Review of the taxonomy of the
spiny-tailed lizards of Arabia
(Reptilia: Agamidae: Leiolepidinae: Uromastyx)
Thomas M. Wilms and Wolfgang Böhme
FAUNA OF ARABIA 23: 435–468 Date of publication: 15.07.2007
Abstract : 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 Uromastyx leptieni.
مراجعة تصنيفية
للسحالي الذيل في شبه الجزيرة العربية
(الزواحف العظائيات: Agamidae: Leiolepidinae Uromastyx)
توماس م. ويلمز وولفاجانج بومة
ملخص: يوجد حاليًا في شبه الجزيرة العربية 6 أنواع من الجنس Uromastyx (الزواحف العظائيات: Leiolepidinae) تتمثل في 3 أنساب من تطورية وهي U. yemenensis و U. benti, U. leptieni, U. ornata, U. thomasi, Uromastyx aegyptia يقدم البحث مراجعة لتصنيف هذه السحالي ويعرض نتائج جديدة تتعلق بالشكل والبيئة و النوع Uromastyx leptieni.
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 genetic 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 represent 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. ornata). 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.
Fig. 1: Distribution range of the genus Uromastyx (after Wilms 2001).
Fig. 3: Distribution of Uromastyx aegyptia aegyptia and Uromastyx aegyptia microlepis (after Wilms & Böhme 2000 a). For the distribution 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.
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.
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 populations west of the left dotted line belong to U. a. microlepis and those east of the right dotted line belong to U. leptieni; the populations between both lines (indicated by question marks) need further study to clarify their taxonomic status. — Localities: 1: Shanayel; 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 photographs 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).
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.
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 molecular 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 morphologically it is grouped within the Afro-Arabian U. ocellata group. Another difference between the two approaches is the placement of the U. aegyptia group within the cladogram. Morphologically, 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.
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