(Lacerta Erythrura Schinz 1833, and Acanthodactylus Vulgaris
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HERPETOLOGICAL JOURNAL, Vol. 5, pp. 271-280 (1995) CONTRIBUTION TO THE SYSTEMATICS OF THE LIZARD ACANTHODACTYLUS ER YTHR UR US (SAURIA, LACERTIDAE) IN MOROCCO JACQUES BONS AND PHILIPPE GENIEZ Laboratoire de Biogeographie et Ecologie des Ve rtebres, Ecole Pratique des Ha utes Etudes, Un iversite Montpellier II, 34095 Mo ntpellier Cedex 05, France We have analysed several scalation characters and the geographic distribution of lizards of the Acanthodactylus erythrurus group to verify the validity of these criteria. These data are collated with biogeography to demonstrate the existence of two distinct species within what are known as common fringe-toed lizards: Acanthodactylus erythrurus, consisting of three subspecies, and Acanthodactylus lineomaculatus, monotypic and endemic to Morocco. Hypotheses concerning the population history of these animals are proposed. INTRODUCTION coast. However, it does not appear to be known from Tunisia. At the beginning of the last century common fringe Morphologically, it is characterised by the presence toed lizards were described from Europe (Lacerta of three complete rows of scales round the fingers and erythrura Schinz 1833, and Acanthodactylus vulgaris small dorsal scales, either keeled or unkeeled, on the Dumeril & Bibron 1839) and from North Africa back, with the underside of the tail bright red in juve (Acanthodactylus lineomaculatus Dumeril & Bibron niles and subadults. The combination of these three 1839). Since then, they have been the subject of many characters distinguishes Acanthodactylus erythrurus investigations. Boulenger' s revision in 1878 was the within the genus. Three subspecies have been recog first attempt to introduce some order to the genus nised in Morocco (Pasteur & Bons, 1960; Bons & Acanthodactylus, often considered as one of the most Girot, 1962): difficult in the Palearctic. The increase in the number - ssp. belli Gray 1845, in the Rif, on the coast and of specimens collected, and the use of often contradic plateaux east of the Atlases, the southern slopes of the tory terminology by different authors, led Salvador High Atlas and the far west of the Anti-Atlas (this sub (1982) and Arnold (1983) to make a revision of the species is the only representative of Acanthodactylus numerous species that this genus comprises and to erythrurus in Algeria; Salvador, 1982). clarify the phylogenesis. Other authors have confined - ssp. atlanticus Boulenger 1918, in the Middle their research to a single group, e.g. Squalli-Houssaini Atlas, the northern slopes of the High Atlas and the (1991) on the fringe-toed I izards of the erythrurus plains situated north of the High Atlas (endemic to group. Morocco). Since we had access to abundant material from - ssp. lineomaculatus Dumeril & Bibron 1839 on North Africa and precise data on these animals' ecol the Atlantic coast from Tangiers to Essaouira (ende ic ogy and distribution in Morocco, our intention was to � to Morocco). readdress the problem, restricting our analysis to cer The nominate subspecies is confined to the Iberian tain scalation characters that, in our opinion, are peninsula. discriminative. These characters were then collated The two recent revisers of the genus, Salvador with the ecological and biogeographical characteristics ( 1982) and Arnold( 1983 ), do not retain the subspecies of the populations being studied. We restricted our study to the different forms of the common fringe-toed atlanticus. They consider it as intermediate between A. erythrurus belli and A. erythrurus lineomaculatus. lizard in Morocco because it is the country where the . Whereas Squalli Houssaini (1991) , without adopting a widest diversity of forms and habits is found. definite position, considers that the Moroccan subspe The common fringe-toed lizard, Acanthodactylus cies have little taxonomic value and are only a erythrurus (Schinz 1833), is the only representative of reflection of their distribution, and that the Iberian the genus occuring in Europe, where it is confined to fringe-toed lizards are sufficiently differentiated from the southern two thirds of the Iberian Peninsula ex their Moroccan counterparts to merit a distinct specific tending its range as far north as Gerona (Barbadillo status. Escriva, 1987), Zaragoza, Burgos (old record) and Leon (old record) (Salvador in Bohme, 1981 ). It is also MATERIALS AND METHODS the only fringe-toed lizard to occupy the whole of Mo We examined 496 Moroccan individuals from 22 rocco, north and west of the Atlases. This distribution localities, or groups of localities, throughout the coun stretches eastwards along a large part of the Algerian try. For comparative purposes, we added 11 individuals 272 J. BONS AND P. GENIEZ ,• t . SPAIN I + I ALGERIA e Mamora Forest e Zaen; Forest (franc e e Region of Lakes • Azrou Tarmilate, Oulm�s e • Tioumliline, Agdal Plateau e Doukkala MO ROCCO e Haouz, Marrakech Essaouira • • Jbel Saghro Jbel Siroua e 40 km after Tazenakht - Taroudannt 0 100 200 km. Oued Noun FIG. I. Location of populations studied. from Andalusia (Spain) (Fig. 1). The samples varied in Jbel Siroua (2 individuals) size depending on the localities, ranging from 1 to 69 Oued Noun ( 1 individual) specimens (average : 22.55). The examined samples are as follows : SPAIN Benidorm (1 individual) MOROCCO Almeria (2 individuals) Larache (47 individuals) Hues car (2 individuals) Mehdiya, Kenitra (36 individuals) Torre de la Higuera (3 individuals) Mamora Forest ( 41 individuals) unknown locality (3 individuals) Rabat-Agdal (55 individuals) Temara, Sables d'or (14 individuals) Five scale characters were monitored: (1) dorsal Zaers Forest (47 individuals) scales strongly keeled or not; (2) position of the Casablanca (47 individuals) subocular in relation to the edge of the upper lip; (3) Doukkala (4 individuals) number of scales and granules arising from the frag Essaouira (6 individuals) mentation of the fi rst supraocular, right and left; ( 4) Region of Lakes ( 5 individuals) number of interprefrontal granules; and (5) internasal Ifrane (3 4 individuals) divided or not. Azrou (17 individuals) We then compared the percentage of individuals Tioumliline, Agdal Plateau (9 individuals) within each population that presented different con Tarmilate, Oulmes (69 individuals) figurations of each character. Haouz, Marrakech (3 individuals) The specimens examined come from the collection Rif (18 individuals) of the Laboratoire de Biogeographie et Ecologie des Eastern Morocco (13 individuals) Vertebres de l'E.P.H.E., Montpellier, France. The re Eastern High Atlas ( 13 individuals) sults of these examinations were collated with Jbel Saghro ( 4 individuals) numerous observations made in the field in Morocco 40 km W. of Tazenakht (11 individuals) and the Iberian Peninsula. SYSTEMATICS OF ACANTHODACTYLUS 273 ( SPAIN e SPAIN � ALGERIA \ .�o�\ ALGERIA •�-�JI e -,-.ii0 MoRocco �- -•I .- -� MOROCCO • 0 ·�· j . �j= • .. ' ...• •• 00 o • FIG. 2. Texture of dorsal scales, expressed in percentages of individuals per site. RESULTS The results of the pholidotic analyses are ordered according to scale characters. DORSAL SCALATION (FIG. 2) 2 An examination of the dorsal scales reveals a parti tion of our samples into two groups: (I) lizards from FIG. 3. Position of subocular scale, expressed in percentages the Atlantic coastal area, between Tangiers and of individuals per site. Essaouira, possess strongly and sharply keeled dorsal scales, starting on the back and sides of the neck; (2) Position 4 : The subocular is in wide or narrow contact lizards from all other localities (Morocco and Spain) with the upper lip. This subocular/edge of lip contact is possess smooth scales on the anterior part of the back. characteristic of the form be/Ii. In some populations they become tectiform or weakly The analysis of our samples shows that only animals keeled at the rear of the back. from the Rif, the southern slopes of the High Atlas and the We did not observe any individuals presenting in east of Morocco are 100% consistent with position 4. termediate characters between these two types of Position 3 is characteristic of animals from the Mid scalation, nor any mixed populations. Consequently, dle Atlas and the Haouz plain, in percentages varying strongly keeled dorsal scales make it possible to distin from 87 to 100%. The other situations are infrequent in guish with certainty Atlantic coast animals fromall the these regions: 0 to 8.7% of position 1, 0 to 20% of posi other common fringe-toed lizards examined from Mo tion 4. rocco and Spain. A later examination of a sample (c. The coastal animals are distributed along a band 30 individuals) from the Aures in Algeria, put at our running north-east/south-west. In localities north of disposal by Laurent Chirio, also confirmed our find Rabat they mostly (75 to 95%) possess a small ings for Algeria. trapezoidal upper labial (position 2) that is usually ab sent (57.1 to 95%) in specimens from south of Rabat POSITION OF THE SUBOCULAR (FIG. 3) (position 1 ). Four positions of this scale in relation to the upper Consequently the traditional criterion of the posi lip were observed. This is the most commonly used tion of the subocular used to distinguish subspecies of character for the recog1.1ition and distinction of the Acanthodactylus erythrurus should be used with care. North African subspecies (Bons & Girot, 1962). Position 4 certainly enables animals from the Rif, east Position 1: The subocular is wedged between the 4th ern Morocco, the southern slopes of the High Atlas