Hemidactylus dawudazraqi, Moravec, Jiří, Kratochvíl, Lukáš, Amr, Zuair S., Jandzik, David, Šmíd, Jiří & Gvoždík, Václav, 2011

Moravec, Jiří, Kratochvíl, Lukáš, Amr, Zuair S., Jandzik, David, Šmíd, Jiří & Gvoždík, Václav, 2011, High genetic differentiation within the Hemidactylus turcicus complex (Reptilia: Gekkonidae) in the Levant, with comments on the phylogeny and systematics of the genus, Zootaxa 2894, pp. 21-38 : 28-32

Hemidactylus dawudazraqi sp. n.

Figs. 3 (A–B), 4 (A–E), 5 (D)

Hemidactylus turcica — Flower (1933). Incorrect subsequent spelling.

Hemidactylus turcicus turcicus — Werner (1971), Disi (1996, 2002), Moravec and Böhme (1997), Disi and Amr (1998), Disi et al. (1999, 2001, 2004), Carranza and Arnold (2006), Amr et al. (2007).

Hemidactylus turcicus lavadeserticus — Carranza and Arnold (2006).

Holotype. NMP 6 V 74134/1, adult male, Azraq, 31°49.770’N, 36°48.433’E, ca. 515 m a.s.l., Jordan, collected on 1–2 July 2006 by L. Kratochvíl, GenBank Acc. No. HQ833753 (Cytb).

Paratypes. NMP 6 V 74134/2–17, six adult males and ten adult females, the same locality and collecting data as the holotype; NMP 6 V 35541, subadult male, Azraq, 31°50’N, 36°49’E, ca. 510 m a.s.l., Jordan, collected on 16 May 1996 by J. Moravec; NMP 6 V 72130/1–3, Dair al Khaf, 32°19’N, 36°53’E, ca. 1120 m a.s.l., one adult male and two adult females, Jordan, collected on 3 June 2004 by D. Modrý; NMP 6 V 72131, subadult specimen, Jawa, 32°20’N, 37°02’E, ca. 935 m a.s.l., Jordan, collected on 4 June 2004 by D. Modrý; NMP 6 V 72740/1–2, two adult females, Jawa, 32°20’N, 37°02’E, ca. 935 m a.s.l., Jordan, collected on 27 June 2005 by M. Abu Baker and D. Modrý; NMP 6 V 70457, subadult specimen, Rashiedeh, 32°40’N, 36°50’E, ca. 1425 m a.s.l., Muhafazat of Sweida, Syria, collected on 14 May 1999 by J. Moravec

Referred material. NMP 6 V 70616, adult female, Azraq, 31°50’N, 36°49’E, ca. 510 m a.s.l., Jordan, collected on May 1997 by D. Modrý; NMP 6 V 74138/1–6, two adult females, four subadult specimens, Azraq, 31°49.770’N, 36°48.433’E, ca. 515 m a.s.l., Jordan, collected on 1–2 July 2006 by L. Kratochvíl; NMP 6 V 74135/1–7, five adult females and two subadult specimens, Wadi Mujib 31°26.023’N, 35°47.489’E, ca. 795 m a.s.l., Jordan, collected on 21–22 June 2006 by L. Kratochvíl; NMP 6 V 74136/1–7, five adult females and two subadult specimens, Little Petra 30°22’N, 35°27’E, ca. 1081 m a.s.l., Jordan, collected on 27 June 2006 by L. Kratochvíl; NMP 6 V 74137, adult male, Petra 30°19.318’N, 35°27.968’E, ca. 1013 m a.s.l., Jordan, collected on 28 June 2006 by L. Kratochvíl.

Diagnosis. A species of the Arid species group of Hemidactylus as revealed from mtDNA analyses, which can be distinguished by the following molecular and morphological characters: (1) diagnostic nucleotide substitutions in Cytb, from all other Levantine taxa in positions 28 A (adenine) → G (guanine), 29 T (thymine) → C (cytosine), 175 A → G, 176 C → A, 246 T → C, 426 C → A, 531 C → T, 564 T → C, 663 A → C, 792 C → A, 985 G → T (GenBank Acc. Nos. HQ833749 – HQ833758 ); (2) small size, SVL 40.1–47.8 mm in males, 41.4–49.9 mm in females; (3) robust head, head depth 44.9–56.4 % of head length, head width 74.3–90.7 % of head length; (4) long tail, tail length 119.8–140.9 % of SVL; (5) nasals separated by a small scale in 92 % of individuals; (6) large anterior postmentals in contact with 1st and less frequently also with the 2nd lower labials, both postmentals in contact with the 2nd lower labials in 8 %; (7) 8–11 upper labials; (8) 6-8 lower labials; (9) 12–15 rows of large, round, conical, slightly keeled, dorsal tubercles; (10) 6–7 lamellae under the 1st toe and 9–12 lamellae under the 4th toe; (11) 5–8 tail segments bearing 6 tubercles; (12) 6–8 preanal pores in males; (13) in life, dorsum pinkish or yellowish white to yellowish orange with a pattern of irregular light brown to orange brown crossbars, head with dark longitudinal streak in loreal and postocular area, tail with a conspicuous pattern of 9–11 dark brown to black transverse bands on yellowish white to white background.

Comparisons. The new species can be distinguished from other Levantine species of the Arid species group of Hemidactylus by following combination of characters (see also Table 3): from H. turcicus by smaller size (maximal size 47.8 mm vs. 54.1 mm in males and 49.9 mm vs. 56.2 mm in females), significantly longer tail relatively to SVL (TL 119.9–140,9 vs. 103.0–121.4 % of SVL) (ANCOVA, tail length as dependent variable, SVL as a covariate, species as factor; species: F (1, 17) = 14.456, p = 0.0014), higher number of lamellae under the 4th toe (9–12 vs. 8–11), and genetic divergence of 10.0 % in Cytb (uncorrected p -distances); from H. lavadeserticus by robust head and body (head depth 44.9–56.4 % vs. 35.0–47.0 % of head length), larger relative tail length (119.8–140.9 % vs. 114.1–117.4 % of SVL), low frequency of contact of both postmentals with the 2nd lower labials (8 % vs. 100 %), lower average number of lamellae under the 1st toe (6–7 vs. 7–8), larger and more prominent dorsal tubercles, higher number of tail segments bearing 6 tubercles (5-8 vs. 2–6), and genetic divergence of 11.1 % in Cytb; from H. mindiae by robust head and body (head depth 44.9–56.4 % vs. 33.9–47.3 % of head length), lower number of upper labials (8–11 vs. 10–13), low frequency of contact of both postmentals with the 2nd lower labials (8 % vs. 80 %), higher number of preanal pores in males (6–8 vs. 4–6), and genetic divergence of 8.4 % in Cytb.

Description of the holotype. Adult male ( Figs. 3 A–B), SVL 46.4 mm, head length 10.9 mm, head width 9.5 mm, head depth 6 mm, tail length 60.6 mm. Upper labials (left/right) 9/9, rows of dorsal tubercles 14, lamellae under the 1st toe 7/7, lamellae under the 4th toe 11/11, tail segments bearing six tubercles. Nostril surrounded by rostral, three subequal nasals and the 1st upper labial. Uppermost nasals separated by one smaller scale. Mental large, pentagonal and deeply impacted between anterior postmentals. Anterior postmentals large, nearly as long as wide, shorter than mental, in punctual contact behind the symphysial, in contact with the 1st lower labial (left) and the 1st and 2nd (punctually) lower labials (right). Posterior postmentals smaller, in contact with the 1st and 2nd lower labials (left) and the 2nd lower labial (right). Digits moderately dilated. Dorsal tubercles round, prominent, feebly keeled, in 14 longitudinal rows. Tail tubercles on the anterior six tail segments slightly larger and obviously keeled. Scales on underside of tail enlarged and imbricate. In alcohol, whitish gray dorsally, with five inconspicuous dark crossbars on the neck and body, and with nine dark transverse bands on tail.

Variation. As mentioned in the part on molecular phylogeny of H. turcicus (s. l.), the new species shows relatively high intraspecific genetic differentiation, forming at least four sublineages (N, W1, W2, and S; Fig. 2 .). In comparison with the population from southern Syria and northern Jordan (sublineage N), the animals from Wadi Mujib (sublineage W1) and Petra and Little Petra (sublineage S) have less robust head and body, relatively larger eyes and smaller and narrower dorsal and especially tail tubercles. The tendency towards depressed head and body and smaller dorsal and tail tubercles appears to be higher in sublineage S (comparative voucher specimens of sublineage W2 were not at our disposal). This variation could reflect differences in habitats of the individual H. dawudazraqi sublineages. Whereas the representatives of sublineage N were collected predominantly on the ground in open areas with stony or loamy-sandy substrates, populations belonging to sublineage W1 and especially sublineage S were associated with rocky areas, caves and rock crevices. Similarly, the new species displays a colour variation corresponding to the substrate character. Individuals from basalt areas (Jawa and Dair al Khaf; Fig. 4 C–D) have yellowish orange to orange brown colouration in contrast to the light pinkish to yellowish white ground colour of the specimens inhabiting light substrata ( Fig. 4 E).

Distribution and ecology. The known range of H. dawudazraqi reaches from southern Syria to southwestern Jordan ( Fig. 6 ). The northernmost locality lies ca. 20 km W of the type locality of H. lavadeserticus and the southernmost locality is situated ca 75 km N of the known Jordanian occurrence of H. mindiae . We can expect that the range of the new species probably covers wider areas of southern Syria and northern and central Jordan.

The type locality lies at the edge of the oasis Azraq, which is situated at the border between basalt lava areas of northern Jordan and stony to loamy-sandy desert of central Jordan. At this place, H. dawudazraqi was collected predominantly in open desert habitat characterised by light loamy-sandy substrate and scattered herbaceous and bush vegetation ( Fig. 4 F). Here, the adult and subadult specimens were frequently encountered on open ground by night. This terrestrial mode of life corresponds well with the find of a multiple egg clutch containing nine eggs of H. dawudazraqi deposited under a flat stone lying on the ground in an open arid area (L. Kratochvíl, pers. com., own obs.) and with the reports that the geckos were observed in deep horizontal burrows in association with termites of the family Hodotermitidae in the Azraq Nature Reserve ( Disi and Amr 1998, Disi et al. 1999). Rarely, the individual specimens of H. dawudazraqi were also collected on the walls of small houses at the periphery of the town of Azraq (a synantropic mode of life was also observed at the Syrian locality of Rashiedeh). Other reptiles found in sympatry with H. dawudazraqi included Mesalina brevirostris Blanford , M. guttulata ( Lichtenstein) , Trachylepis vittata (Olivier) , Trapelus pallidus agnetae (Werner) , Pseudotrapelus sinaitus werneri Moravec , Chamaeleo chamaeleon (Linneaus) , Spalerosophis diadema (Schlegel) , and three other species of geckos ( Bunopus tuberculatus Blanford ; Cyrtopodion scabrum (Heyden) and Stenodactylus grandiceps Haas ) were observed near Azraq (J. Moravec, L. Kratochvíl, V. Gvoždík, pers. obs.).

As mentioned in the chapter about variation, geckos from Wadi Mujib, Little Petra and Petra were predominantly rock dwellers looking for shelters in rock crevices and caves.

Etymology. The specific name is a patronym for our colleague and friend David Modrý in recognition of his important contributions to the knowledge of the Jordanian herpetofauna. The name is used in its Arabic form as a compound of Arabic Dawud (David) and Azraq (the name of the type locality meaning “Blue” in English and “Modrý” in Czech).

Figures

FIGURE 2. Maximum likelihood tree of H. turcicus and the Levantine taxa of Hemidactylus based on the complete mitochondrial Cytb (1137 bp) and 303 bp-long sequences from Carranza and Arnold (2006). Short sequences which did not possess unique haplotypes were only allocated into one of the two sublineages of H. turcicus. Letters in circles correspond to the geographic origin of the sublineages of H. dawudazraqi sp. n. Numbers above branches are ML bootstrap values and Bayesian posterior probabilities, if above 50 %. The tree was rooted by H. haitianus (HdC 1), H. cf. fasciatus (Hd 30), H. cf. yerburii (Hd 60), Hemidactylus sp. 2 (Hd 90) and Hemidactylus sp. 3 (Hd 59).

FIGURE 3. Holotype of Hemidactylus dawudazraqi sp. n. (NMP 6 V 74134 / 1), (A) dorsal view, and (B) detail of the head.

FIGURE 4. Hemidactylus dawudazraqi sp. n., (A) adult male from the type locality (uncollected), (B) detail of the head of the same specimen. (C) Subadult paratype of Hemidactylus dawudazraqi sp. n. (NMP 6 V 70457) from Rashiedeh (Syria). (D), adult female paratype of Hemidactylus dawudazraqi sp. n. (NMP 6 V 7213 / 2) from Dair al Khaf (Jordan). (E) Adult female of Hemidactylus dawudazraqi sp. n. from Wadi Mujib (Jordan) (uncollected). (F) Type locality of Hemidactylus dawudazraqi sp. n., Azraq (Jordan).

FIGURE 6. Schematic map showing distributions of individual Hemidactylus species and their forms in the Mediterranean and Levant as inferred from the molecular analyses (see also Figs. 1 and 2).