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(Reptilia, Agamidae) in Western Sahara: De Rev. Esp. Herp. ( 1998) 12:97-109 97 Chorological analysis and morphological variations of Saurians of the genus Uromastyx (Reptilia, Agamidae) in western Sabara. Description of two new taxa. 1 2 3 JOSÉ ANTONIO MATE0 ·3, PHILIPPE ÜENIEZ , LUIS FELIPE LÓPEZ-JURAD0 & JACQUES BONS2 I Estación Biológica de Doñana-CSIC, Apartado 1056, E-4108 Sevilla, Spain. 2laboratoire de Biogéographie et Ecologie des Vertébrés-EPHE, Université Montpellier 2, Place Eugene Bataillon, F-34095 Montpellier, France. 3 Departamento de Biología, Universidad de las Palmas de Gran Canaria, 35017 las Palmas de Gran Canaria, Spain. E-mail: luisfelipe. lopez@biologia. ulpgc. es Abstract: The description of a new species ofthe genus Uromastyx is proposed on the basis oftwo specimens from the Adrar Souttouf in Western Sahara. This taxon differs greatly from U. acanthinura on account its larger size, the much larger number of scales, the arrangement of tubercules on its upper thighs, the different habitus and colouring. These morphological features mean it closely resembles U. aegyptia. The existence of a relictual U. aegyptia-group throughout the Sahara is suggested. In addition, the morphological variations in Spiny-tailed agamas (or Mastigures) ofthe Uromastyx acanthinura group in the west ofthe Sahara are briefl y analysed. This produces evidence for the existence of a species proper to Western Sahara and surrounding areas, Uromastyx jlavifasciata, represented by two subspecies: U. f jlavifasciata in the north and U. f obscura subsp. nov. in the south. The latter new form is characterised by uniformly black colouring, even in active individuals. This work also demonstrates that Uromastyx acanthinura werneri does not penetrate Western Sahara and that its distribution is parapatric with that of U. jlavifasciata. Lastly, the presence of U. maliensis is suspected in the Adrar Atar (Mauritania) and the Adrar Souttouf (Western Sahara). Key words: Agamidae, Uromastyx, chorology, systematics, Sahara. Resumen: Análisis corológico y variaciones morfológicas en los saurios del género Uromastyx (Reptilia, Agamidae) de la región occidental del Sahara. Descripción de dos nuevos taxones.- Se describe una nueva especie del género Uromastyx sobre la base de dos especímenes procedentes del Adrar Sutuf (Sahara Occidental). Los individuos del nuevo taxón difieren fuertemente de Uromastyx acanthinura por el gran tamaño que llegan a alcanzar los adultos, su siempre mayor número de escamas, la disposición diferente de los tubérculos en los muslos, y por su diseño y coloración. Sus características generales se asemejan a las de Uromastyx aegyptia, un hecho que sugiere la posible existencia de lagartos del complejo U. aegyptia en poblaciones relícticas a lo largo del desierto del Sahara. Se han analizado también las variaciones morfológicas de los lagartos de cola espinosa del grupo Uromastyx acanthinura en la región occidental del Sahara, cuyos resultados ponen de manifiesto la existencia de una especie propia al Sahara Occidental y sus alrededores, Uromastyxjlavifasciata, representada por dos subespecies: U.fjlavifasciata en el norte y Uf obscura subsp. noven el sur. Esta última forma se caracteriza por su coloración negra homogénea incluso en animales activos. Este trabajo también demuestra que Uromastyx acanthinura werneri no penetra en el Sahara Occidental y que su distribución es parapátrica con U.jlavifasciata. Finalmente, nuestras observaciones también dan como posible la existencia de Uromastyx maliensis en el Adrar Atar (Mauritania) y en el Adrar Sutuf (Sahara Occidental). Palabras Clave: Agamidae, Uromastyx, corología, sistemática, Sahara. INTRODUCTION ran wind- dunes) the two can be seen to com­ plement each other quite closely (Figure 1). The genus Uromastyx, of the family As SCHLEICH et al. ( 1996) point out, it is clear Agamidae and the subfamily Uromastycinae, that Spiny-tailed agamas (or Mastigures) comprises ten species (WILMS, 1995;JOGER& avoid the ergs which may well constitute LAMBERT, 1996). lt has a typically Saharo­ barriers for these animals (GRENOT & VER­ Sindian range covering the whole of the Sa­ NET, 1972). As is evidenced by the distribu­ hara, Somalia, Ethiopia, the Arabian penín• tion maps of the genus published by WILMS sula, Palestine, Syria, lrak, lran, southern Af­ ( 1995), the emplacement of these ergs pro­ ghanistan, Pakistan and north-west India bably gave rise to the isolation of Uromastyx (WILMS, 1995). In Africa, lizards ofthe genus in the west ofthe Sahara from those in central Uromastyx are not evenly distributed through­ Sahara. The range of Spiny-tailed agamas in out the Sahara and there are whole areas the west Sahara corresponds to an area orien­ where this animal is unknown. lf the range of ted north-east / south-west starting in south­ the genus Uromastyx in the western half ofthe east Morocco and extending south-westwards Sahara is compared with a map of ergs (Saha- over the entire foothills of the Moroccan Anti- 98 MATEO et al. 20· 1o·E :.i - c ·-1\-.• r_ ( - • .¡; ./ --a.__,--.. a ·~r?¿.F7;· •.• '·<--..-·" 1 'lit., .. > • 'i:7''t. z " , ·*~••,.. ~· ~""';i..~!i ,,,.~,rr>~:~i~~ ..., ,,,.il~... ~. ... \,..··:' l it~~~~:.., ... .,· .•;; •• o 1>o<?a • • • • .., . ~~.. • !,:.. ,, • . / !':·-'_.;, .. ., .-,-;...-:.,·.··,•,~ • .-­ ... ~··· ': ...... • • - )]f,~ff!;;)r? •• .,= • • • z • • <:) •• •• / ·-·-- ·-- . --- ./ , . - · - · ___. ..-. 10W o 10 E Figure 1.- Distribution map of ergs (in grey) and of Uromastyx (full circles), al i taxons confused. The dashed line is the isoline for 200 mm of anual rainfall. Figure 1.- Disposición de los ergs (punteado) y de los lagartos del género Uromastyx (puntos negros). La línea punteada corresponde a la isoyeta de 200 mm anuales. Atlas, and covering Western Sahara, north­ nise any particular forms and ascribe the east Mauritania and the Adrar Atar region also forms nigriventris Rothschild & Hartert, 1912, in Mauritania. lt is limited to the north-west nigerrimus Hartert, 1913, werneri L. Müller. by the Moroccan Atlas axis, to the west by Sa­ 1922, andpluriscutata Féjerváry, 1927, toma­ haran zones under strong oceanic influences nifestations of the species' considerable mor­ (Saharan bioclimatic zone with warm win­ phological and chromatic variation. They do ters), to the south and east by a succession of not comment on the chromatic differences, ergs which probably constitute an impassable observed by VALVERDE ( 1957), between barrier: the Trarza and Ouarane sands ( cf. AN­ animals from south Morocco and those from GEL & LOTHE, 1938), the Majabat al-Koubra the Western Sahara. In VALYERDE's opinion, (cf. MONOD, 1958), the Iguidi, Chech and Er­ Spiny-tailed agamas from the former Spanish Raoui ergs and, lastly, the Great Western erg colony are sufficiently distinct to deserve a (Figure l ). separate subspecific status. Five years later, MERTENS ( 1962) suggested the existence of Historical background to the systematics of three subspecies in the western area: Uromas­ Spiny tailed agamas (or Mastigures) in the tyx acanthinura acanthinura Bell, 1825, for west of the Sabara north-west Africa, U. a. werneri for north­ In the north ofthis western region, i1.e. Mo­ westAlgeriaand eastern Morocco(wadi Mou­ rocco, PASTEUR & BONS ( 1962) do not recog- louya valley) and U. a. flavifasciata Mertens, Uromastix FROM WESTERN SAHARA 99 1962. This last form was described on the nomy ofSpiny-tailed agamas from the west of basis of a specimen which this author reported the Sahara and, in particular, that in the Wes­ to have come from 50 km. north-east ofDakar tern Sahara this is almost completely un­ (Senegal). MERTENS (1962) also mentioned known. The examination of a large sample of the existence of this taxon in Western Sahara Spiny-tailed agamas from the west of the Sa­ (after the illustrations published by VALVER­ hara has enabled us to demonstrate the impor­ DE, 1957), and recalled the occurence of U tance of this region for understanding the phy­ acanthinura in the Adrar ofMauritania but he logeography of this genus, the existence of a stated that it had not yet been established whe­ new species ofthe genus Uromastyx as well as ther or not these Mauritanian populations large morphological and chromatic variations belonged to the subspecies flavifasciata. The within the Uromastyx acanthinura group. terra typica of U a. flavifasciata was rightly questioned by BóHME ( 1978) who suggested MATERIAL ANO METHODS locating it at Atar, in the Adrar of Mauritania, but without giving reasons for this, possibly We examined 154 Spiny tailed agamas the result of a report by GRENOT (1974, from the following scientific collections: Esta­ p.162). Probably following the two latter au­ ción Biológica de Doñana-Sevilla, (E.B.D.), thors, WILMS & BóHME (1993) and SCHLEICH (n=75), Muséum National d'Histoire Nature­ et al. ( 1996) recognised the existence of Uro­ lle de Paris (M.N.H.N.), (n= 40), Laboratoire mastyx a. flavifasciata in the Adrar Atar. de Biogéographie et Ecologie des Vertébrés Within the species Uromastyx acanthinura, de l'E.P.H.E., Montpellier (E.P.H.E), (n= 37), WILMS & BóHME (1993) recognised four sub­ Departamento de Biología University of Las species: Uromastyx a. acanthinura in north­ Palmas (U.L.P.G.C), (n= 2). Of these 154 west Sahara; U a. geyri in south central Sa­ animals, 124 come from the area of the pre­ hara; U a. dispar in southern east Sahara; and sent publication, namely 60 from Morocco U a. flavifasciata only in the Atar region of and north-west Algeria, 58 from Western Mauritania. Curiously, WILMS ( 1995) consi­ Sahara, 5 from Mauritania and one from the dered Uromastyx acanthinura flavifasciata Hamada of Tindouf in Algeria. The other 30 synonymous with the nominal form and individuals were from regions and taxons because of lack of information he did not which are outside the framework of this study; pronounce on the status of U a. werneri. At they were used for purposes of comparison. the same time, he omitted from his U They are Uromastyx a. acanthinura (2 speci­ acanthinura distribution map the whole of mens from central and east Algeria), U a. Western Sahara, a region where Spiny-tailed geyri (8 individuals from south Algeria and agamas are well represented ( cf.
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