus Hd41 NMP6V70163/2 Egypt Sharm el Sheik; Sinai 2 27.885, 34.317 KC818724 HQ833759 JQ957148 - KC818981 KF647606
Hemidactylus granosus Hd96 NMP6V70163/1 Egypt Sharm el Sheik; Sinai 2 27.885, 34.317 KC818724 HQ833759 - - - KF647607
Hemidactylus granosus Hd97 NMP6V70163/3 Egypt Sharm el Sheik; Sinai 2 27.885, 34.317 KC818724 HQ833759 - - - KF647608
Hemidactylus granosus HSA63 ZFMK 94084 Saudi Arabia Al Wajh 3 26.208, 36.4976 KC818724 HQ833759 KF647576 KF647589 KF647596 KF647610
Hemidactylus granosus HSA64 ZFMK 94085 Saudi Arabia Al Wajh 3 26.208, 36.4976 KF647571 - - - - -
Hemidactylus granosus HSA65 ZFMK 94086 Saudi Arabia 15 km S of Al Wajh 4 26.123, 36.5689 KF647570 KF647581 KF647574 KF647590 KF647601 KF647610
Hemidactylus granosus HSA66 ZFMK 94087 Saudi Arabia 15 km S of Al Wajh 4 26.123, 36.5689 KC818724 - - - - -
Hemidactylus granosus HSA67 ZFMK 94088 Saudi Arabia 15 km S of Al Wajh 4 26.123, 36.5689 KF647569 - - - - -
Hemidactylus granosus HSA68 TUZC-R8 Saudi Arabia 15 km S of Al Wajh 4 26.123, 36.5689 KF647570 - - - - -
Hemidactylus granosus HSA69 ZFMK 94089 Saudi Arabia 15 km S of Al Wajh 4 26.123, 36.5689 KF647570 - - - - -
Hemidactylus granosus HSA70 TUZC-R9 Saudi Arabia 72 km N of Umluj 5 25.614, 36.9867 KF647569 KF647582 JQ957148 KF647591 KF647600 KF647609
Hemidactylus granosus HSA62 TUZC-R10 Saudi Arabia 180 km W of Hail 6 26.883, 40.0874 KF647569 KF647585 JQ957148 KF647588 KF647602 KF647609
Hemidactylus granosus HSA61 IBES10001 Saudi Arabia Al Ghat 7 26.054, 45.0003 KF647569 KF647585 JQ957148 KF647588 KF647599 KF647610
Hemidactylus granosus HSA57 IBES10183 Saudi Arabia 30 km NE of Alhawiyah 8 21.624, 40.7094 KF647568 KF647580 - - KF647597 KF647610
Hemidactylus granosus HSA58 ZFMK 94090 Saudi Arabia 30 km NE of Alhawiyah 8 21.624, 40.7094 KF647569 - - - - -
Hemidactylus granosus HSA59 TUZC-R11 Saudi Arabia 30 km NE of Alhawiyah 8 21.624, 40.7094 KF647569 - - - - -
Hemidactylus granosus HSA60 IBES10344 Saudi Arabia 30 km NE of Alhawiyah 8 21.624, 40.7094 KF647569 KF647583 - - KF647598 KF647610
Hemidactylus granosus HSA54 IBES10150 Saudi Arabia 20 km S of Ashayrah 9 21.602, 40.6911 KF647568 KF647584 KF647576 KF647588 KF647595 KF647609
Hemidactylus granosus HSA55 ZFMK 94091 Saudi Arabia 20 km S of Ashayrah 9 21.602, 40.6911 KF647569 KF647584 KF647575 KF647588 KF647596 KF647610
Hemidactylus granosus HSA56 IBES10363 Saudi Arabia 20 km S of Ashayrah 9 21.602, 40.6911 KF647569 - - - - -
Hemidactylus granosus ZFMK 87236 ZFMK 87236 Saudi Arabia Taif National Wildlife Research Center 10 21.25, 40.96 KF647569 - - - - -
Hemidactylus saba BJ27 NHM-BS N41914 Yemen Marib 17 14.9, 45.5 KF647567 - KF647573 - - KF647605
Hemidactylus saba BJ28 NHM-BS N41913 Yemen Marib 17 14.9, 45.5 KF647567 KF647579 KF647573 KF647586 - KF647605
Hemidactylus saba BJ29 NHM-BS N41912 Yemen Marib 17 14.9, 45.5 KF647567 - KF647573 KF647587 KF647594 KF647605
Hemidactylus ulii sp. n. JS48 NMP6V 74834/1 Yemen Wadi Zabid 11 14.147, 43.517 KC818730 KC818881 KC818789 KC818943 KC819001 KC819062
Hemidactylus ulii sp. n. JS49 NMP6V 74834/2 Yemen Wadi Zabid 11 14.147, 43.517 KC818731 KC818882 KC818789 - KF647603 KF647614
Hemidactylus ulii sp. n. JS45 not collected Yemen Al Hababi 12 13.333, 43.722 KC818728 KC818878 - - - KF647612
Hemidactylus ulii sp. n. JS46 NMP6V 74833/1 Yemen Al Hababi 12 13.333, 43.722 KC818728 KC818879 KC818789 - - KF647613
Hemidactylus ulii sp. n. JS47 NMP6V 74833/2 Yemen Al Hababi 12 13.333, 43.722 KC818729 KC818880 KC818789 KC818942 KC819001 KC819061
Hemidactylus ulii sp. n. JS37 NMP6V 74832/1 Yemen 3 km S of Najd an Nashamah 13 13.358, 43.957 KC818727 KC818876 KF647578 KC818943 - KF647611
Hemidactylus ulii sp. n. JS38 NMP6V 74832/2 Yemen 3 km S of Najd an Nashamah 13 13.358, 43.957 KC818727 KC818877 KC818789 KF647593 - KF647614
Hemidactylus ulii sp. n. JS32 NMP6V 74835 Yemen 35 km W of Lahij 14 13.032, 44.558 KC818726 KC818875 KC818788 KC818941 KC819000 KC819060
Hemidactylus ulii sp. n. BJ09 NHM-BS N41916 Yemen Radman 15 14.1, 45.283 KF647572 - KF647577 KF647592 - KC819059
Hemidactylus ulii sp. n. JS17 NMP6V 74831/1 Yemen Al Hadr 16 13.877, 45.8 KC818725 KC818874 KC818787 KC818940 KC818999 KC819059
Hemidactylus ulii sp. n. JS18 NMP6V 74831/2 Yemen Al Hadr 16 13.877, 45.8 KC818725 - KC818789 - KF647604 KC819059
Hemidactylus angulatus JS123 NMP6V 74845/2 Ethiopia Arba Minch - 6.034, 37.564 KC818659 KC818807 KC818747 KC818903 KC818956 KC819018
Hemidactylus flaviviridis JS111 not collected Pakistan Okara - 30.811, 73.457 KC818676 KC818822 JQ957126 JQ957253 KC818965 KC819026
Hemidactylus flaviviridis JS113 not collected India Haridwar - 29.964, 78.201 KC818676 KC818823 JQ957126 JQ957253 KC818966 KC819027
Hemidactylus flaviviridis JS119 not collected Oman Jalan Bani Bu Hassan - 22.089, 59.278 JQ957119 JQ957183 KC818754 KC818911 KC818967 KC819028
Phylogenetic analyses and haplotype networks construction

The final dataset consisted of 36 ingroup individuals. Specimen numbers, localities, and GenBank accession numbers of all genes sequenced are presented in Table 1. The alignment of all concatenated genes was 4012 bp long. The software jModelTest 2.1.1 (Guindon and Gascuel 2003; Darriba et al. 2012) was used to assess the best-fitting model of nucleotide substitution for each gene separately under the Akaike information criterion [AIC, Akaike (1973)]. The best-fitting models were selected as follows: 12S – GTR+G; cytb – GTR+I+G; cmos – HKY+I; mc1r – TIM2+I; rag1 – HKY+I; rag2 – TrN+I). Phylogenetic analyses were performed using maximum likelihood (ML) and Bayesian inference (BI) methods. In order to detect the potential effect of the nuclear genes on the tree topology and nodal support, independent analyses were run on two datasets: (1) a dataset containing mtDNA genes only (12S, cytb), and (2) a concatenated dataset of all mtDNA and nDNA genes. Sequences of nuclear genes were not phased; heterozygous positions were coded according to the IUPAC ambiguity codes. Gaps were treated as missing data. Three specimens of Hemidactylus flaviviridis and one of Hemidactylus angulatus, representatives of two different clades of Hemidactylus (Carranza and Arnold 2006), were used to root the trees. Uncorrected genetic distances (p distances) were calculated in MEGA 5 (Tamura et al. 2011). Almost complete cytb sequences (1127 bp) of the new species from Yemen deposited in GenBank (Šmíd et al. 2013) were used to calculate p distances within this species, whereas an alignment of 307 bp was used to obtain intraspecific p distances within Hemidactylus saba and the new species from Saudi Arabia and Sinai, and also interspecific p distances between these three species.

Maximum likelihood analyses of both datasets were performed in RAxML 7.0.3 (Stamatakis 2006) using raxmlGUI (Silvestro and Michalak 2012) graphical extension with parameters estimated independently for each partition, GTR+I+G model of nucleotide evolution and a heuristic search with 100 random addition replicates. Support of the tree nodes was assessed by bootstrap analysis with 1000 pseudoreplications (Felsenstein 1985).

The BI analyses were run in MrBayes 3.2.1 (Ronquist et al. 2012). Appropriate equivalents of the best-fitting models were specified to each partition (gene) and all parameters were unlinked across partitions. Analyses were performed with two runs and four chains for each run for 107 generations, with sampling interval of 1000 generations. Appropriate sampling was confirmed by examining the stationarity of log likelihood (lnL) values and the value of average standard deviations of the split frequencies. Convergence between two simultaneous runs was confirmed by the PSRF (potential scale reduction factor) value. From 104 sampled trees, 25% were discarded as a burn-in and a majority-rule consensus tree was produced from the remaining ones, with posterior probabilities (pp) of each clade embedded. Nodes with ML bootstrap values ≥ 70% and pp values ≥ 0.95 were considered highly supported (Huelsenbeck and Rannala 2004).

Heterozygous positions in nuclear genes were identified based on the presence of double peaks in chromatograms and using the Heterozygote Plugin in Geneious. For the purpose of haplotype network construction, haplotypes from sequences with more than one heterozygous position were resolved in PHASE 2.1.1 (Stephens et al. 2001). Input data for PHASE were prepared in SeqPHASE (Flot 2010). In order to include as much data as possible, sequences of all Hemidactylus species from the Arid clade used in our previous study (Šmíd et al. 2013) were combined with the newly produced sequences and phased together (data not shown). In the case of rag1, the original alignment was trimmed to 846 bp, the length at which sequences of all individuals did not contain any N ends that would give misleading results in the allele reconstruction (Joly et al. 2007). PHASE was run under default settings except the probability threshold, which was set to 0.7. Haplotype networks of the four nuclear markers (cmos, mc1r, rag1, rag2) were drawn using TCS 1.21 (Clement et al. 2000) with 95% connection limit.

Material for morphological analyses

Material for morphological comparison included 225 specimens of 8 Hemidactylus species and one subspecies (Appendix) and was obtained from the following collections: National Museum Prague, Czech Republic (NMP); Natural History Museum in Braunschweig, Germany (NHM-BS); Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt, Germany (SMF); Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK); Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy (MSNG); Museo Civico di Storia Naturale di Milano, Milano, Italy (MSNM); Museo Civico di Storia Naturale, Carmagnola, Italy (MCCI); Università di Firenze, Museo Zoologico “La Specola”, Firenze, Italy (MZUF); British Museum of Natural History, London, UK (BMNH); California Academy of Sciences, San Francisco, USA (CAS); Taif University Zoological Collection, Taif, Saudi Arabia (TUZC); Institute of Evolutionary Biology Collection, Barcelona, Spain (IBES); Tomas Mazuch private collection, Dříteč, Czech Republic (TMHC); L. Kratochvíl collection (JEM); J. Šmíd collection (JS); Sherif Baha El Din private collection, Cairo, Egypt (SMB). Names of localities and governorates are spelled according to Google Earth (http://www.google.com/earth/). All coordinates are in WGS84 geographic coordinate system. Table of localities in a CSV text format and high-resolution photographs of all individuals analyzed in this study (397 pictures in total) have been deposited in MorphoBank (Project 1006; http://www.morphobank.org).

Morphological characters

The following measurements were taken with Powerfix digital calliper to the nearest 0.1 mm: snout-vent length (SVL), measured from tip of snout to vent; head length (HL), measured from tip of snout to retroarticular process of jaw; head width (HW), taken at the widest part of the head; head depth (HD), maximum depth of head; left eye diameter (E), measured horizontally; axilla-groin distance (AG), measured from posterior end of front limb insertion to anterior end of hind limb insertion; tail length (TL), measured from vent to tip of original tail. In addition to these metric characters, the following meristic characters were examined using a dissecting microscope: number of upper and lower labials (left/right); contact of nasals; number of infralabials in contact with first postmentals; mutual position of first postmentals; number of longitudinal rows of enlarged dorsal tubercles; number of lamellae under the first and fourth toe including unpaired proximal ones; and number of preanal pores in males. Terminology and diagnostic characters follow Moravec and Böhme (1997) and Moravec et al. (2011).


Phylogenetic analyses of both datasets resulted in trees presented in Fig. 2. Tree topology remains congruent with that showed in Šmíd et al. (2013). The three species form a well-supported monophyletic group (mtDNA: ML bootstrap 85/ Bayesian pp 1; mtDNA + nDNA: 100/1) to which we will refer to as the ‘Hemidactylus saba species group’ [support of individual species: Hemidactylus saba (100/1; 100/1), Hemidactylus sp. 1 from Sinai and Saudi Arabia (100/1; 100/1), Hemidactylus sp. 4 from Yemen (83/1; 100/1)]. The performed analyses did not resolve the topology within this species group despite the inclusion of more individuals and additional genetic data in comparison with previous works (Moravec et al. 2011; Šmíd et al. 2013). Therefore, with the current knowledge, this group remains polytomic. There is no genetic variability within Hemidactylus saba (all three specimens analyzed originate from the same locality) in both of the studied mtDNA genes and a very little variability in nDNA (mc1r and rag1 only) (Fig. 3). The species from Sinai and Saudi Arabia also shows very little variation in mtDNA (intraspecific p distance max. 1.3% in both 12S and cytb), but it varies in sequences of all the nDNA genes studied (Fig. 3). On the other hand, the unnamed Hemidactylus from Yemen exhibits relatively deep intraspecific differentiation into three well supported lineages. Uncorrected genetic distances between these lineages are up to 6.3% in cytb and up to 4.2% in 12S (Fig. 2). Moreover, the nDNA genes show a high level of genetic differentiation (Fig. 3). Intra- and interspecific genetic distances in both mtDNA genes analyzed between all three species are shown in Fig. 2. The results of the nuclear networks indicate that all alleles for all four independent loci are specific for each species.

Figure 2.

Maximum likelihood trees of mtDNA and mtDNA + nDNA datasets of the ‘Hemidactylus saba species group’. ML bootstrap values/Bayesian posterior probabilities are indicated by the nodes. Hemidactylus flaviviridis and Hemidactylus angulatus were used as outgroups. At the sides, schematic networks showing intra- and interspecific uncorrected p distances (in %) in the sequences of 12S and cytb. * intraspecific distances within Hemidactylus ulii sp. n. are based on an alignment of 1127 bp, all other values for cytb are calculated for an alignment of 307 bp.

Figure 3.

Nuclear allele networks of the four loci analyzed (cmos, mc1r, rag1, rag2). Circle sizes are proportional to the number of alleles. Small white circles represent mutational steps. Position of alleles BJ09a and BJ09b in the mc1r network is indicated by dashed lines because the sequence of the sample BJ09 (voucher NHM-BS N41916) was 108 bp shorter than the rest of the alignment and haplotype network reconstructions based on both 666 bp and 558 bp alignments linked these alleles to JS32b and JS32a, respectively.

The results of the molecular analyses, together with a unique combination of morphological features (see below) confirm the earlier conclusion that the newly recognized Hemidactylus sp. 1 and Hemidactylus sp. 4 represent two separate species, whose taxonomy and nomenclature need to be resolved.

Redescription of Hemidactylus granosus Heyden, 1827


Figs 4, 5
Hemidactylus granosus Heyden, 1827: p. 17; tab. 5, fig. 1. Lectotype SMF 8723 designated by Mertens (1967); collected by E. Rüppell 1827.
Hemidactylus turcicus (Linnaeus, 1758) – Boettger (1893: 29; part.); Anderson (1898: 80; part.); Salvador (1981: 84; part.); Baha El Din (2006: 66; part.).
Hemidactylus turcicus turcicus (Linnaeus, 1758) – Loveridge (1947: 143; part.); Mertens and Wermuth (1960: 79; part.); Baha El Din (2005: 19; part.); Mertens (1967: 55).
Hemidactylus verrucosus (Cuvier, 1829 [corr. Hemidactylus verrucosus Gray, 1831]) – Rüppell (1845: 300; part.).
Hemidactylus sp. 1 – Moravec et al. (2011: 24); Carranza and Arnold (2012: 17); Šmíd et al. (2013: 3).
Terra typica (Heyden 1827): “Egypten, Arabien, und Abyssinien”.
Terra typica restricta [by lectotype designation by Mertens (1967)]: “Arabia petraea” = Sinai, Egypt.
Material examined.

SMF 8723 (lectotype, adult male), Petr. Arabica [Arabia petraea], collected by E. Rüppell in 1827 (MorphoBank M305565M305594); NMP6V 70163/1 (adult female, MorphoBank M305520M305528), NMP6V 70163/2 (adult male, MorphoBank M305529M305542), NMP6V 70163/3–4 (adult females, MorphoBank M305543M305554, M305555M305564), Egypt, South Sinai governorate, Sharm el-Sheikh (27.885°N, 34.317°E), ca. 30 m a.s.l., collected by R. Kovář and R. Víta in 1996; ZFMK 94084, ZFMK 94085 (adult females, MorphoBank M305744M305760, M305761M305775), Saudi Arabia, Tabuk province, Al Wajh (26.2076°N, 36.4976°E), 5 m a.s.l., 31. V. 2012; ZFMK 94086 (adult female, MorphoBank M305778M305791), ZFMK 94088, ZFMK 94089 (adult males, M305793M305799, M305807, M305822M305827, M305828M305841), Saudi Arabia, Tabuk province, 15 km S of Al Wajh (26.1226°N, 36.5689°E), 25 m a.s.l., 31. V. 2012; TUZC-R10 (adult female, MorphoBank M305728M305743), Saudi Arabia, Hail province, 180 km N of Hail (26.8831°N, 40.0874°E), 1020 m a.s.l., 30. V. 2012; IBES10183, TUZC-R11 (adult males, MorphoBank M305656M305671, M305688M305701), ZFMK 94090, IBES10344 (adult females, MorphoBank M305672M305687, M305702M305717), Saudi Arabia, Makkah province, 30 km NE of Alhawiyah (21.6244°N, 40.7094°E), 1295 m a.s.l., 28. V. 2012; IBES10150, IBES10363 (adult males, MorphoBank M305615M305628, M305643M305655), ZFMK 94091 (adult female, MorphoBank M305629M305642), Saudi Arabia, Makkah province, 20 km S of Ashayrah (21.6022°N, 40.6911°E), 1316 m a.s.l., 28. V. 2012. All Saudi specimens were collected by M. Shobrak, S. Carranza and T. Wilms.

Referred material.

SMB 10660, Egypt, Suez governorate, Ayoun Musa (29.875°N, 32.649°E), ca. 12 m a.s.l., collected by S. Baha El Din, date unknown; TUZC-R9, Saudi Arabia, Tabuk province, 72 km N of Umluj (25.614°N, 36.9867°E), 19 m a.s.l., 31. V. 2012; IBES10001, Saudi Arabia, Riyadh province, Al Ghat (26.0545°N, 45.0003°E), 776 m a.s.l., 29. V. 2012; ZFMK 94087, TUZC-R8, Saudi Arabia, Tabuk province, 15 km S of Al Wajh (26.1226°N, 36.5689°E), 25 m a.s.l., 31. V. 2012; ZFMK 87236, Saudi Arabia, Makkah province, Taif National Wildlife Research Center (21.25°N, 40.96°E), 25. VI. 2007 by T. Wilms. These specimens were used for the molecular analyses only.

Status and nomenclature.

Heyden (1827) described Hemidactylus granosus as a new species occurring in Egypt, Arabia and Abyssinia (Ethiopia and Eritrea). Although not explicitly mentioned by the author, the description was apparently based on four specimens collected by Rüppell currently deposited in the Senckenberg Naturmuseum Frankfurt (collection numbers SMF 8723–8726). Heyden did not diagnose the new species against Hemidactylus turcicus (Linnaeus, 1758) and in respect to our today’s knowledge on the morphological variation in Hemidactylus the description of Hemidactylus granosus is very general. Traditionally, Hemidactylus turcicus has been considered a common species widely distributed across the Mediterranean and the Middle East. As the general diagnostic characters of Hemidactylus granosus given by Heyden (1827) were also applicable to Hemidactylus turcicus at that time, the name Hemidactylus granosus Heyden, 1827 was considered its junior synonym (e.g. Boulenger 1885, Loveridge 1947, Mertens and Wermuth 1960, Mertens 1967, Salvador 1981, Baha El Din 2006).

Recent examination (by JŠ) of four specimens collected by Rüppell (SMF 8723–8726) has shown that one of them [SMF 8723 designated by Mertens (1967) as lectotype of Hemidactylus granosus; for description see below] corresponds morphologically to Hemidactylus sp. 1 from Sinai. The other three specimens from this series morphologically correspond to Hemidactylus robustus Heyden, 1827 (SMF 8725, 8726) and Hemidactylus cf. granosus (SMF 8724), an animal superficially resembling Hemidactylus granosus but differing from the members of the ‘Hemidactylus saba species group’ in several important characters (see below). These findings lead to the conclusion that Hemidactylus granosus Heyden, 1827 is a valid taxon and needs to be resurrected from the synonymy of Hemidactylus turcicus. In the light of current knowledge, the range of Hemidactylus turcicus does not include a large part of Egypt, being restricted mostly to northern Egypt including Sinai and its Red Sea coast. The species is also missing in Arabia (sensu lato) and Ethiopia (Carranza and Arnold 2006; Moravec et al. 2011; Rato et al. 2011; Šmíd et al. 2013).


Hemidactylus granosus is a member of the ‘Hemidactylus saba species group’ within the Arabian radiation of the Arid clade as evidenced by the mtDNA and nDNA analyses. The species has the following combination of molecular and morphological characters: (1) Uncorrected genetic distance from Hemidactylus saba: 9.9–10.2% in 12S, 14.5–15.5% in cytb; from Hemidactylus sp. 4: 10.2–12.3% in 12S, 11.2–13.5% in cytb; (2) small size, SVL 39.0–53.2 mm in males, 40.6–53.3 mm in females; (3) rather elongated head, head length 24–28% of SVL, head width 68–86% of head length, head depth 33–47% of head length; (4) tail length 107–130% of SVL; (5) uppermost nasals separated by a small shield in 89% of specimens; (6) large anterior postmentals in wide mutual contact, and always in contact with the 1st and 2nd lower labial; (7) 9–11 upper labials; (8) 7–9 lower labials; (9) 14–15 longitudinal rows of enlarged, subtriangular, distinctly keeled dorsal tubercles; (10) 7–8 lamellae under the 1st toe and 10–13 under the 4th toe; (11) ca. 6–8 tail segments bearing 6 pointed tubercles; (12) 4–7 preanal pores in males forming a continuous row on the left and right side; (13) subcaudals enlarged; (14) in life, dorsum pale buff with dark brown spots tending to form transverse bands or X-shaped markings, dark horizontal stripe in prefrontal and temporal region, tail with ca. 10–13 dark brown transverse bands, venter white.

Description of the lectotype.

SMF 8723, adult male [erroneously determined as female by Mertens (1967)]. Head and body moderately depressed (Fig. 4). Upper labials (10/10), lower labials (8/7). Nostril between rostral, three subequal nasals and in punctual contact with first upper labial. Uppermost nasals separated by a small inserted scale. Mental triangular, as long as wide. Anterior postmentals long, in a broad contact with each other, both in contact with the 1st and 2nd lower labial reaching in about one fourth of the width of the 2nd labial. Second postmentals almost round, touching only the 2nd lower labial (Fig. 5). Two enlarged scales behind each second postmental, the lateral ones in contact with the 3rd lower labial. Eye moderate (E/HL=0.26). Head long, distinctly separated from body by a slender neck. Crescent-shaped ear opening. Interorbital region, crown of head and temporal area above the level of ear opening covered by round smooth tubercles. Dorsal region of the specimen is slightly scarred so it is not possible to count the enlarged tubercles on both sides precisely, but there are seven longitudinal rows of large, keeled and caudally pointed tubercles on the left side from which we infer there were originally 14 rows on both sides together. Lower arms, thighs and lower legs with prominent tubercles without keels. Tail original with 6 segments bearing 6 pointed tubercles, broken into three pieces, subcaudals enlarged from just after the hemipenial bulges. Lamellae under the 1st toe 7/7, lamellae under the 4th toe 11/11. Four preanal pores in a continuous row. No femoral pores or enlarged femoral scales. Colour (in alcohol) faded due to long fixation.

Figure 4.

Male lectotype of Hemidactylus granosus (SMF 8723) from Sinai, Egypt. General habitus, lateral and ventral view of the head, precloacal region with preanal pores, right hind leg. Scale refers to the uppermost picture only.

Figure 5.

Schematic drawing of the chin region of the lectotype and a new specimen from Sinai of Hemidactylus granosus, the holotype of Hemidactylus ulii sp. n., and Hemidactylus turcicus from Sinai.

Measurements (in mm): SVL 51.5, HL 12.9, HW 9.8, HD 6.0, E 3.3, AG 23.7.

Paralectotype SMF 8724 differs from other individuals of Hemidactylus granosus in having relatively high head (HD 50% of HL), lower number of lower labials (6), uppermost nasals in wide contact, first postmentals in contact with 1st lower labials, and 2 preanal pores.


Hemidactylus granosus can be distinguished from other member of the ‘Hemidactylus saba species group’ and from other congeners distributed in Sinai and the Red Sea coast by the following set of characters (see also Table 2).

Table 2.

Morphological comparison among members of the ‘Hemidactylus saba species group’ and with other Hemidactylus species from Sinai and SW Yemen. The values are given as follows: sample size, mean ± standard deviation above, min. – max. value below.

Species / Character Hemidactylus saba species group Hemidactylus robustus Hemidactylus turcicus Hemidactylus mindiae Hemidactylus jumailiae Hemidactylus yerburii yerburii Hemidactylus yerburii montanus
Hemidactylus granosus Hemidactylus saba Hemidactylus ulii sp. n.
Upper labials 18 9.4 ± 0.5 3 9.3 ± 0.8 10 9.3 ± 0.8 27 9.4 ± 0.7 33 8.2 ± 0.5 5 10.8 ± 0.8 18 9.8 ± 0.7 51 10.3 ± 0.7 57 10.2 ± 0.7
9–11 8–10 8–10 8–11 7 - 10 10 - 12 8–12 9–12 8–12
Lower labials 18 7.4 ± 0.4 3 7.7 ± 0.6 10 8.0 ± 0.6 27 7.7 ± 0.6 33 6.7 ± 0.5 5 8.1 ± 0.4 18 8.2 ± 0.6 51 7.9 ± 0.5 57 7.8 ± 0.6
7–9 7–8 7–9 6–9 6–8 7–9 7–10 6–9 6–10
Nasals in contact (%) 18 11 3 33.3 10 40 27 22.2 33 21.2 5 0 18 5.5 51 7.8 57 5.3
1st postmental in contact with 2nd lower labial (%) 18 100 3 33.3 10 100 27 70.3 33 12.1 5 80 18 83.3 51 98 57 89.5
Rows of dorsal tubercles 18 14.1 ± 0.2 3 14 ± 0.0 10 14.1 ± 1.0 27 14.8 ± 1.2 33 13.8 ± 0.7 5 12.4 ± 0.9 15 14 ± 1.4 46 15.3 ± 1.1 53 15.2 ± 1.2
14–15 14–14 12–16 13–18 12–16 12–14 12–16 13–18 12–18
Pores 8 5.6 ± 1.1 1 6 2 8 ± 0.0 9 6.1 ± 0.8 13 7.2 ± 1.4 1 4 9 7.2 ± 1.1 23 13.7 ± 2.2 27 11.2 ± 1.1
4–7 8 - 8 5–8 6–10 6–9 10–18 9–13
Lamellae under 1st toe 18 7.4 ± 0.5 3 8.2 ± 0.3 10 5.4 ± 0.5 27 6.1 ± 0.5 32 6.5 ± 0.5 5 6.2 ± 0.3 18 6.9 ± 0.7 51 6.7 ± 0.4 57 6.3 ± 0.4
7–8 8–9 5–6 5–8 6–7 6–7 6–8 6–8 5–7
Lamellae under 4th toe 18 11.5 ± 0.7 3 11.2 ± 0.3 10 8.6 ± 0.5 27 10.1 ± 0.7 32 9.7 ± 0.6 5 10 ± 0.0 18 10.9 ± 0.8 51 10.4 ± 0.6 57 10.2 ± 0.5
10 - 13 11–12 8–9 8–12 8–11 10–10 9–12 9–12 9–11
SVL (males) 8 46.8 ± 5.9 1 58.3 2 38.6 ± 2.6 8 41.8 ± 2.3 13 46.0 ± 5.8 1 49.3 8 48.4 ± 4.1 23 58.5 ± 7.1 25 56.5 ± 5.7
39.0–53.2 36.8–40.4 37.0–43.7 37.3–54.1 40.0–54.2 43.6–74.9 45.2–65.3
SVL (females) 10 49.0 ± 3.5 2 53.5 ± 7.9 2 40.1 ± 0.9 16 43.6 ± 4.7 18 49.2 ± 5.1 4 46.2 ± 11.4 8 48.6 ± 3.3 23 55.7 ± 5.3 30 52.6 ± 5.1
40.6–53.3 47.9–59.1 39.4–40.7 32.7–50.1 39.4–56.2 35.6–56.6 43.1–54.0 43.6–62.1 42.4–64.1

From Hemidactylus saba by having distinctly keeled dorsal tubercles (smooth in Hemidactylus saba), and lower number of lamellae under the 1st toe (7–8 vs. 8–9).

From Hemidactylus sp. 4 (described below) by its larger size (max. SVL 53.2 mm vs. 40.4 mm in males, 53.3 mm vs. 40.7 mm in females), in having more frequently separated uppermost nasals (100% vs. 60% of specimens), lower number of preanal pores in males (4–7 vs. 8), and higher number of lamellae under the 1st (7–8 vs. 5–6) and 4th (10–13 vs. 8–9) toe.

From Hemidactylus flaviviridis by its smaller size (max. SVL 53.2 mm in males and 53.3 mm in females vs. up to 90 mm [Anderson (1999); sexes not distinguished]), by the presence of enlarged dorsal tubercles, and the absence of femoral pores in males.

From Hemidactylus mindiae by the lower number of supralabials (9–11 vs. 10–12), by having anterior postmentals in wide contact (punctual in Hemidactylus mindiae) and keeled dorsal tubercles (smooth in Hemidactylus mindiae).

From Hemidactylus robustus by the larger size of males (max. SVL 53.2 mm vs. 43.7 mm), longer tail (tail length 53.0–64.8 mm vs. 40.9–48.7 mm), and lower number of preanal pores in males (4–7 vs. 5–8).

From Hemidactylus turcicus by its higher number of upper labials (9–11 vs. 7–10), in having anterior postmentals more frequently in contact with 2nd lower labial (100% vs. 12.1%), in having anterior postmentals in wide mutual contact behind the mental scale (contact punctual in 67% specimens of Hemidactylus turcicus), and by the lower number of preanal pores in males (4–7 vs. 6–10).


Specimens with intact tail vary in number of tail segments bearing 6 pointed tubercles (7–8). The original portion of the tail of the female NMP6V 70163/4 is very wide at the base, separated from cloacal region by a basal constriction. One specimen (IBES10212) is the only animal with 15 longitudinal rows of enlarged tubercles. Another one (IBES10284) has uppermost nasals in wide contact. Most striking is the variation in the number of preanal pores in males. Whereas the lectotype and the only male from Sinai (NMP6V 70163/2) have both 4 pores, all males from Saudi Arabia have 6–7 pores. There seems to be clinal variability in this character, males from NW of the known range (Fig. 6) possess only 4 preanal pores, all animals from the eastern Red Sea coast in Saudi Arabia have 6 pores and a single individual from the southern limit of the range has 7 pores.

Figure 6.

Distribution map of Hemidactylus granosus, Hemidactylus saba and Hemidactylus ulii sp. n. For the list of locality names and their corresponding numbers in the map see Table 1.

Coloration (in life) pale buff dorsally (Fig. 7). Conspicuous dark brown horizontal stripe in loreal and temporal area, terminated at the level of ear from where it continues in a series of dark patches on the neck. Four barely visible X-shaped markings on dorsum formed mainly by dark brown enlarged tubercles (first on nape, second across scapulae, third in lumbal region, and fourth just in front of the anterior insertion of hind limbs). Isolated dark brown stripe runs across body in the place of posterior insertion of hind limbs. Regenerated tails are uniformly buff from above. Dorsum, sides of chin, underside of front and hind limbs and underside of tail with faint stipple visible under magnification. Belly white. Tips of fingers and toes black behind insertion of terminal phalanges. Coloration is consistent among all specimens and varies only in distinctness of the markings.

Figure 7.

Live specimens of Hemidactylus granosus from Saudi Arabia. A IBES10344, 30 km NE of Alhawiyah (loc. number 8) B TUZC-R10, 180 km W of Hail (6) C ZFMK 94091, 20 km S of Ashayrah (9) D ZFMK 94086, 15 km S of Al Wajh (4).

There is a very low variation in mtDNA between specimens from Sinai and Saudi Arabia (max. 1.3% in both 12S and cytb). All animals from Sinai share the same haplotypes in 12S and also cytb gene. All four nuclear loci studied show some degree of intraspecific variation (Fig. 3).

Distribution and ecology.

Eduard Rüppell collected the original series in 1827 when he began his marine biological studies of the Red Sea and travelled from Egypt to Eritrea. There is no specific information that he went to Arabia as well (Rüppell 1826–1828; Klausewitz 2002; Wagner 2008); therefore the original distribution of Hemidactylus granosus described as “Egypt, Arabia, and Abyssinia [Ethiopia and Eritrea]” by Heyden (1827) was probably too general and incorrect. Because there were no other specimens assignable with certainty to Hemidactylus granosus apart from the four individuals collected in Sinai (SMF 8723–8726, for their current status see ‘Status and nomenclature’ section) (Boettger 1893), one of which became the lectotype after Mertens’ (1967) designation, Sinai could be considered the only reliable locality for Hemidactylus granosus. Here, Hemidactylus granosus is also confirmed from two coastal localities in south and west Sinai and from coastal and inland regions in western and central Saudi Arabia (Fig. 6). Nevertheless, a wider distribution of the species along the Red Sea coast can be expected. According to Baha El Din (2005), Hemidactylus geckos inhabiting the interior lowland of Sinai and the Eastern Desert in Egypt stand out in having notably coarse scalation. Interestingly, the areas with occurrence of animals with coarse scalation correspond with the presence of individuals with low numbers of preanal pores (Baha El Din 2005), which is typical for the Sinai populations of Hemidactylus granosus.

In 1996, when the NMP specimens were collected, the locality in Sharm el-Sheikh was formed by a crop field supplied with drain water from nearby habitations. Geckos were found during the day under unused empty barrels and also inside buildings. Other species syntopic with Hemidactylus granosus in Sharm el-Sheikh were: Hemidactylus turcicus, Chalcides ocellatus (Forskål, 1775), Stenodactylus sthenodactylus (Lichtenstein, 1823), and Ptyodactylus hasselquistii (Donndorff, 1798) (R. Víta in litt, 2013). However, when visited again in 2010, the locality had changed dramatically (R. Víta in litt, 2013). The whole area was under heavy development and the irrigation channels had disappeared. The current conditions at the place are unknown to us. In 2011 JM surveyed a neighbouring urban area east of this locality. It was covered by a mosaic of tourist resorts and abandoned ruderal plots. In dry anthropogenic habitats (e.g. rubbish dumps, road ditches, old walls and buildings, abandoned construction sites, natural but heavily disturbed open areas, etc.) dominated two very abundant gecko species. Ptyodactylus hasselquistii occupied primarily various vertical surfaces whereas Cyrtopodion scabrum (Heyden, 1827) prevailed on the ground. Tropiocolotes nattereri Steindachner, 1901 was found in dry and relatively well-preserved natural places. Hemidactylus turcicus was occasionally encountered in more humid artificial habitats in parks and hotel gardens. Specimens from Saudi Arabia were mostly collected during the day inside concrete tunnels under roads. In some of the tunnels they were syntopic with Ptyodactylus hasselquistii. One specimen was also collected on the walls of the Taif National Wildlife Research Centre, where it was also syntopic with Ptyodactylus hasselquistii.

Hemidactylus ulii sp. n.



Figs 5, 7, 8
Hemidactylus turcicusRösler and Wranik (1998: 120; part.).
Hemidactylus sp. ‘OTU7’ – Busais and Joger (2011a: 27); Busais and Joger (2011b: 268); Carranza and Arnold (2012: 95).
Hemidactylus sp. 4 – Moravec et al. (2011: 25); Šmíd et al. (2013: 3).

NMP6V 74833/2, adult male (MorphoBank M305892M305902), Yemen, Ta’izz governorate, Al Hababi (13.333°N, 43.722°E), 463 m a.s.l.; collected by L. Kratochvíl, 28. X. 2007.


NMP6V 74833/1 (adult male, MorphoBank M305884M305891), same collecting data as holotype; NMP6V 74831/1–2 (one adult and one subadult female, MorphoBank M305854M305863, M305864M305870), Yemen, Abyan governorate, Al Hadr (13.877°N, 45.8°E), 1151 m a.s.l., collected by L. Kratochvíl on 22. X. 2005; NMP6V 74832/1–2 (two subadult females, MorphoBank M305871M305875, M305876M305883), Yemen, Ta’izz governorate, ca. 3 km S of Najd an Nashamah by road (13.358°N, 43.957°E), 1182 m a.s.l., collected by L. Kratochvíl on 26. X. 2007; NMP6V 74834/1–2 (one adult and one subadult female, MorphoBank M305903M305911), Yemen, Dhamar governorate, Wadi Zabid (14.147°N, 43.517°E), 292 m a.s.l., collected by L. Kratochvíl on 29. X. 2007; NHM-BS N41916 (juvenile, MorphoBank M305842M305852), Yemen, Al Bayda’ governorate, Radman (14.1°N, 45.283°E), collected by W. Mustafa on 13. XI. 2007.

Referred material.

NMP6V 74835 (juvenile), Yemen, Lahij governorate, wadi 35 km W of Lahij (13.032°N, 44.558°E), 297 m a.s.l., collected by L. Kratochvíl on 25. X. 2007; JEM476 (juvenile), same collecting data as holotype; All juvenile specimens were used for comparison of meristic characters and included in the molecular analyses.


A small species of the ‘Hemidactylus saba species group’ withinthe Arabian radiation of the Arid clade of Hemidactylus, as evidenced by the mtDNA and nDNA analyses. The new species is characterized by the following combination of molecular and morphological characters: (1) Uncorrected genetic distances from Hemidactylus saba: 9.9–10.7% in 12S, 13.5–14.9% in cytb; from Hemidactylus granosus: 10.2–12.3% in 12S, 11.2–13.5% in cytb; (2) small size with a maximum recorded SVL 40.7 mm (36.8–40.4 mm in males, 39.4–40.7 mm in females); (3) moderately robust head, head length 28–30% of SVL, head width 70–75% of head length, head depth 37–46% of head length; (4) tail length 116% of SVL (only 1 specimen with intact tail); (5) uppermost nasals separated by a small shield (60% specimens) or in wide contact (40%); (6) large anterior postmentals in wide mutual contact in 90% of individuals, and in contact with the 1st and 2nd lower labial (scarcely and unilaterally with the 1st lower labial only); (7) 8–10 upper labials; (8) 7–9 lower labials; (9) dorsum with 12-16 longitudinal rows of enlarged, slightly keeled, conical tubercles; (10) 5–6 lamellae under the 1st toe and 8–9 lamellae under the 4th toe; (11) ca. 6–8 tail segments bearing 6 tubercles; (12) 8 preanal pores in one continuous row in males; (13) subcaudals enlarged; (14) in alcohol dorsum brownish grey with a pattern of more or less conspicuous dark transverse bands starting on the nape, tail with 9 dark brown transverse bands.


Hemidactylus ulii sp. n. can be distinguished from the other members of the ‘Hemidactylus saba species group’ and from all other congeners distributed in the region by the following combination of characters (see also Table 2):

From Hemidactylus granosus by its smaller size (max. SVL 40.4 mm vs. 53.2 mm in males, 40.7 mm vs. 53.3 mm in females), by having less frequently separated uppermost nasals (60% vs. 89% of specimens), higher number of preanal pores in males (8 vs. 4–7), and lower number of lamellae under the 1st (5–6 vs. 7–8) and 4th (8–9 vs. 10–13) toe.

From Hemidactylus saba by its smaller size (max. SVL 40.4 mm vs. 58.3 mm in males, 40.7 mm vs. 59.1 mm in females), higher number of preanal pores in males (8 vs. 6), and lower number of lamellae under the 1st (5–6 vs. 8–9) and 4th (8–9 vs. 11–12) toe.

From Hemidactylus flaviviridis by its smaller size (maximum SVL 40.4 mm in males, 40.7 mm in females vs. up to 90 mm [Anderson (1999); sexes not distinguished]), the presence of enlarged dorsal tubercles, and the absence of femoral pores in males.

From Hemidactylus jumailiae by its smaller size (max. SVL 40.4 mm vs. 54.2 mm in males, 40.7 mm vs. 54.0 mm in females), lower frequency of separated uppermost nasals (60% vs. 95%), in having conical and at least slightly keeled dorsal tubercles (vs. non-protruding and smooth tubercles), and lower number of lamellae under the 1st (5–6 vs. 6–8) and 4th (8–9 vs. 9–12) toe.

From Hemidactylus robustus by its smaller size (max. SVL 40.4 mm vs. 43.7 mm in males, 40.7 mm vs. 50.1 mm in females), and lower number of lamellae under the 4th toe (8–9 vs. 8–12).

From Hemidactylus sinaitus by the presence of enlarged tile-like subcaudals and in having separated uppermost nasals (60% vs. 9% of specimens).

From Hemidactylus yerburii montanus by its smaller size (maximum SVL 40.4 mm vs. 65.3 mm in males, 40.7 mm vs. 64.1 mm in females), lower number of preanal pores in males (8 vs. 9–13), and lower number of lamellae under the 4th toe (8–9 vs. 9–11).

From Hemidactylus yerburii yerburii by its smaller size (maximum SVL 40.4 mm vs. 74.9 mm in males, 40.7 mm vs. 62.1 mm in females), lower number of supralabials (8–10 vs. 9–12), lower frequency of having separated uppermost nasals (60% vs. 92%), lower number of preanal pores in males (8 vs. 10–18), and lower number of lamellae under the 1st (5–6 vs. 6–8) and 4th (8–9 vs. 9–12) toe.

Description of holotype.

NMP6V 74833/2, adult male. Body slightly depressed to cylindrical (Fig. 8). Upper labials 8/8, lower labials 7/7. Nostril between rostral, three nasals and in punctual contact with the first upper labial. Uppermost nasals separated by a small inserted shield. Mental almost triangular. Anterior postmentals large and very long, in wide mutual contact behind mental, in contact with the 1st lower labial (left) and the 1st and 2nd lower labials (right) (Fig. 5). Posterior postmentals smaller, in contact with the 1st and 2nd (left) and the 2nd (right) lower labial. Eye moderate (E/HL=0.24). Supraciliar granules with prominent projections, which form a comb-like structure above the eyes. Parietal and temporal region covered with round pointed regularly distributed tubercles. Ear opening oval. Dorsum with 14 longitudinal rows of enlarged, prominent, caudally pointed tubercles bearing distinct longitudinal keels. Thighs and lower legs with scattered enlarged tubercles. Tail partially regenerated from about half of its original length (estimate), original part relatively thick without basal constriction. Conical and keeled tail tubercles on tail segments forming regular whorls. Each whorl separated from the next one by four small scales. Subcaudals enlarged, tile-like. Regenerated part of the tail with small uniform scales without tubercles. Lamellae under the 1st toe 6/6, lamellae under the 4th toe 8/8. Eight preanal pores, no femoral pores or enlarged femoral scales.

Figure 8.

Holotype of Hemidactylus ulii sp. n. (NMP6V 74833/2, male) from Al Hababi, Yemen. General habitus, lateral and ventral view of the head, precloacal region with preanal pores, right hind leg. Scale refers to the uppermost picture only.

Measurements (in mm): SVL 40.4, HL 11.5, HW 8.6, HD 5.2, E 2.8, AG 16.2.

Coloration of holotype in preservative.

Overall dorsal coloration brownish grey. An indistinct dark horizontal stripe in loreal and temporal area. Seven dark brown transverse bands across the nape and body, the one in scapular region being the most conspicuous. Dark brown bands also on the original part of the tail. Belly whitish.


The paratypes (Fig. 9) differ from the holotype in the following features: number of upper labials 8–10; number of lower labials 7–9; four paratypes (NMP6V 74831/1, NMP6V 74832/1–2, NMP6V 748333/1) have uppermost nasals in wide contact; anterior postmentals in contact with 2nd lower labials on both sides (except of NMP6V 74832/1 where the arrangement is the same as in the holotype); longitudinal rows of enlarged tubercles 12–16; lamellae under the 1st toe 5–6, lamellae under the 4th toe 8–9. The intact tail of the paratype NMP6V 74833/1 has 7 segments bearing at least six enlarged spine-like tubercles and 9 dark brown transverse bands widening towards the tail tip.

Figure 9.

Four (out of eight) paratypes of Hemidactylus ulii sp. n. A NMP6V 74833/1, male B NMP6V 74834/1, female C NMP6V 74831/1, female D NMP6V 74832/1, subadult female.

Measurements of paratypes (in mm): NMP6V 74831/1: SVL 40.7, HL 11.5, HW 8.2, HD 4.9, E 3.0, AG 19.0; NMP6V 74831/2: SVL 32.0, HL 9.3, HW 6.6, HD 3.7, E 2.1, AG 12.7; NMP6V 74832/1: SVL 32.7, HL 9.7, HW 7.0, HD 3.4, E 2.3, AG 14.3; NMP6V 74832/2: SVL 32.9, HL 9.3, HW 6.7, HD 3.6, E 2.4, AG 13.5; NMP6V 74833/1: SVL 36.8, HL 10.7, HW 8.0, HD 4.5, E 2.4, AG 14.1, TL 42.5; NMP6V 74834/1: SVL 39.4, HL 11.1, HW 8.1, HD 4.4, E 2.7, AG 16.7; NMP6V 74834/2: SVL 32.0, HL 9.5, HW 6.7, HD 3.9, E 2.5, AG 13.8; NHM-BS N41916: juvenile, not measured.

As already mentioned (Results), the level of genetic variability within Hemidactylus ulii sp. n. is very high. The species is divided into three well supported sublineages which reflect the geographic origin of the samples. Although there is a certain geographic separation corresponding with these sublineages, the exact limits are not distinct and also morphological variation among paratypes is not congruent with geography.


The species epithet “ulii” is a patronym for Prof. Ulrich Joger, a German herpetologist known as Uli among friends, in recognition of his important contribution to the knowledge of the herpetofauna of the Western Palearctic.

Distribution and ecology.

Hemidactylus ulii sp. n. is known from inland mid-altitude areas (292–1182 m) of southwestern Yemen (Fig. 6). Most specimens were collected in open dry wadis with scattered rocks and boulders, in stony deserts and also in the vicinity of villages in gardens and irrigated cropland fields.

The following reptile specieswere found to occur in sympatry with Hemidactylus ulii: Bunopus spatalurus Anderson, 1901; Hemidactylus yerburii yerburii Anderson, 1895; Pristurus crucifer (Valenciennes, 1861); Pristurus flavipunctatus Rüppell, 1835; Pristurus rupestris Blanford, 1874; Ptyodactylus sp.; Tropiocolotes scorteccii Cherchi and Spano, 1963; Acanthodactylus sp.; Chamaeleo arabicus Matschie, 1893; Pseudotrapelus sinaitus (Heyden, 1827); Trapelus flavimaculatus Rüppell, 1835; and Pelomedusa subrufa (Bonnaterre, 1789).


Previous phylogenetic studies of the Arid clade of Hemidactylus disclosed an extraordinarily rich diversity within this genus in the Arabian Peninsula (Moravec et al. 2011; Carranza and Arnold 2012; Šmíd et al. 2013). The latter work, besides of showing the phylogenetic relationships among individual species of the Arid clade, highlighted the high level of genetic differentiation and existence of several yet undescribed taxa within this genus. The ‘Hemidactylus saba species group’ as defined herein represents one of the monophyletic groups within the Arabian radiation. All three species forming this group – Hemidactylus granosus, Hemidactylus saba, and Hemidactylus ulii sp. n.– are well defined and distinguishable both genetically and morphologically from each other, as well as from other Hemidactylus species that occur in the same area. Geographically, Hemidactylus saba and Hemidactylus ulii sp. n. are confined to the foothills and submontane areas of southwestern Yemen, where they occupy mid-altitude elevations (292–1182 m in Hemidactylus ulii sp. n., 1180 m in Hemidactylus saba). In comparison, Hemidactylus granosus has a much wider distribution, spanning from northeastern Egypt to central Saudi Arabia. It was found from the sea-level up to almost 1600 m in the Asir Mountains, which stretch along the eastern Red Sea coast of the Arabian Peninsula. Its occurrence in eastern Egypt is also likely based on observations of Baha El Din (2005, 2006), who reported morphologically variable populations of Hemidactylus turcicus (sensu lato) in these regions attributable to Hemidactylus granosus (see Distribution and ecology). The distribution of Hemidactylus granosus in the coastal Sinai and Saudi Arabia near important marine junctions together with the genetic uniformity of this species indicates extensive gene flow between these populations. It may be the result of recent colonization event(s), their inadvertent human-mediated transportation or perpetual contact of populations in a continuous range. The continuous range of Hemidactylus granosus along the Hijaz and Asir Mountains in western Arabia confirms that these mountain ranges can serve as a corridor providing connection between the eastern Mediterranean and southern Arabia (Scott 1942; Gvoždík et al. 2010).

The highlands of southwestern Saudi Arabia and Yemen are known to host a high number of endemic taxa (Balletto et al. 1985; Arnold 1986; Gasperetti 1988; Harrison and Bates 1991; Gasperetti et al. 1993). The genus Hemidactylus also shows a high rate of speciation and endemicity in the area. Currently, there are eight species and one subspecies known from the Yemen highlands, which makes Hemidactylus one of the most specious reptile genera in the area (Fritz and Schütte 1987; Busais and Joger 2011b; Šmíd et al. 2013; Uetz 2013). As new genetic and morphological data are becoming available from Arabia even more new species are to be expected (Moravec et al. 2011; Šmíd et al. 2013), thus fulfilling the prognosis of Baha El Din (2005) and the models of Ficetola et al. (2013) which suggested that the Red Sea region is likely to contribute significantly to the diversity of Hemidactylus.


We thank the following curators for granting access to collections under their care: U. Joger (NHM-BS), G. Köhler and his assistant L. Acker (SMF), R. Sindaco and G. Boano (MCCI), G. Doria (MSNG), S. Scali (MSNM), A. Nistri (MZUF), J. Vindum (CAS), B. Clarke and E. N. Arnold (BMNH), and T. Mazuch. We are very indebted to R. Kovář and R. Víta for collecting the Sinai material of Hemidactylus granosus, to S. Baha El Din for providing tissue sample of specimen SMB 10660 of the same species and to J. Červenka for field assistance in Yemen. We are grateful to two anonymous reviewers for their helpful comments. The study was supported by the NAKI project of the Ministry of Culture of the Czech Republic (# DF12P01OVV021 MKČR to JŠ and JM), by grant CGL2012-36970 to SC from the Ministerio de Economía y Competitividad, Spain (co-funded by FEDER). We are thankful to the Deanship of academic research at Taif University for funding the sample collection in Saudi Arabia (Grant no. 1-433-2108) and to Omer Baeshen, Environment Protection Agency, Sana'a, Republic of Yemen for issuing the collecting permit (Ref 10/2007).

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Specimens examined

Hemidactylus flaviviridis (8 individuals) - NMP6V 74858 (Oman, Jalan Bani Bu Hasan); NMP6V 74859/1–5 (Pakistan, Multan); NMP6V 74856 (Pakistan, Rakhni); NMP6V 74857 (Pakistan, Sukkur)

Hemidactylus jumailiae (18 individuals) - NMP6V 74818/1 (Yemen, near Al Bayda [At Dageeg]); NMP6V 74819 (Yemen, Sana’a); NHM-BS N41788, NHM-BS N41890 (paratype), NHM-BS N41891, NHM-BS N41893 (holotype), NHM-BS N41894 (paratype), NHM-BS N41897 (paratype) (Yemen, Ibb); NHM-BS N41898 (paratype, the same number as one of Hemidactylus yerburii montanus paratypes, Busais and Joger 2011b), NHM-BS N41899 (paratype) (Yemen, Thamar); BMNH1982.1143–44 (Yemen, Al Nabi Shuaib, 30 Km W. of Sana’a); BMNH1982.1145 (Yemen, Sana’a); BMNH1982.1146 (Yemen, Wadi Ahger, 45 Km. W. of Sana’a); BMNH1952.1.3.52 (Yemen, Sana’a); MSNG-YEM02, MSNG-YEM03 (Yemen, El Menghil); MCCI-R814 (Yemen, Hababah)

Hemidactylus mindiae (5 individuals) - NMP6V 71323/1–2 (Jordan, Jabal Ghazali); NMP6V 72739/1–3 (Jordan, Wadi Ramm Nughra Radet Salem)

Hemidactylus robustus (27 individuals) - SMF 8720 (lectotype), SMF 8721 (“Abyssinia” [Ethiopia and Eritrea]); SMF 8725–8726 – redetermined from Hemidactylus granosus (Egypt, Sinai); JS210, TMHC2012.07.092, TMHC2012.07.100 (Ethiopia, Jijiga), CAS130512 – redetermined from Hemidactylus macropholis as it is in the CAS catalogue (Kenya, vicinity of Mandera); NMP6V 74820 (Iran, Bandar Lengeh); NMP6V 74821/1–2 (Yemen, Wadi Zabid); NMP6V 74829 (Yemen, Bir Ali); JS144 (Kenya, Garissa); NMP6V 74867/1–3 (Oman, Muscat); NMP6V 74868 (Oman, Salalah); NMP6V 74869/1–7 (Oman, Mughsayl); NMP6V 74870/1–2 (Oman, Shisr); MCCI–R815 (Yemen, Zabid)

Hemidactylus saba (3 individuals) - NHM-BS N41912 (holotype, MorphoBank M305478M305492), NHM-BS N41913 (paratype, MorphoBank M305493M305504), NHM-BS N41914 (paratype, MorphoBank M305505M305519) (Yemen, Marib)

Hemidactylus sinaitus (23 individuals) - BMNH82.8.16.27 (holotype, probably from Suakin, Sudan); BMNH97.10.28.83–85 (Sudan, Durrur, N of Suakin); BMNH97.10.28.87 (Sudan, Wadi Haifa); BMNH1974.3931 (Ethiopia, Mule River?, Danakil); BMNH1937.12.5.293–294 (Somalia, Borama district); BMNH95.5.23.7 (Yemen, Sheikh Osman, near Aden); BMNH1945.12.12.14 (Yemen, Bir Fadhl, Aden); NMP6V 74809/1–4 (Sudan, Wad Ben Naga); NMP6V 74810 (Sudan, 15 km SE Atbara); MZUF28645–646 (Yemen, Moka); MZUF10914, MSNM521 (Eritrea, Isola [island] Sheik-Said); MSNM523–524 (Eritrea, Ailet); CAS174021–022 (Sudan, Assalaya)

Hemidactylus turcicus (33 individuals) - NMP6V 34747 (Syria, Baniyas); NMP6V 34748/1–3 (Syria, Palmyra); NMP6V 34749 (Syria, Salkhad); NMP6V 70648/1–4 (Turkey, Kaş); NMP6V 70668 (Greece, Kastellorizo, St. Georgies); NMP6V 71056 (Egypt, Bahariya); NMP6V 71587/1–3 (Cyprus, Famagusta); NMP6V 71592/1–2 (Cyprus, Yali); NMP6V 72497 (Syria); NMP6V 74046/1–2 (Syria, Cyrrhus); NMP6V 74047/1–2 (Turkey, Antakya); NMP6V 74050 (Greece, Crete, Kavros); NMP6V 74131/1–3 (Syria, Palmyra); NMP6V 73626/1–3 (Turkey, Finike); NMP6V 70269 (Italy, Sardinia, Cagliari); NMP6V 72073 (Greece, Korfu, Nicos); NMP6V 74167 (Greece, Crete, Kavros); NMP6V 70667 (Greece, Kastellorizo); NMP6V 70163/5 (Egypt, Sharm el-Sheikh)

Hemidactylus yerburii yerburii (51 individuals) - NMP6V 74827/1–4 (Yemen, Jabel Habeshi); NMP6V 74825/1–2 (Yemen, Al Turbah); NMP6V 74826 (Yemen, N of Lahij, Wadi Tuban); NMP6V 74823/1–3 (Yemen, 14 km NW of Al Turbah); NMP6V 74824/1–2 (Yemen, 3 km S of Najd an Nashamah); NMP6V 74828/1–3 (Yemen, Al Hababi); NMP6V 74822/1–5 (Yemen, near Zinjubar); MSNG-YEM01 (Yemen, Ta’izz); MSNG-YEM05, MSNG-YEM06 (Yemen, Vahren); NHM-BS N41856–59, NHM-BS N41861–64, NHM-BS N41866, NHM-BS N41868–69, NHM-BS N41888 (Yemen, Tour Albaha); NHM-BS N41860 (Yemen, Lahij); NHM-BS N41871–72 (Yemen, Radfan); NHM-BS N41873 (Yemen, Shihr); NHM-BS N41875 (Yemen, Ariab); NHM-BS N41876–77, NHM-BS N41879–86 (Yemen, Lowder); NHM-BS N41887 (Yemen, Aden)

Hemidactylus yerburii montanus (57 individuals) - NMP6V 74802 (Yemen, Jabal Bura); NHM-BS N41751–52 (paratypes), NHM-BS N41758 (paratype), NHM-BS N41762–63, NHM-BS N41765–66, NHM-BS N41768–69, NHM-BS N41770 (paratype), NHM-BS N41772–74, NHM-BS N41779, NHM-BS N41783 (paratype), NHM-BS N41785 (paratype), NHM-BS N41791 (paratype), NHM-BS N41793 (paratype), NHM-BS N41797–800 (paratypes), NHM-BS N41802–06 (paratypes), NHM-BS N41807 (paratype), NHM-BS N41809 (paratype), NHM-BS N41811–15 (paratypes), NHM-BS N41818 (paratype), NHM-BS N41821 (paratype), NHM-BS N41823 (paratype), NHM-BS N41836 (holotype), NHM-BS N41839, NHM-BS N41840 (paratype), NHM-BS N41842 (paratype), NHM-BS N41843, NHM-BS N41844 (paratype), NHM-BS N41846, NHM-BS N41848, NHM-BS N41851–52, NHM-BS N41867 (paratype) (Yemen, Ibb); NHM-BS N41771 (paratype) (Yemen, Yareem); NHM-BS N41789–90 (Yemen, Thamar); NHM-BS N41833–34 (paratypes) (Yemen, Wadah); NHM-BS N41853–55 (paratypes) (Yemen, Sana’a).