review of the taxonomy of the spiny-tailed lizards of arabia

FAUNA OF ARABIA 23: 435–468 Date of publication: 15.07.2007

Review of the taxonomy of the spiny-tailed lizards of Arabia

(Reptilia: Agamidae: Leiolepidinae: Uromastyx)Thomas M. Wilms and Wolfgang Böhme

A b s t r a c t : Currently six species of the genus Uromastyx (Reptilia: Agamidae: Leiolepidinae), representing three phylogenetic lineages, are known to occur in Arabia: Uromastyx aegyptia, U. benti, U. leptieni, U. ornata, U. thomasi and U. yemenensis. The present paper gives an overview of the taxonomy of these lizards and presents new data on the morphology and ecology of Uro-mastyx leptieni.

Authors’ addresses: Thomas M. Wilms, Reptilium – Terrarien- und WüstenZoo, Werner-Heisenberg-Str. 1, 76829 Landau, Germany; e-mail: [email protected] Wolfgang Böhme, Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany; e-mail: [email protected]

Authors’ addresses: Thomas M. Wilms, Reptilium – Terrarien- und WüstenZoo, Werner-Heisenberg-Str. 1, 76829 Landau, Germany; e-mail: [email protected] Wolfgang Böhme, Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany; e-mail: [email protected]

INTRODUCTION

Spiny-tailed agamas are small to medium-sized, ground- or rock-dwelling lizards. Most species reach a maximum length of 25-50 cm, and only species of the Uromastyx aegyptia group can reach a total length of up to 70 cm or more. The animals have a bulky, depressed body and strong, short limbs. The tail is covered by spiny scales, arranged in distinct whorls. The tympanum is visible. None of the species has a nuchal or dorsal crest or a gular pouch or fan. Only a transverse fold at the throat is present (gular fold). Body scales are small and mostly homogenous, but some species have enlarged tubercular scales on the body and/or limbs.

The main diagnostic character of the genus is the highly specialised tooth-like bony structure replacing the incisor teeth in the upper jaw in adults. This transformation of the premaxillary bone to a tooth-like structure is an autapomorphy of the genus Uromastyx, convergent in Sphenodon, which has also a convergently acrodont dentition. Owing to morphological, ethological and ge-

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436 T.M. WilMs & W. BöhMe

netic synapomorphies, butterfly agamas of the genus Leiolepis are considered to be the closest living relatives of spiny-tailed lizards and both genera are therefore placed in the subfamily Leiolepidinae (synonym: Uromastycinae) (PeTers 1971, Moody 1980, BöhMe 1988, schMiTz et al. 2001, AMer & KuMAzAWA 2005).

Lizards of the genus Uromastyx occur in the desert belt of the Old World between 5°N and 35°N. Their range covers an enormous land mass, including northern Africa, the Middle East, Arabia, Iran, Iraq, Afghanistan, Pakistan and north-western India (Fig. 1). Today, 17 species are recognised, of which six species (with three additional subspecies) are native to Arabia (Kingdom of Saudi Arabia, Yemen, Sultanate of Oman, United Arab Emirates, Qatar and Kuwait (WilMs 2001, 2005; WilMs & schMiTz 2007).

The species of spiny-tailed lizards currently known to occur in Arabia are: Uromastyx aegyptia (Forskål, 1775), U. leptieni Wilms & Böhme, 2000, U. benti (Anderson, 1894), U. ornata von Heyden, 1827, U. yemenensis Wilms & Schmitz, 2007 and U. thomasi Parker, 1930. They rep-resent three different phylogenetic groups, with species compositions listed below (WilMs 2001, WilMs & schMiTz 2007).

Uromastyx habitats are generally characterised by high temperatures, low precipitation, sparse vegetation and marked seasonal changes (Plates 1-2). However, in the coastal regions of south and south-western Arabia (Asir and Hejaz Mountains, Hadramaut and Dhofar), Uromastyx habitats are more mesic because of the influence of the summer monsoon, resulting in high air moisture and elevated precipitation. These habitats are often well vegetated. Uromastyx species are mostly found on firm soil or on rock surfaces that provide shelter in the form of cracks and crevices. They also need, at least for part of the year, a sufficient amount of vegetation as a food source. Uromastyx live mainly in rocky desert and semi-desert habitats that are covered with rocks, scattered stones or gravel. Large continuous dune areas (erg) with drifting sand do not offer suitable habitats and represent range barriers. However, this does not prevent Uromastyx from travelling a few kilometres across sand from one gravel plain to another. Most species are digging, ground-dwelling animals, while some species live in rocky or mountainous areas and retreat into cracks and crevices. Some species have even been observed climbing acacia trees (U. aegyptia, U. dispar, U. ocellata and U. or-nata). Uromastyx are opportunistic herbivores, which means, that they feed mainly on plants and to a lesser extent on animals. By far the largest part of the diet is leaves, flowers, sprouts and seeds of various plant species. Spiny-tailed agamas are strictly diurnal.

MATERIALS AND METHODS

Morphological sampling

The taxonomy presented in the present paper is based on a revision of the genus Uromastyx (WilMs 1998; WilMs & BöhMe 2000 a, 2000 b, 2000 c; WilMs & schMiTz 2007), in which a total of 625 specimens have been examined, including 180 specimens of Arabian species. The specimens are kept in museum collections indicated by the following abbreviations:

BMNH British Museum of Natural History, London, UK (now: The Natural History Museum, London, UK, NHM)

MHNG Muséum d’histoire naturelle de la Ville de Genève, Geneva, SwitzerlandMNHNP Muséum national d’Histoire naturelle, Paris, FranceMTD Staatliches Museum für Tierkunde, Dresden, Germany

Review of the taxonomy of the spiny-tailed lizards of Arabia 437

MZUF Università di Firenze, Museo Zoologico “La Specola”, Florence, ItalyNHMW Naturhistorisches Museum Wien, Vienna, AustriaSMF Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a.M., GermanyZFMK Zoologisches Forschungsmuseum Koenig, Bonn, GermanyZMB Museum für Naturkunde der Humboldt-Universität zu Berlin, GermanyZMH Zoologisches Institut und Zoologisches Museum der Universität Hamburg, GermanyZSM Zoologische Staatssammlung München, Munich, Germany

Lists of examined specimens have been published by WilMs & BöhMe (2000 a, 2000 b, 2000 c), WilMs et al. (2002) and WilMs & schMiTz (2007).

To obtain new information on the morphology and ecology of Uromastyx leptieni, data were collected by Thomas Wilms and Brien Holmes at a site near Abu Dhabi International Airport (United Arab Emirates) between 11 June 2005 and 16 July 2005, including morphometric and meristic data of 82 live specimens. These specimens were subsequently released in the course of a translocation project, during which a total of 213 specimens were translocated (Tourenq 2005, C. Drew pers. comm.).

Fig. 1: Distribution range of the genus Uromastyx (after WilMs 2001).


Characters

For each museum specimen the following data were routinely recorded:– snout-vent-length (SVL)– tail length– head width, measured between the anterior margins of the ear openings– head length, measured from the tip of the snout to the anterior margin of the ear opening

on the left side– tail width between the 4th and 5th whorl– maximum tail width at the 5th whorl– number of scales around mid-body– number of scales between gular and inguinal fold (= ventrals)– number of gular scales (from the mental scale to a line between the anterior margins of the

ear openings)

Plate 1: Habitat of Uromastyx thomasi at Ras Hilf, Masirah Island, Oman. Photo: T. Wilms.

Plate 2: Habitat of Uromastyx aegyptia microlepis in Mahazat as-Sayd, Saudi Arabia. Photo: F. Krupp.


– number of scales from the mid of the lower end of the ear opening to the mental scale (left and right)

– number of scales between supralabial and enlarged subocular scale– number of scales around the 5th whorl– number of tail whorls– number of scales beneath the 4th toe on the left side– number of preanofemoral pores on either side– number of enlarged scales at the anterior margin of the ear opening– number of scales from the upper to the lower end of the left ear opening (approximately

three scale rows before the anterior margin of the ear opening)– number of scales from the upper end of the left ear opening to the first enlarged subocular

scale– intercalary scales between the whorls (presence or absence)– presence or absence of enlarged tubercular scales at the flanks and on the dorsum

With the exception of the number of scales around the mid-body, the same measurements and scale counts as in museums specimens were taken from the 82 live Uromastyx leptieni.

Statistical analyses

The statistical packages Excel 2000, SPSS (10.0) and R 2.2.1 (R develoPMenT core TeAM 2005) were used to run the analyses. Two multivariate procedures were selected to evaluate data and to explore relationships within the Uromastyx aegyptia group (sensu WilMs & BöhMe 2000 a).

Hierarchical cluster analysis

Hierarchical cluster algorithms find successive clusters using previously established clusters and thus grouping similar objects together. Because of the many different methods to calculate dis-tances in a given matrix and the large number of clustering algorithms this method is much more subjective than the ordination techniques such as PCA (rAsTegAr-PouyAni 2005). In the present study the use of average linkage method has produced the highest cophenetic correlation coef-ficients and the highest γ-coefficients. Both coefficients provide a measure for the reliability of a cluster analysis and, according to hAndl (2002), a γ-coefficient larger than 0.70 indicates that the fit of the analysis is satisfactory or better. A cophenetic correlation coefficient above 0.750 is con-sidered good fit (rAsTegAr-PouyAni 2005).

Principal component analysis (PCA)

Principal component analysis (PCA) is a multivariate procedure which rotates the data such that maxi-mum variabilities are projected onto the axes. Essentially, a set of correlated variables are transformed into a set of uncorrelated variables which are ordered by reducing variability. The uncorrelated variables are linear combinations of the original variables, and the last of these variables can be removed with min-imum loss of real data. The main use of PCA is to reduce the dimensionality of a dataset while retaining as much information as is possible. Results of PCA do not depend on prior specimen classification.


SYSTEMATIC ACCOUNT

Genus Uromastyx Merrem, 1820

Within the genus Uromastyx, six different species groups have been identified, three of which occur in Arabia (WilMs 1998; for a proposed phylogeny of the Arabian taxa see Fig. 2):

Uromastyx hardwickii groupUromastyx asmussi groupUromastyx princeps groupUromastyx ocellata groupUromastyx aegyptia groupUromastyx acanthinura group

The Arabian representatives of three of these groups (U. princeps group, U. ocellata group and U. aegyptia group) can be identified by means of the following key.

Key to the Arabian species of the genus Uromastyx

1 Preanal and femoral pores absent .............................................................................. 2– Preanal and femoral pores present ............................................................................. 4

2 143-187 scales around mid-body (average 160.05 ± 8.98), 66-86 ventral scales Uromastyxbenti– 163-227 scales around mid-body (average 192.53 ± 16.63), 79-97 ventral scales ...... 3

3 Ground colour light brown with five distinct cross-bands on the back (see Plate 15)Uromastyxyemenensisyemenensis

– Ground colour dark brown with light brown dots tending to form transverse rows on the back (see Plate 17) Uromastyxyemenensisshobraki

4 Tail short, up to 35 % of snout-vent-length, disc-shaped Uromastyxthomasi– Tail long, more than 60 % of snout-vent-length, 22-27 whorls ................................. 5

5 247-322 scales around mid-body .............................................................................. 6– 138-193 scales around mid-body .............................................................................. 7

6 110-150 ventral scales, preserved juveniles reddish brown with a dark vermiculation or with small irregularly arranged dark and light spots Uromastyxleptieni

– 126-193 ventral scales, preserved juveniles greyish brown with yellowish dots on the back arranged in transverse rows ............................................................................... 8

7 Ratio of tail length to maximum tail width at the 5th whorl: 3.61-5.30 Uromastyxornataornata– Ratio of tail length to maximum tail width at the 5th whorl: 3.03-3.96

Uromastyxornataphilbyi

8 126-158 ventral scales, enlarged lateral tubercles present Uromastyxaegyptiaaegyptia– 149-193 ventral scales, enlarged lateral tubercles absent Uromastyxaegyptiamicrolepis


Uromastyxaegyptia group

Within the Uromastyx aegyptia group, four different taxa have been described: the recently discov-ered U. occidentalis Mateo, Geniez, López-Jurado & Bons, 1998, from Western Sahara and U. lep-tieni Wilms & Böhme, 2000 from Oman and the United Arab Emirates and the two well-known taxa U. aegyptia Forskål, 1775, and U. microlepis Blanford, 1874. WilMs & BöhMe (2000 a) provide a detailed discussion of the taxonomic history of the taxa of the Uromastyx aegyptia group and recognise three taxa as valid species (U. aegyptia, U. leptieni and U. occidentalis). The fourth nominal taxon, microlepis, has been treated as a subspecies of U. aegyptia.

The main characters to distinguish the members of the Uromastyx aegyptia group from other Uromastyx species groups are their large body size (exceeding 700 mm in adult specimens), com-bined with extraordinarily small body scales.

Uromastyxaegyptia (Forskål, 1775) – Egyptian spiny-tailed lizardLacerta aegyptia Forskål, 1775. — Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in

itinere orientali observavit Petrus Forskål: 13.

Two subspecies are recognised: Uromastyx aegyptia aegyptia and Uromastyx aegyptia microlepis.

Uromastyxaegyptiaaegyptia (Forskål, 1775) Plate 3Lacerta aegyptia Forskål, 1775. — Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in

itinere orientali observavit Petrus Forskål: 13.Lacerta harbai Forskål, 1775. — Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in

itinere orientali observavit Petrus Forskål: 9 (?; synonymy fide MerreM 1820).Stellio spinipes Daudin, 1802. — Histoire naturelle, générale et particulière des reptiles. Vol. 4: 31.Uromastyx spinipes. — MerreM 1820; Versuch eines Systems der Amphibien – Tentamen systematis amphibiorum: 56

(n. comb.).Lacerta herbai. — MerreM 1820; Versuch eines Systems der Amphibien – Tentamen systematis amphibiorum: 56 (nomen

substitutum pro Lacerta harbai Forskål, 1775).Mastigura spinipes. — Fleming 1822; The Philosophy of Zoology; or, A General View of the Structure, Functions, and Clas-

sification of Animals. Vol. 2: 277 (n. comb.).Uromastix spinipes. — Boulenger 1885; Catalogue of the Lizards in the British Museum (Nat. Hist.). Vol. 1: 407.Uromastix aegyptius. — Anderson 1896; A Contribution to the Herpetology of Arabia, with a Preliminary List of the Reptiles

and Batrachians of Egypt: 79, 85.Uromastyx aegyptica. — Khalil & Hussein 1962; Bulletin of the Zoological Society of Egypt 17: 80.

Fig. 2: Proposed phy-logeny of the Arabian taxa of the genus Uro-mastyx based on mor-phological characters.


Ty p e s p e c i m e n : Neotype, adult , Egypt, Suez at the road to Cairo, VIII.1982, I. Re-hak, ZFMK 44216, designated by WilMs & BöhMe (2000 a).

D e s c r i p t i o n : S i z e : Large-growing taxon, maximum total length exceeding 700 mm.S c a l a t i o n : 247-322 scales around mid-body, 126-158 scales between gular and inguinal

fold, 33-59 gular scales, 24-31 scales from the mid of the lower end of the ear opening to the men-tal scale. On both sides 4-7 scales between supralabial and enlarged subocular scale; 29-46 scales around 5th whorl; 20-23 tail whorls; 16-20 scales beneath 4th left toe; 14-20 preanofemoral pores on either side.

Head covered with irregularly arranged scales of different size; the smallest situated above the eyes. In the occipital region slightly pointed scales. Scales of the neck very small. Nostrils big. An-terior margins of the ear openings covered with enlarged triangular, pointed scales (2-7 on either side). Scales of the underside of the head small. Scales of the chest, the belly and the underside of the extremities smooth. Scales of the soles slightly keeled. Subdigital scales with 3 or 4 distinct keels. Scales of the back small and smooth. At the flanks enlarged lateral tubercles. Lateral sac-ral region with distinctly enlarged tubercular scales. Scales of the upper side of the upper arms smooth, the scales of the forearm smooth or slightly pointed. Scales on the back of the upper arms distinctly enlarged, some slightly keeled. Dorsal parts of the hind legs covered with enlarged tuber-cular scales, between which are very small and smooth scales. Scales on the upper side of the feet enlarged and distinctly keeled.

C o l o u r : Uromastyx aegyptia aegyptia has the ability to undergo a physiological colour change. At high temperatures the animals have a light brown to light grey colour, with a black throat and small black dots on the neck. Some individuals have an entirely black to dark blue head. At low temperatures the animals are dark grey to nearly black. Juveniles have characteristic trans-verse rows of yellow to orange ocelli on their back. The main colour of the body is greyish brown.

D i s t r i b u t i o n : See Fig. 3. The nominotypic subspecies inhabits northern Egypt east of the River Nile, the Sinai Peninsula, Palestine and extreme north-western Saudi Arabia (Wadi Sa-wawin, Jebel as-Sinfa).

Uromastyxaegyptiamicrolepis Blanford, 1874 Plates 4-5Uromastix microlepis Blanford, 1874. — Proceedings of the Zoological Society of London (1874): 658.Uromastyx aegyptius microlepis. — MerTens 1956; Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 111

(1): 93.Uromastyx microlepis microlepis. — Arnold 1987 (ex errore); Proceedings of the Symposium on the Fauna and Zoogeography

of the Middle East; Beihefte zum TAVO A 28: 249.

Ty p e s p e c i m e n : Lectotype, adult , Iraq, Basrah, without date, Capt. Phillips, BMNH 1946.8.14.55, designated by WilMs & BöhMe (2000 a).

D e s c r i p t i o n : S i z e : Large-growing taxon, maximum total length exceeding 700 mm.S c a l a t i o n : 255-391 scales around mid-body, 149-193 scales between gular and inguinal

fold, 38-65 gular scales, 27-49 scales from the mid of the lower end of the ear opening to the men-tal scale. On both sides 5-8 scales between supralabial and enlarged subocular scale; 30-43 scales around 5th whorl; 20-24 tail whorls; 14-23 scales beneath 4th left toe; 13-21 preanofemoral pores on either side.

Head covered with irregularly arranged scales of different size; the smallest situated above the eyes. In the occipital region slightly pointed and convex scales. Scales of the neck very small. Nos-trils big. Anterior margins of the ear openings covered with enlarged triangular, pointed scales (3-9 on either side). Scales of the underside of the head small. Scales of the chest, the belly and the un-derside of the extremities smooth. Scales of the soles slightly keeled. Subdigital scales with 3-4 dis-


tinct keels. Scales of the back very small and smooth. No enlarged lateral tubercles; occasionally specimens from the northern parts of the area can have slightly enlarged tubercles exclusively at the rear parts of the body, but they are much smaller than tubercles in U. a. aegyptia. Scales of the upper side of the upper arms smooth, those of the forearm smooth or slightly pointed. Scales on the back of the upper arms distinctly enlarged, some slightly keeled. Dorsal parts of the hind legs covered with enlarged tubercular scales, between which are very small and smooth scales. Scales on the upper side of the feet enlarged and distinctly keeled.

Fig. 3: Distribution of Uromastyx aegyptia aegyptia and Uromastyx aegyptia microlepis (after WilMs & BöhMe 2000 a). For the distri-bution of U. a. microlepis in the UAE see Fig. 4. — Localities: 1: Wadi Sawawin at the Jebel as-Sinfa region; 2: Wadi Araba (Palestine and Jordan); 3: vicinity of Hazeva (Araba Valley, Palestine); 4: Sinai (Jebel el-Tik); 5: El Turkmania near Maghara; 6: 16 km W of Feiran Oasis; 7: Suez, Jebel Suez; 8: between Suez and Ismailia; 9: Baltim; 10: 120 km S of Cairo, 35 km S of Cairo, Khanka N of Cairo; 11: Wadi Hof (Helwan), Wadi Digla; 12: Kutamiya region, Wadi Iseili; 13: vicinity of Hurghada; 14: Jol, Hadramaut; 15: Bin Khautar, Hadramaut; 16: Wadi Qitbit; 17: Jiddat al-Harasis; 18: 100 km from Muscat; 19: Abu Dhabi (al-Hamran, Bada Zaid); 20: Ras al-Barr, Bahrain; 21: al-Jubail, 70 km N of Bahrain; 22: Dhahran; 23: Harad; 24: 100 km N of Riyadh; 25: al-Quwayiyah; 26: Wadi Sirra; 27: Wadi Fatimah; 28: Jeddah; 29: Yanbu al-Bahr; 30: Madinah; 31: Anaiza; 32: as-Sa’ira; 33: ad-Dibdibah; 34: Wadi al-Miyah; 35: between al-Gaisumah (= al-Qaysumah) and Turaif; 36: Kuwait; 37: Sirri Island; 38: Rudkhaneh-ye Shapur; 39: Fao; 40: 50 km W of Basrah, 60 km W of Basrah, Basrah; 41: W of Karbala; 42: Baghdad; 43: Kirkuk; 44: ar-Rutbah.


C o l o u r : Uromastyx aegyptia microlepis has the ability to undergo a physiological colour change. At high temperatures the animals are a light brown to yellow or greenish colour, with a black throat and small black dots on the neck and dorsum. Some individuals have an entirely black to dark blue head. At low temperatures the animals are dark grey, nearly black. Juveniles have characteristic transverse rows of yellow to orange ocelli on their back. The main colouration of the body is greyish brown.

D i s t r i b u t i o n : See Fig. 3 and (for details of the distribution in the UAE) Fig. 4. The border between the ranges of the subspecies U. a. aegyptia and U. a. microlepis is obviously east of Wadi Araba in Palestine and Jordan, and east of Wadi Sawawin in the Jebel as-Sinfa region of Saudi Arabia.

Uromastyx aegyptia microlepis lives in the deserts and semi-deserts of Arabia (Saudi Arabia, Yemen, Oman, United Arab Emirates, Qatar, Kuwait) and in Jordan, Syria, Iraq and coastal Iran.

Uromastyxleptieni Wilms & Böhme, 2000 – Leptien’s spiny-tailed lizard Plates 6-8Uromastyx leptieni Wilms & Böhme, 2000. — Herpetozoa 13 (3/4): 142.

Ty p e s p e c i m e n : Holotype, adult , United Arab Emirates, Wadi Siji, VI.1983, R. Lep-tien, ZFMK 52398.

D e s c r i p t i o n : S i z e : Maximum length was given as 53 cm (SVL 28.3 cm, paratype BMNH 85.II.7.4) by WilMs & BöhMe (2000 a). The largest specimen measured in summer 2005 was a male of 67.5 cm total length (SVL 37.5 cm), weighing 1562 g. The largest female had 58.3 cm total length (SVL 33.7 cm), weighing 978 g.

S c a l a t i o n : 238-294 scales around mid-body, 110-150 scales between gular and inguinal fold, 34-48 gular scales, 30-37 scales from the mid of the lower end of the ear opening to the men-tal scale. On both sides 5-9 scales between supralabial and enlarged subocular scale; 29-44 scales around 5th whorl; 19-24 tail whorls; 16-22 scales beneath 4th left toe; 12-19 preanofemoral pores on either side (in some specimens very small or absent).

Head covered with irregularly arranged scales of different size; the smallest situated above the eyes. Occipital scales and scales on the neck small and slightly pointed; on the neck, mostly on its sides, intermixed with enlarged triangular, pointed scales. Anterior margins of the ear openings covered with enlarged pointed scales (3-7 on either side). Scales of the dorsal parts of the forelegs slightly keeled. Scales of the ventral parts of the upper arm very small; on the forearm triangular and keeled. Scales of the soles distinctly keeled; each subdigital scale with 3-4 keels. Forelegs and back without enlarged tubercles. Flank scalation with a row of enlarged tubercular scales from the sacral region almost to the insertion of the forelegs. Dorsal scales are small and smooth, but some specimens may have a series of enlarged plate-like scales from the sacral region to half the body-length along the vertebral column. The diameter of these plates may reach three to four times the diameter of surrounding scales.

C o l o u r : In preserved specimens the main colour is olive-beige with small dark dots. Neck, head and throat are black. In some specimens the throat is marbled with black and orange. Ventral part of the forelegs, chest and belly are marbled with grey. Ventral parts of the hind legs and first half of the tail are grey. Colouration of juveniles is red-brown to dark brown with a dark brown to black net-like pattern or with small irregularly arranged dark and light spots. Active specimens are pale brown to almost entirely white, while cold specimens are grey with a dark grey to almost black head.

D i s t r i b u t i o n : See Fig. 4. Uromastyx leptieni is known from east of the Hajar al-Gharbi mountains in northern Oman (vicinity of Muscat to Musandam Peninsula), and from the north-eastern United Arab Emirates. The westernmost locality in the UAE is not Jebel Ali, as stated by


Plates 3‑8: Photographs of Uromastyx specimens (photos by T. Wilms, unless stated otherwise). 3: Uromastyx aegyptia aegyptia from Safawi, Jordan (photo: D. Modry). 4‑5: Uromastyx aegyptia microlepis; 4: specimen from the vicinity of Abu Kamal, Syria (photo: D. Modry); 5: extraordinary coloured specimen from Mahazat as-Sayd, Saudi Arabia. 6‑8: Uromastyx leptieni from the United Arab Emirates; 6: adult specimen from Jebel Jayah (photo: E.N. Arnold); 7: juvenile from study site near Abu Dhabi International Airport; 8: subadult from same locality.

WilMs & BöhMe (2000 a), but approximately at 24°27'N 54°38'E (study site near Abu Dhabi In-ternational Airport). In the UAE the nearest records of U. aegyptia microlepis are Shanayel, located at the Hmeem road, and the above-mentioned study site. The distance between these two localities is approximately 50 km. In Oman, the closest records of the two species are Muscat for U. leptieni and a locality about 100 km inland from Muscat for U. a. microlepis.


New information on the morphology and ecology of Uromastyx leptieni

The following paragraphs present observations and data collected on 82 live specimens of U. lep-tieni by Thomas Wilms and Brien Holmes at a site near Abu Dhabi International Airport (simply referred to as “study site” in the following) in the period between 11 June 2005 and 16 July 2005.

N a t u r a l h i s t o r y n o t e s : Little is known about the natural history of Uromastyx leptieni. cunninghAM (2000, 2001 a, 2001 b) provided data on the ecology of a Uromastyx population from the vicinity of al-Ain (“Dhub valley” at 23°25'N 55°35'E). Based on the photographs pub-lished by cunninghAM (2000, 2001 a), we are inclined to assign these animals to U. leptieni. This assumption is supported by the known distribution of U. leptieni in the UAE and the fact that specimens of Uromastyx from Sweihan, al-Khazna and al-Ain (near Jebel Hafeet) belong without doubt to U. leptieni (T. Wilms pers. obs.).

B u r r o w s p e c i f i c s : Our own observations show that most U. leptieni burrows enter the soil in a straight line for up to 1 m, with a slope of about 30-45°. Further down, the burrow in most cases turns to the left or to the right at approximately a right angle and then descends in the

Fig. 4: Distribution of Uromastyx aegyptia microlepis in the United Arab Emirates and of Uromastyx leptieni in the United Arab Emirates and Oman (based on BAhA el din 1996, WilMs & BöhMe 2000 a, Tourenq 2005, pers. obs.). It is assumed that popula-tions west of the left dotted line belong to U. a. microlepis and those east of the right dotted line belong to U. leptieni; the popula-tions between both lines (indicated by question marks) need further study to clarify their taxonomic status. — Localities: 1: Sha-nayel; 2: study site near Abu Dhabi International Airport; 3: al-Khazna; 4: Sweihan; 5: al-Ain; 6: Munayi; 7: vicinity of ar-Rustaq; 8: Muscat. — Identification: U. a. microlepis: specimens from near Jebel Dannah, from 23°45'N 53°33'E and from al-Hamran / Bada Zaid (Abu Dhabi) examined (WilMs & BöhMe 2000 a); specimens from Shanayel (locality 1) identified on the basis of photo-graphs taken by C. Tourenq and I. Barcelo. Uromastyx leptieni: specimens from the numbered localities 2-5 identified by pers. obs. (June/July 2005); from Jebel Ali, Tawi Bil Khabis, Jebel Jayah and Wadi Siji examined (WilMs & BöhMe 2000 a).


form of a spiral or in a zigzag line. Reported burrow length is more than 180 cm with a depth of 90-120 cm (cunninghAM 2001 a), which can be confirmed by our own observations.

Data loggers were used to measure air temperatures in the habitat (at a height of 1 m above ground level) and soil temperatures at the surface outside the burrows (just in front of the entrance) as well as inside (80 cm deep in the burrow) during a period between 20 June and 11 July 2005. Air temperatures varied between a maximum of 51.8 °C and a minimum of 22.5 °C, while the soil surface temperatures were between 55.4 °C and 22.1 °C. Within the burrow the temperature was relatively stable, with a maximum of 40.1 °C and a minimum of 32.8 °C, thus providing ambient temperatures to the lizards and allowing survival even when temperatures outside the shelters may reach lethal values (Fig. 5). Apart from shelter against high temperatures, the burrows offer protec-tion from drought. All dug-up burrows entered the moist soil layer, which could be important for hydroregulation, as Uromastyx skin seems to have the ability to absorb water from moist surround-ings (KevorK & Al-uThMAn 1972).

Existing burrows are obviously reoccupied after the death of the former occupant. In the vi-cinity of some burrows, remains of large Uromastyx can be found, excavated by the successor. This finding implies the high ecological value of the burrows, supposedly because of the difficulties to dig into extraordinarily hard soil. The habitats exploited by Uromastyx leptieni for burrowing at the study site consisted of aeolianite outcrops, gravel plains (partly sandy) and low hills. We observed that more than one burrow was regularly used by a particular animal.

D a i l y a c t i v i t y : So far, only data concerning daily activity during summer are available. According to cunninghAM (2000), who provided data for May and June, the animals spend an average of 81.2 % of the daylight hours underground. During our observation period, a total of 117 activity records have been taken (see Fig. 6). The diagram shows that U. leptieni exhibits a bimodal activity rhythm during summer time with the highest activity before 11:30 and a resting

Fig. 5: Temperatures inside and outside of a Uromastyx leptieni burrow at the study site near Abu Dhabi International Airport be-tween 20 June and 11 July 2005.


period during the hottest time of the day. Afternoon activity is very limited and restricted to the time after 15:30.

B o d y t e m p e r a t u r e s : During the observation period, body temperatures (cloacal tempera-tures) of four hand-caught Uromastyx leptieni were measured within less than one minute of cap-ture (only animals caught without chasing were included). The cloacal temperature of these active animals was between 35.8 °C and 42.2 °C (see Table 1).

To obtain information on the maximum cloacal temperature tolerated, measurements were taken in 32 lizards caught in cage traps, which had been exposed to the sun. Cloacal temperature ranged between 34.9 °C and 47.3 °C. Despite a cloacal temperature of 47.3 °C, which is close to the lethal temperature in U. acanthinura (48-50 °C according to verneT et al. 1988), this speci-men appeared healthy and alert.

D i e t : According to cunninghAM (2000, 2001 a), Uromastyx leptieni feeds on 10 different plant species at “Dhub valley”: Pennisetum divisium, Stipagrostis plumosa, Aerva javanica, Leptadenia py-rotechnica, Moltkiopsis ciliata, Heliotropium kotschyi, Haloxylon salicornicum, Citrullus colocynthis, Taverniera cuneifolia and Polygala erioptera, of which grasses of the family Gramineae were pre-ferred. This preference for grasses is confirmed by our observations, which suggest that U. leptieni avoids feeding on the abundant Zygophyllum qatarense. A very limited amount of animal matter is consumed. Obviously, in June and July feeding occurs almost exclusively before 09:00.

At the study site, faecal samples were collected and U. leptieni tracks followed to determine food preferences. The longest measured track of a specimen collecting plant material was 387 m. This specimen, a large female (total length 51.8 cm, weight 782 g), started at her burrow, grazed on several grasses and bushes before returning to her burrow.

P o p u l a t i o n s t r u c t u r e a n d d e n s i t y : Within the sample of 82 specimens caught at the study site and subsequently examined in detail, the sex ratio of males to females was 1 : 1.43

Fig. 6: Activity profile of a Uromastyx leptieni population at the study site near Abu Dhabi International Airport between 15 and 29 June 2005 in correlation with mean air temperatures.


with only 12 % of the animals being juveniles. Males generally grow larger than females (t-test: t = 4.57, α = 0.01) with a maximum total length of 67.5 cm (females: 58.3 cm). There is also a significant difference in body weight between adult males and females (t-test: t = 5.84, α = 0.01), with adult males (total length larger than 40 cm) ranging between 384 g and 1572 g (mean: 999 g), while adult females (total length larger than 39 cm) range between 278 g and 983 g (mean: 599 g). Fig. 7 shows total length of juvenile, male and female Uromastyx leptieni plotted against weight.

It is well known that Uromastyx females loose a considerable portion (approx. 30-45 %) of their body weight after oviposition (zAri 1999, WilMs 2001). Some of the females caught were very skinny and emaciated, so that a recent oviposition could be supposed. This assumption was supported by oviposition observed in some of the females caught at the study site, which were kept at the Breeding Centre of Endangered Arabian Wildlife, Sharjah (D. Egan, pers. comm.). This ob-servation indicates that reproduction of U. leptieni in the UAE takes place at least until July.

During the course of the translocation project, in total 213 specimens of U. leptieni were caught, and this number is considered to be nearly the whole adult population of the area in question. Fig. 8 gives an overview of the distribution of the Uromastyx population on the study site. Three core areas with exceptionally high densities of Uromastyx have been identified within

Table 1: Cloacal temperatures of four hand-caught Uromastyx leptieni at the time of capture, in comparison with soil and air tem-peratures.

Time of capture Cloacal temperature Soil temperature Air temperature09:25 40.5 °C 41.1 °C 39.3 °C10:15 42.2 °C 49.7 °C 40.0 °C11:00 36.5 °C [no record] [no record]11:05 35.8 °C 32.6 °C 35.1 °C

Fig. 7: Scatter plot of total length against body weight of 82 individuals of Uromastyx leptieni from the study site near Abu Dhabi International Airport. Males and females are indicated by the respective symbols, juveniles by circles.


the site, covering 160 ha (Area 1), 130 ha (Area 2) and 13 ha (Area 3). Despite intensive search-ing, only a few specimens have been found outside these areas. With 64 individuals of U. leptieni caught in Area 1, the density was one animal per 2.5 ha, while in Area 2 (131 individuals) and Area 3 (14 individuals) the density was one animal per approximately one hectare (0.99 and 0.93, respectively).

Vegetation density in Area 2 was assessed by randomly placing six vegetation plots (each 100 m2). Vegetation cover in three plots was lower than 1 %; in the other three plots it ranged be-tween 2 % and 3 %. Area 1 was severely degraded and had a lower vegetation cover than Area 2.

Multivariate analyses of the taxa of the Uromastyx aegyptia complex

Cluster analysisDistance phenograms based on metric and meristic characters of individual adult specimens of Uromastyx aegyptia and U. leptieni (41 males, 48 females; definition of variables V 1 to V 9 see

Fig. 8: Satellite image of the study site near Abu Dhabi International Airport, with outlines of the three core areas and capture lo-calities of Uromastyx leptieni superimposed.


Table 2) were calculated, for males and females separately, using the average linkage method (Fig. 9 ). The cophenetic correlation coefficient is 0.842 for males and 0.857 for females, while the γ-coefficient is 0.712 for males and 0.739 for females.

In males (left phenogram), one specimen from Egypt (number 1; BMNH 97.10.28.213, Baltim Delta) is separated from all other specimens at the first major dichotomy. We consider this animal an aberrant specimen because of its extraordinarily long tail (102.83 % of the SVL). The second dichotomy separates two branches, of which one encompasses all but one individual U. leptieni and the other branch the remaining U. aegyptia and the single U. leptieni (number 20).

In females (right phenogram), three U. leptieni specimens (numbers 17, 30 and 48) are separated by the first dichotomy from all other specimens. All three specimens have very long

Fig. 9: Distance phenogram resulting from the cluster analysis of 41 adult males (left) and 48 adult females (right) of the Arabian taxa of the Uromastyx aegyptia complex.


tails (101.79 %, 109.26 % and 112.30 % of the SVL, respectively). The second major dichotomy separates roughly U. leptieni from U. aegyptia, with three individual U. aegyptia (numbers 3, 4 and 6) clustering together with the majority of U. leptieni and one individual U. leptieni (number 11) clustering with the remaining U. aegyptia.

Because females of closely related Uromastyx species are generally more similar to one another than males, it is not surprising that the cluster analysis of the datasets of individual females of the three nominal taxa did not show such a high degree of congruence with the supposed taxonomic arrangement as those of males did.

Principal component analysis (PCA)A principal component analysis was run using the variables V 1 to V 9 for the same individual males and females separately (for factor loading on the first three principal components see Ta-ble 3). In the projection of the first two principal components the male specimens (Fig. 10, above) group together according to prior specimen classification, providing evidence that the operational taxonomic units (OTUs) involved represent in fact the three taxa U. a. aegyptia, U. a. microlepis and U. leptieni. The PCA run with the datasets of individual females (Fig. 10, below) did not clearly separate the OTUs but has produced an area of broad overlapping. This result is in accord-ance with the result of the hierarchical cluster analysis, showing that females of different OTUs are much more similar among each other than are males.

Variable DefinitionV 1 [snout-vent-length] × 100 / [tail length]V 2 [head width] / [head length]V 3 number of tail whorlsV 4 number of scales beneath the 4th toe on the left sideV 5 [maximum tail width at the 5th whorl] – [tail width between the 4th and 5th whorl] / 2V 6 [tail length] / [maximum tail width at the 5th whorl]V 7 number of gular scalesV 8 number of scales between gular and inguinal foldV 9 number of scales around the 5th whorl

Males FemalesVariable Component 1 Component 2 Component 3 Component 1 Component 2 Component 3

V 1 0.189 0.615 –0.155 –0.348 0.339 –0.298V 2 0.456 –0.229 –0.406 0.190 0.387V 3 0.787 –0.282 –0.487V 4 0.322 –0.365 0.460 –0.135V 5 0.299 0.203 –0.357 –0.223 0.297 0.573V 6 0.332 0.552 –0.451 –0.291V 7 –0.424 0.450 0.398 0.261 –0.288V 8 –0.509 0.136 –0.233 0.439 0.386 –0.233V 9 –0.148 0.147 0.566

Eigen values 2.579 1.611 1.131 2.504 1.479 1.308Accumulated percent of trace 29.4 % 47.8 % 60.6 % 28.4 % 45.2 % 60.0 %

Table 2: Definition of variables based on metric and meristic characters. To eliminate allometric effects, proportions have been calculated using metric values.

Table 3: Factor loading on the first three principal components from a correlation matrix of variables V 1 to V 9 for individual males and females.


Fig. 10: Ordina-tion of the individual 41 adult males (above) and 48 adult females (below) of the Arabian taxa of the Uromastyx aegyptia complex on the first two principal com-ponents (Variables: V 1 to V 9, see Table 2).


Uromastyxocellata group

This group consists of four species (U. benti Anderson, 1894, U. ocellata Lichtenstein, 1823, U. ornata von Heyden, 1827 and U. yemenensis Wilms & Schmitz, 2007) inhabiting the African and Arabian littoral regions of the Red Sea as well as the coastal areas of the Gulf of Aden in Yemen and southern Oman (WilMs & BöhMe 2000 c). One species, Uromastyx macfadyeni, that has been previously assigned to this group due to morphological similarities, is most probably not related to the other members of the Uromastyx ocellata group (AMer & KuMAzAWA 2005, WilMs & schMiTz 2007).

Three of the four species within the group, U. benti, U. ornata and U. yemenensis, are known to occur in Arabia. The taxonomic history in this group is confusing, leading to difficulties in the proper identification of the species in question. According to WerMuTh (1967), all taxa deserve specific rank, but Arnold (1986) treats the taxa ornata and philbyi as subspecies of Uromastyx ocel-lata, and lAnzA (1988) regards macfadyeni likewise as a subspecies of U. ocellata. WilMs (1995) follows this point of view and treats ornata, philbyi and macfadyeni as subspecies of U. ocellata.

According to Moody (1987), “ocellatus and ornatus [are] nearly indistinguishable” (see also Jo-ger 1987), and schäTTi & gAsPereTTi (1994) stated that they “consider benti, ornata and philbyi to be conspecific with U. ocellata Lichtenstein”. schäTTi & desvoignes (1999) follow this concept and regard U. benti, U. ornata, and U. philbyi as junior synonyms of U. ocellata. The examination of 148 specimens of the Uromastyx ocellata group, including type material of all involved taxa, by WilMs & BöhMe (2000 c) revealed that in fact four species (Uromastyx ocellata, U. benti, U. mac-fadyeni and U. ornata as well as the recently discovered U. yemenensis) are involved and that they can easily be distinguished. Of these species, U. macfadyeni is obviously not closely related to the other taxa, because it forms a strongly supported clade basal to and clearly outside the rest of the species of the Uromastyx ocellata group (WilMs & schMiTz 2007). For a detailed discussion of the taxonomy and a key to the species of the Uromastyx ocellata group, see WilMs & BöhMe (2000 c).

Uromastyxbenti (Anderson, 1894) – Bent’s spiny-tailed lizard Plate 9Aporoscelis benti Anderson, 1894. — Annals and Magazine of Natural History (6) 14: 376.Uromastix (Aporoscelis) benti. — Anderson 1896; A Contribution to the Herpetology of Arabia, with a Preliminary List of the

Reptiles and Batrachians of Egypt: 33, 63, 79, 85.Uromastix simonyi Steindachner, 1899. — Anzeiger der Akademie der Wissenschaften in Wien, Mathematisch-Naturwissen-

schaftliche Klasse 36: 143.Uromastix philbyi. — hAAs & BATTersBy 1959; Copeia 1959: 202 (synonymy fide Arnold 1986).Uromastyx thomasi. — Al-BAdry & Al-sAfAdi 1982; Proceedings of the Egyptian Academy of Sciences 34: 66 (synonymy fide

schäTTi 1989).Uromastyx ocellata benti. — schäTTi & gAsPereTTi 1994; Fauna of Saudi Arabia 14: 369.Uromastyx ocellata. — schäTTi & desvoignes 1999; The Herpetofauna of Southern Yemen and the Sokotra Archipelago: 39.

Ty p e s p e c i m e n : Lectotype, adult , Yemen, Wadi Hadramaut, without date, J. Ander-son, BMNH 1946.8.11.72, designated by WilMs & BöhMe (2000 c).

D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 360 mm, maximum SVL 196 mm (NHMW 16174).

S c a l a t i o n : 143-187 scales around mid-body, 66-86 scales between gular and inguinal fold, 23-33 gular scales, 19-27 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 4-7 scales between supralabial and enlarged subocular scale; 28-38 scales around 5th whorl; 22-26 tail whorls; 11-15 scales beneath 4th left toe. No preanofemoral pores.

Head covered with irregularly arranged smooth scales of different size. Anterior margins of the ear openings covered with enlarged scales (3-6 on either side). No enlarged tubercular scales


on neck, forelegs or body. Body scales small and smooth. Thighs without enlarged tubercular scales. Dorsal parts of the lower leg covered with very small scales, beneath the knee conical tu-bercular scales and/or enlarged keeled scales. Ventral parts of the lower leg covered with smooth scales, resembling the scales of the soles in size and shape. Tail long and narrow, approximately 82.41 ± 5.48 % of SVL. Median scales of the upper side of the tail strongly keeled, dorsolateral scales of the tail with well-developed spines.

C o l o u r : Males: In preserved specimens, ground colour of back, tail and hind legs yellowish brown. Tail without distinct pattern, hind legs with a turquoise and orange colouration. Back with a pattern consisting of dark brown lines and dots, as well as 7-9 rows of ocelli (ivory-coloured with dark brown edges). Dorsal side of the forelegs anthracite-coloured with orange and green colour elements. Hands yellowish brown. Head orange or blue, marbled with dark brown and black. Un-derside of the head anthracite-coloured with some orange dots. Ventral parts of forelegs and chest marbled with grey. Belly with narrow grey/anthracite cross-bands. — Females: much paler in col-ouration, with a yellowish brown ground colour and a pattern of small dark brown lines and dots.

D i s t r i b u t i o n : See Fig. 11. Uromastyx benti occurs in southern and south-eastern Yemen, from the vicinity of Azzan eastwards to the Hadramaut Valley and along the coast of the Arabian Sea. In the Sultanate of Oman this species is only known from the vicinity of Mirbat in south-western Oman (seufer et al. 1998, WilMs & hulBerT 2000).

Uromastyxornata von Heyden, 1827 – Ornate spiny-tailed lizardUromastyx ornatus von Heyden, 1827. — Atlas zu der Reise im nördlichen Afrika von Eduard Rüppell. Abtheilung 1. Rep-

tilien: 1.

Two subspecies are recognised: Uromastyx ornata ornata and Uromastyx ornata philbyi.

Fig. 11: Distribution of Uromastyx benti (after WilMs & schMiTz 2007). — Localities: 1: Azzan; 2: al-Mukalla; 3: Ras Fartak; 4: vicinity of Mirbat.


Uromastyxornataornata von Heyden, 1827 Plates 10-14Uromastyx ornatus von Heyden, 1827. — Atlas zu der Reise im nördlichen Afrika von Eduard Rüppell. Abtheilung 1. Rep-

tilien: 1.Uromastix oronatus [sic]. — Farag & Banaja 1980 (typographic error); Bulletin of the Faculty of Science, King Abdulaziz

University 4: 12.Uromastyx ocellatus ornatus. — Arnold 1986; Fauna of Saudi Arabia 8: 393.Uromastyx ocellata ornata. — lAnzA 1988; Biogeographia 14: 420.Uromastyx ocellata. — schäTTi & desvoignes 1999; The Herpetofauna of Southern Yemen and the Sokotra Archipelago: 39.

Plates 9‑14: Photographs of Uromastyx specimens (photos by T. Wilms, unless stated otherwise). 9: Uromastyx benti, male from the vicinity of Mirbat, Oman. 10‑14: Uromastyx ornata ornata from Sinai Peninsula, Egypt; 10: male from Wadi Feiran (photo: D. Modry); 11: adult male, blue colour phase; 12: adult male, red colour phase; 13: pregnant female; 14: hatchling.


Ty p e s p e c i m e n : Holotype, female, Saudi Arabia, Mohila (= al-Muwaylih), 1828, E. Rüppell, SMF 10403.

D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 368 mm, maximum SVL 196 mm (BMNH 97.10.28.199).

S c a l a t i o n : 149-185 scales around mid-body, 75-99 scales between gular and inguinal fold, 22-31 gular scales, 21-27 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 3-5 scales between supralabial and enlarged subocular scale; 19-25 scales around 5th whorl; 20-23 tail whorls; 11-15 scales beneath 4th left toe; 7-14 preanofemoral pores on either side.

Scalation homogenous, without enlarged tubercular scales on head, neck and back. Head cov-ered with irregularly arranged scales of different size; the smallest situated above the eyes. In the occipital region slightly pointed and slightly keeled scales. Scales of the neck very small. Anterior margins of the ear openings covered with enlarged triangular, pointed scales (5-7 on either side). Scales of the underside of the head small and smooth. Scales of the chest, the belly and the under-side of the extremities smooth. Scales of the soles slightly keeled. Subdigital scales with 3-4 keels. Scales of the back small and smooth. No tubercles on body and forearms. Scales of the forearm smooth or slightly keeled. Dorsal parts of the hind legs covered with enlarged conical tubercular scales. Between those scales very small and smooth scales. Scales on the upper side of the feet en-larged and smooth or slightly keeled.

C o l o u r : Uromastyx ornata is a very variable species. Colour of the males is green, blue or red, with an irregularly reddish brown net-like pattern and yellow spots on the back. Sometimes yellow cross-bands are present. Venter with dark pattern. Females are not as colourful as males. They are light brown with dark brown spots and sometimes light yellow or light red spots. Belly without pattern, yellowish or white.

Uromastyxornataphilbyi Parker, 1938Uromastyx philbyi Parker, 1938. — Annals and Magazine of Natural History (11) 1: 484.Uromastyx ocellatus philbyi. — Arnold 1986; Fauna of Saudi Arabia 8: 416.Uromastyx ornatus philbyi. — Arnold 1987 (ex errore); Proceedings of the Symposium on the Fauna and Zoogeography of

the Middle East; Beihefte zum TAVO A 28: 252.Uromastyx ocellata philbyi. — schäTTi & gAsPereTTi 1994; Fauna of Saudi Arabia 14: 369.Uromastyx ocellata. — schäTTi & desvoignes 1999; The Herpetofauna of Southern Yemen and the Sokotra Archipelago: 39.

Ty p e s p e c i m e n : Holotype, , Saudi Arabia, southern Hejaz between mountains and Rub al-Khali, between Makkah and Shabwa, H.St.J.B. Philby, BMNH 1946.8.14.65 (former number: BMNH 1938.2.1.1).

D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 341 mm, maximum SVL 205 mm (MZUF 27906).

S c a l a t i o n : 138-193 scales around mid-body, 69-96 scales between gular and inguinal fold, 17-31 gular scales, 18-22 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 3-5 scales between supralabial and enlarged subocular scale; 22-29 scales around 5th whorl; 17-22 tail whorls; 11-14 scales beneath 4th left toe; 7-14 preanofemoral pores on either side.

Scalation homogenous and similar to that of Uromastyx o. ornata. Anterior margins of the ear openings covered with 5-7 enlarged scales.

C o l o u r : Similar to that of Uromastyx o. ornata.D i s t r i b u t i o n : See Fig. 12. Uromastyx o. ornata occurs on the Sinai Peninsula and in adja-

cent north-western Saudi Arabia, as well as at the extreme southern tip of Israel (area of Elat). Uro-


mastyx o. philbyi occurs in the mountains of western Saudi Arabia, from the Jebel as-Sinfa region to the southern Hejaz. WilMs & BöhMe (2000 c) provide a discussion of the overlapping distribu-tion of the two subspecies and point out that, on the basis of the available collections, a decision whether philbyi deserves specific rank or not is impossible.

Fig. 12: Distribution of Uromastyx ornata ornata and Uromastyx ornata philbyi (after WilMs & BöhMe 2000 c); sites with sympat-ric occurrence of both subspecies are shown. — Localities: 1: Sharm el-Sheikh; 2: Dahab; 3: Tor; 4: Mount Sinai, Wadi Nassen, Kara’ah; 5: Wadi Feiran; 6: Elat; 7: Wadi Makfa; 8: Mohila (= al-Muwaylih); 9: Jebel as-Sinfa; 10: Jeddah, Burayman, Bazzah; 11: Makkah; 12: Taif; 13: Jebel Lababa (18°01'N 42°02'E); 14: southern Hejaz (between Makkah and Shabwa); 15: Ju’ Amlah (approx. 26 km NW of Sa’dah).

Plates 15‑22: Photographs of Uromastyx specimens (photos by T. Wilms, unless stated otherwise). 15‑16: Uromastyx yemenensis yemenensis; 15: holotype in dorsal view; 16: same in ventral view. 17‑19: Uromastyx yemenensis shobraki; 17: holotype in dorsal view; 18: same in ventral view; 19: male from northern Yemen. 20‑22: Uromastyx thomasi from Ras Hilf, Masirah Island, Oman; 20: male; 21: mating pair (photo: F. Hulbert); 22: hatchling.


Uromastyxyemenensis Wilms & Schmitz, 2007 – Southern Arabian spiny-tailed lizardUromastyx yemenensis Wilms & Schmitz, 2007. — Zootaxa 1394: 12.

Two subspecies are recognised: Uromastyx yemenensis yemenensis and Uromastyx yemenensis shobraki.

Uromastyxyemenensisyemenensis Wilms & Schmitz, 2007 Plates 15-16Uromastyx yemenensis yemenensis Wilms & Schmitz, 2007. — Zootaxa 1394: 12.

Ty p e s p e c i m e n : Holotype, adult , Republic of Yemen, Abyan Governorate, vicinity of Lodar (= Lawdar), don. 1985, I. Haikal, ZFMK 47861.

D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 332 mm (BMNH 1946.8.11.68).

S c a l a t i o n : 208-227 scales around mid-body, 86-85 scales between gular and inguinal fold, 32-36 gular scales, 24-28 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 4-6 scales between supralabial and enlarged subocular scale; 37-40 scales around 5th whorl; 23-26 tail whorls; 15-18 scales beneath 4th left toe; no preanofemoral pores.

Anterior margins of the ear openings covered with 3-6 enlarged scales. For a more detailed description see WilMs & schMiTz (2007).

C o l o u r : Males: In preserved specimens, ground colour of the back, tail and hind legs yel-lowish brown. Tail without distinct pattern, hind legs with very small dark brown dots. Back with a pattern consisting of dark brown lines and dots; five distinct cross-bands without or with very few patterns on the back. Dorsal side of the forelegs anthracite-coloured. Hands yellowish brown. Head yellowish brown, marbled with dark brown. Underside of the head anthracite-coloured with some yellowish brown dots. Ventral parts of forelegs and chest marbled with grey. Belly with narrow grey/anthracite cross-bands. — Females are much paler in colour, with a yellowish brown ground colour with a pattern of small dark brown lines and dots. Five pale cross-bands on the back. The ground colour of the ventral side is a light yellowish brown.

Uromastyxyemenensisshobraki Wilms & Schmitz, 2007 Plates 17-19Uromastyx yemenensis shobraki Wilms & Schmitz, 2007. — Zootaxa 1394: 16.

Ty p e s p e c i m e n : Holotype, adult , Yemen, Mafraq al-Mukha, 13.5 km W of Mafraq, 5-6.IV.1988, B. Schätti, ZFMK 48681.

D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 393 mm (MHNG 2538.47).

S c a l a t i o n : 163-207 scales around mid body, 79-97 scales between gular and inguinal fold, 25-33 gular scales, 23-31 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 3-5 scales between supralabial and enlarged subocular scale; 32-39 scales around 5th whorl; 24-27 tail whorls; 15-17 scales beneath 4th left toe; no preanofemoral pores.

Anterior margins of the ear openings covered with 3-5 enlarged scales. For a more detailed description see WilMs & schMiTz (2007).

C o l o u r : In preserved specimens, dorsal surface of head, body and hind limbs dark brown, tail lighter. Light brown roundish dots (diameter 4 or 5 scales) are present on the dorsum, tending to form transverse rows. In addition, irregular light brown dots are present on the whole dorsum. Colour of the hands not different from the colour of the forearm. Head dark brown, with light brown pattern. Ventral side of the specimen yellowish brown. Ventral side of head and chest mar-bled with anthracite and dark brown. There is an anthracite-coloured stripe-like pattern on the sides of the belly.

D i s t r i b u t i o n : See Fig. 13. Uromastyx yemenensis is distributed in the south-western parts of Yemen, with the range extending from the vicinity of Lawdar west to the Red Sea and from


Zabid south to the Gulf of Aden. The areas of the subspecies U. y. yemenensis and U. y. shobraki are separated by a series of mountains and mountain ranges (example: Jebel Manar, Jebel Sawraq, Jebel Sabir, Jebel Dawran) with heights up to 3,200 m (WilMs & schMiTz 2007).

Uromastyxprinceps group

This group includes the Arabian species Uromastyx thomasi and the Somalian species U. princeps (O’Shaughnessy, 1880). The shared derived character is the extraordinarily short and broad tail with only 9-14 annuli.

Uromastyxthomasi Parker, 1930 – Thomas’s spiny-tailed lizard Plates 20-22Uromastix thomasi Parker, 1930. — Annals and Magazine of Natural History (10) 6: 595.

Ty p e s p e c i m e n : Holotype, , Oman, Rub al-Khali, Dhofar, Bu Ju’ay, without date, B. Thomas, BMNH 1946.8.14.43 (former number: BMNH 1930.6.30.2).

D e s c r i p t i o n : S i z e : Small species, maximum total length approximately 24 cm, maxi-mum SVL approximately 19 cm.

S c a l a t i o n : 125-150 scales around mid-body, 72-100 scales between gular and inguinal fold, 25-36 gular scales, 19-25 scales from the mid of the lower end of the ear opening to the men-tal scale. On both sides 2-4 scales between supralabial and enlarged subocular scale; 28-34 scales around 5th whorl; 11-13 tail whorls; 13-18 scales beneath 4th left toe; 12-19 preanofemoral pores on either side.

Snout short. Head, neck and body without enlarged tubercular scales. Head covered with ir-regularly arranged scales of different size; the smallest situated above the eyes. Scales of the under-

Fig. 13: Distribution of Uromastyx yemenensis yemenensis and Uromastyx yemenensis shobraki. — Localities: 1: Oued (= Wadi) Zabid; 2: Mafraq al-Mukha; 3: Ta’izz; 4: Oued (= Wadi) Tiban; 5: Lahj; 6: 50 mi from Aden; 7: Zinjibar; 8: Lawdar.


side of the head small and smooth. Scales of the chest, the belly and the underside of the extremi-ties smooth. Scales of the soles distinctly keeled. Subdigital scales with 3 or 4 distinct keels. Upper side of forelegs and thighs of hind legs covered with enlarged triangular, slightly keeled scales. Size of the scales of the anterior margin of ear opening equals those of the cheek. Body scales small and subimbricate. Scalation of the back not homogenous with respect to scale size. Smallest scales along the vertebral column, biggest at the flanks. Tail distinctly depressed, short and disc-shaped.

C o l o u r : Yellowish green with a dark net-like pattern. A broad red stripe runs along the vertebral column from the neck to the first half of the tail. In some specimens the head can be an orange to red colour. Ventral side yellowish or white. Neck and sides of the head of the juveniles striped (black and white). Colour of the upper side of the body black with six lighter transverse bands, between which are yellowish to orange-coloured ocelli. Tail dorsally brown with some large black spots. Belly and throat white. Ventral side of the tail white with black dots.

D i s t r i b u t i o n : See Fig. 14. Uromastyx thomasi lives in coastal Oman.

DISCUSSION

The following discussion of the taxonomy of the genus Uromastyx is based on the results of our morphological studies (WilMs & BöhMe 2000 a, 2000 b, 2000 c; WilMs 2001). Recently AMer & KuMAzAWA (2005) found that a phylogeny of the genus Uromastyx, established on the basis of mo-lecular genetics, is mostly consistent with our results, but they found a few discrepancies. In their study, U. macfadyeni clustered with the species of the African U. acanthinura group, whereas mor-phologically it is grouped within the Afro-Arabian U. ocellata group. Another difference between

Fig. 14: Distribution of Uromastyx thomasi (after WilMs 1998). — Localities: 1: between Bawi and Salalah; 2: Thumrait (= Thamar-it); 3: Bu (= Bin) Ju’ay; 4: al-Ajaiz; 5: Ras Duqm, Jiddat al-Harasis; 6: Wadi Hauf (= Wadi Haluf ); 7: Masirah Island.


the two approaches is the placement of the U. aegyptia group within the cladogram. Morphologi-cally, it is regarded as the sister taxon of the U. acanthinura group, while genetically it clusters with the Arabian U. ocellata clade (without U. macfadyeni). Further studies, combining morphological and genetic methods, are needed to establish reliable phylogenetic relationships within the genus.

Uromastyxocellata group

As pointed out previously, the Uromastyx ocellata complex consists of six closely related taxa (ocel-lata, ornata, philbyi, benti, yemenensis and shobraki; but see AMer & KuMAzAWA 2005 for the status of macfadyeni), distributed around the Red Sea as well as in the coastal area of the Gulf of Aden (Djibouti, Somalia, Yemen and Oman), of which four deserve specific rank (U. ocellata, U. ornata, U. benti and U. yemenensis). According to WilMs & BöhMe (2000 c), U. philbyi should be treated as a subspecies of U. ornata until new evidence becomes available.

To provide an overview on the taxonomic problems relating to the proper identification of the taxa within the U. ocellata group, it seems to be necessary to discuss the identity of U. macfadyeni, despite the fact that this species occurs outside the geographical scope of the present paper. Ac-cording to schäTTi (1989), one specimen from Djibouti (MHNG 2394.100) corresponds well with the type specimens of U. macfadyeni from the Guban (northern Somalia). He states that: “Ein Tier der Originalserie (BMNH 1937.12.5.125, Borama) besitzt einzelne, weit auseinanderliegende Ocellen auf dem Rücken, beim Holotypus verschmelzen diese zu einem Netzmuster” [One animal from the original series (BMNH 1937.12.5.125, Borama) has single, widely separated ocelli on the back, in the holotype they merge to a net-like pattern]. The examination of those animals re-vealed that in fact BMNH 1937.12.5.125 (Borama) agreed very well with MHNG 2394.100, but unfortunately is not part of the specimens examined by PArKer (1932). The original description of U. macfadyeni by Parker is based on only four specimens: the holotype BMNH 1946.8.14.54 (former number: BMNH 1925.4.3.1) from near Berbera, British Somaliland; paratype BMNH 1946.8.14.52-53 (former number: BMNH 1931.8.1.141-142) comprising 2 specimens from Dayah Shabell, 24 miles SE of Berbera; and specimen BMNH 1905.10.30.31 from near Berbera (today in the Museum of Comparative Zoology, Harvard University).

Even PArKer (1942) himself contributed to the nomenclatural confusion by identifying 14 Uromastyx collected in the Guban (northern Somalia) between 10°00'N to 11°15'N and 42°40'E to 43°15'E as U. macfadyeni. He did not state any collection numbers, but the only speci-mens from the Guban in the Natural History Museum London are inventoried under the numbers BMNH 1937.12.5.117-130. With the exception of BMNH 1937.12.5.120 (since 1968 deposited in the San Diego State Collection, USA) all of these specimens were examined, and all proved to belong exclusively to U. ocellata.

Apart from the holotype and the two paratypes still in the Natural History Museum Lon-don, only one additional specimen belonging to the taxon U. macfadyeni could be found there (BMNH 1956.1.6.55 from near Heis, 20 miles W of Mait, Somaliland). The synonymisation of U. macfadyeni with U. ocellata by schäTTi & gAsPereTTi (1994) is obviously based on misidenti-fied U. ocellata from the Guban (WilMs & BöhMe 2000 c). In addition to these findings, AMer & KuMAzAWA (2005) and also WilMs & schMiTz (2007) showed that U. macfadyeni is genetically dif-ferentiated from all other species of the U. ocellata complex and should therefore not be recognised as a member of this complex.

Further to the problem of discriminating U. macfadyeni from U. ocellata, some authors have pointed out that “ocellatus and ornatus [are] nearly indistinguishable” (Moody 1987). WilMs &


BöhMe (2000 c) have shown that in fact both species can be distinguished by two main charac-ters: Firstly, the anterior margin of the ear is always covered by large, triangular scales in U. ornata which are always absent in U. ocellata, and secondly U. ocellata has smaller body scales (U. ocellata: 189-256 scales around mid-body; U. ornata: 149-185 scales around mid-body). For a more de-tailed discussion of the differences between those two taxa, leading to the conclusion that both can be separated at specific rank, see WilMs & BöhMe (2000 c).

Another taxonomic problem arose when examining the relationship between U. ornata and the taxon U. o. philbyi. The main character to distinguish U. o. ornata from U. o. philbyi is the relatively broader tail in the latter (ratio of tail length to maximum tail width at the 5th whorl: U. o. ornata 3.61-5.3; U. o. philbyi 3.03-3.96). WilMs & BöhMe (2000) found that the ranges of both phenotypes broadly overlap in the mountains of western Saudi Arabia, based on six specimens of the narrow-tailed phenotype (U. o. ornata-type) from inside the range of U. o. philbyi, namely: BMNH 1986.436 from Jeddah; BMNH 1975.519 from 21°39'N 39°13'E and BMNH 1979.960 from 21°45'N 39°15'E, both in the vicinity of Burayman; BMNH 1976.1748 from Wadi Fatimah; BMNH 1985.884 from km 115 of the Makkah by-pass, 21°15'N 39°55'E; MHNG 2457.34 from Jebel Hababa (= Jebel Lababa), 18°01'N 42°02'E. These localities are 800-1300 km further south than the type locality of U. o. ornata. According to Arnold (1986), the northernmost locality of U. o. philbyi (Jebel as-Sinfa, 27°57'N 35°47'E) is not far from the type locality of U. o. ornata (al-Muwaylih, 27°41'N 35°27'E).

At Burayman (21°40'N 39°10'E; BMNH 1964.296) and at Jebel Lababa (18°01'N 42°02'E; MHNG 2457.33) both phenotypes are obviously sympatric, likewise in the region of Makkah and Jeddah (U. o. philbyi: MHNG 2536.49, MNHP 4318, BMNH 1985.882, BMNH 1985.884, BMNH 1986.434).

Based on the specimens available, a final decision on the taxonomic status of U. o. philbyi is not yet possible. Three hypotheses are suggested:

– Both taxa (U. o. ornata and U. o. philbyi) are the extremes of an intraspecific cline: U. o. philbyi must be synonymised with U. o. ornata.

– There are no intermediate specimens in the region in question (no gene flow between the two sympatric types): U. o. philbyi must be lifted to specific rank.

– There is a secondary contact zone with gene flow between the two types: U. o. philbyi must be treated as a subspecies of Uromastyx ornata.

As yet, none of these hypotheses can be excluded with certainty, so we suggest that until more specimens and/or ecological data are available, U. o. philbyi should be treated as a subspecies of U. ornata.

Uromastyxaegyptia group

WilMs & BöhMe (2000 a) provide an overview of the taxonomy of the taxa of the U. aegyptia group. Three taxa are considered as valid at specific rank (Uromastyx aegyptia Forskål, 1775, Uromastyx leptieni Wilms & Böhme, 2000 and Uromastyx occidentalis Mateo, Geniez, López-Jurado & Bons, 1998) and the taxon microlepis Blanford, 1874 is assigned as a subspecies to U. aegyptia. The valid-ity of U. aegyptia and U. microlepis was under debate for a long time. The diagnostic features (sensu BlAnford 1874) to distinguish both taxa are the lack of enlarged tubercular scales on the flanks and the presence of skin folds at the sides of the neck covered with tubercles in U. microlepis. The second character has been found insufficient to distinguish both taxa properly (WilMs & BöhMe 2000 a).


schMidT (1939) had pointed out that it is sometimes difficult to distinguish between both forms by the presence or absence of enlarged scales on the flanks, due to the variability of this character in some populations. In the following decades U. microlepis was considered to be a valid species by many authors, and MerTens (1956) suggested that U. microlepis could be a subspecies of U. aegyptia. However, until the 1980s this viewpoint was not adopted by other authors. Moody (1987) suggested U. microlepis to be a synonym of the nominotypic subspecies, because “the only character used in earlier literature to diagnose U. aegyptia (spiny scales on the flanks) is variable throughout the range of both species; and no other diagnostic characters have been discovered”. Moody’s point of view was followed by Joger (1987) and schäTTi & gAsPereTTi (1994). Arnold (1980) supposed that U. aegyptia and U. microlepis might be conspecific, and in subsequent papers (Arnold 1986, 1987) he treated U. microlepis as a subspecies of U. aegyptia. leviTon et al. (1992) and WilMs (1995) adopted this viewpoint and pointed out that it would not be justified to syno-nymise U. microlepis with U. aegyptia before more information on the geographic variability of these taxa was available. Anderson (1999) treated U. microlepis as a synonym of U. aegyptia.

According to WilMs & BöhMe (2000 a), all specimens belonging to U. aegyptia have enlarged tubercles scattered in their flank scalation, whereas most of the examined specimens of U. micro-lepis lack this feature. Some specimens of U. microlepis, especially from the northern parts of the range (e.g. Kirkuk, Iraq), have very few slightly enlarged lateral tubercles. These never reach the size of the tubercles found in U. aegyptia and they have a different distribution on the animals’ body. If present in U. microlepis, the lateral tubercles are much smaller than those in U. aegyptia and restricted to the rear parts of the body just before the insertion of the hind legs, whereas in U. aegyptia they are scattered in the whole flank scalation. Anderson (1999) did not find any tu-bercles on the flanks of the specimen he had examined from the Gulf coast of Arabia.

Furthermore, both taxa have different counts of ventral scales between the gular and inguinal fold. The values are 126-158 (mean: 142) for U. aegyptia and 149-193 (mean: 171.8) for U. mi-crolepis. Additionally, U. microlepis is more colourful than U. aegyptia, with a yellow or greenish colour in warmed-up adult specimens.

Further studies are needed to show whether or not both taxa interbreed at the contact zone of their ranges. This potential contact zone should be expected east of Wadi Araba (Jordan and Pales-tine) and east of Wadi Sawawin in the Jebel as-Sinfa region (Saudi Arabia). The specimens exam-ined from both localities are without any doubt members of U. aegyptia, while specimens from east of that region are exclusively U. microlepis. Because of the overall similarity of both taxa and their parapatric distribution we think it is justified to treat U. microlepis as a subspecies of U. aegyptia, at least until reproductive isolation mechanisms between both taxa are demonstrated.

Another confusing fact was the presence of another Uromastyx species from extreme south-eastern Arabia. We believe that Moody’s (1987) conclusion to synonymise U. microlepis with U. aegyptia was based on data obtained from specimens of U. leptieni which indeed have enlarged lateral tubercles. However, specimens from surrounding areas do not have even slightly enlarged lateral tubercles and therefore fit the description of U. microlepis. Uromastyx leptieni can be distin-guished from U. aegyptia by the combination of the number of ventral and mid-body scales (see WilMs & BöhMe 2000 a: Fig. 2) and by the different colouration and pattern of the juveniles. As has been shown in the present paper, it is possible to discriminate U. leptieni from U. aegyptia using a multivariable approach of morphological datasets. Nevertheless, to further clarify the taxo-nomic situation a molecular approach should be applied.

The third species in this group is U. occidentalis, which is only known from two individuals. The main diagnostic character for this species is the lack of preanal and femoral pores. Without any doubt, U. leptieni is the sister taxon of U. aegyptia.


ACKNOWLEDGEMENTS

We would like to thank H.H. Prince Bandar bin Saud, Secretary General, National Commis-sion for Wildlife Conservation and Development (NCWCD), Riyadh, Saudi Arabia; H.E. Prof. Dr A.H. Abuzinada, former Secretary General, NCWCD; Dr M. Shobrak, NCWCD; Prof. Dr I.A. Nader, NCWCD; Dr H. Tatwani, NCWCD; Mr A. bin Amer Al-Kiyumi, Director General of Nature Reserves, Muscat, Oman and Mr S.M. Al-Saady, Director of Research, Muscat, Oman, for their hospitality and help during the fieldwork of T. Wilms and for issuing the permits re-quired. Special thanks are due to Brien Holmes, al-Ain, UAE and Felix Hulbert, Eltville, Germany for their very pleasant companionship in the field.

The first author would also like to thank I. Barcelo, ERWDA, Abu Dhabi, UAE; Dr C. Drew, Parsons International Limited, Abu Dhabi; Dr I. Galal, Jeddah, Saudi Arabia; Dr A.S. Gardner, formerly Al-Koud, Muscat, Oman now Abu Dhabi, UAE; Dr B. Howarth, al-Ain, UAE; Mr P. Paillat, formerly National Wildlife Research Center (NWRC), Taif, Saudi Arabia now National Avian Research Center (NARC), Sweihan, Abu Dhabi, UAE; Dr C. Tourenq, ERWDA, Abu Dhabi, UAE; P. Vercammen and D. Egan, Breeding Centre for Endangered Arabian Wildlife, Sharjah, UAE and the teams of the NWRC, NARC and the King Khalid Wildlife Research Cent-er, Thumamah, Saudi Arabia, for their hospitality and for the support provided.

We are grateful to the curators of the following museums for their hospitality and/or for the loan of important collections: Dr E.N. Arnold, Dr C. McCarthy and Dr B. Clarke (BMNH); Dr F. Tiedemann (NMW); Dr M. Poggesi and Dr C. Corti (MZUF); Dr V. Mahnert, Dr J. Mariaux and Dr A. Schmitz (MHNG); Dr I. Ineich (MNHP); Prof. F.J. Obst and Dr U. Fritz (MTKD); Dr K. Klemmer and Dr G. Köhler (SMF); Dr J. Hallermann (ZMH); Dr R. Günther (ZMB); Dr U. Gruber and Dr F. Glaw (ZSM). Dr D. Modry (Brno, Czech Republic), Dr F. Krupp (Frankfurt a.M., Germany) and Mr F. Hulbert (Eltville, Germany) and Dr E.N. Arnold (BMNH) provided photographs.

Thomas Wilms is especially grateful to his brother, Matthias Wilms, who shared his knowl-edge on computers and mathematical science, and to the Wünstel family (Reptilium Landau) for providing financial resources.

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zAri, T. 1999. On the reproductive biology of the herbivorous spiny-tailed agamid Uromastyx philbyi in western Saudi Arabia. Zool-ogy in the Middle East 19: 123-130.

Manuscript submitted: 03 April 2006 Manuscript accepted: 14 February 2007

review of the taxonomy of the spiny-tailed lizards of arabia

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