taxonID	type	description	language	source
03C1AF1C831CFFF2FCCC3D071265FE1F.taxon	distribution	Distribution (Fig. 13). In the Sahara, south-eastern Algeria, north-eastern Mali, northern Niger, northern Chad, northern Sudan, Egypt, Libya, and southern Tunisia (Salvador, 1982; Nouïra, 1996; Ch. P. Blanc, pers. com.; this study). The characters given above and a careful examination of the pictures published by Bons & Girot (1964) lead us to reject the occurrence of this species in Morocco (cf. Bons & Geniez, 1996), in opposition to the opinions of Bons & Girot (1964), Mellado & Olmedo (1990) and Pérez Mellado (1992). The species is also encountered in Israel, Arabia and Iraq (Salvador, 1982). Diagnosis. (Remarks: the following diagnosis is based on specimens of Acanthodactylus scutellatus audouini only). The largest species of the group (reaching 72.5 mm snout-vent length, mean = 63.9; cf. Table 2, Figs 14 and 15). The subocular in contact with three or (rarely) four supralabials in 99 % of the individuals (Table 3) distinguishes A. s. audouini from A. aureus and A. taghitensis. Dorsal scales usually small and rather numerous (in the Sahara, range: 39 – 69, mean: 53.4; Table 2, see remarks below under ‘ Geographical variation’), nearly equal in size between the dorsum and the flanks, and moderately to strongly carinate (codes 4 or 5 in 86 % of our sample, no individual reaches code 6; Table 3, see Fig. 16). Fewer than two rows of supraciliary granules in 70 % of the specimens, two rows in 28 %, more than two rows in only 2 % (Table 3). Generally 13 or 14 longitudinal rows of ventral scales (in 83 % of the specimens, range: 12 – 16, mean: 13.9; Table 2). Femoral pores rather numerous (range: 16 – 26, mean: 21.1; Table 2). Adult males with a black, highly contrasting, dorsal reticulation. Females with isolated black spots at regular spaces on the back. Pale dorsal spots often lacking in adults, which have a dorsal coloration made of only two colours; most individuals have black spots on their pileus. For separation of A. s. audouini from A. longipes and A. senegalensis, see these species. Acanthodactylus scutellatus audouini can be distinguished from the closely similar A. dumerili by its larger maximum size and by a different pileus coloration. In A. dumerili, as in the remaining species within the scutellatus group, the pileus is either uniformly pale or marked with darker vermiculations or small dots, but not with distinct and well-individualized large black spots as in A. s. audouini. In addition, a higher proportion of adults A. s. audouini have a dorsal coloration consisting of a dark pattern on a uniform background (two colours only on the dorsum, COUL code 4, in 27 % of the specimens; Table 3). When other species within the scutellatus group have only two colours on the dorsum, it is usually light spots on a darker ground colour (code 2, Table 3). According to Nouïra (1996; p. 246), A. s. audouini and A. dumerili also differ in the fragmentation of the cephalic plates, A. s. audouini having the first supraocular (SO 1) usually separated from the second supraocular (SO 2) by a row of proximal granules and a strongly fragmented fourth supraocular (SO 4). We checked the validity of these proposed differences on 116 A. s. audouini (most of them from Tunisia) and 298 A. dumerili. In A. s. audouini, 48 % of the individuals have SO 1 and SO 2 partly or totally separated (including 18 % with completely separated SO 1 and SO 4; Table 4), whereas in A. dumerili only 7 % of the specimens have SO 1 and SO 2 partly separated (none having SO 1 and SO 4 completely separated; Table 4). In A. s. audouini, 30 % of the specimens have an entire SO 4, 33 % have a partially fragmented SO 4 and 37 % have a completely fragmented SO 4. In A. dumerili, 49 % of the specimens have an entire SO 4, 38 % a partially fragmented one, and only 14 % a completely fragmented SO 4 (Table 4). The differences described by Nouïra (1996) are thus real but of limited use when trying to identify single specimens. Geographical variation. Three subspecies have been recognized. The subspecies hardyi is recognized as a valid taxon in most of the recent works on the genus (e. g. Bons & Girot, 1964; Salvador, 1982; Arnold, 1986; Nouïra, 1996). Harris & Arnold (2000) even propose to give specific status to this taxon, although without justification. In addition, Bons & Girot (1964) recognized the subspecies audouini, based on an analysis of large series of specimens. We follow their opinion here, based on a preliminary examination of a number of specimens from Egypt and Israel. Subspecies. Acanthodactylus scutellatus scutellatus in Israel, Sinai and Egypt.	en	Crochet, Pierre-André, Geniez, Philippe, Ineich, Ivan (2003): A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications. Zoological Journal of the Linnean Society 137 (1): 117-155, DOI: 10.1046/j.1096-3642.2003.00044.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2003.00044.x
03C1AF1C831CFFF2FCCC3D071265FE1F.taxon	description	A. s. hardyi Haas, 1957	en	Crochet, Pierre-André, Geniez, Philippe, Ineich, Ivan (2003): A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications. Zoological Journal of the Linnean Society 137 (1): 117-155, DOI: 10.1046/j.1096-3642.2003.00044.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2003.00044.x
03C1AF1C831CFFF2FCCC3D071265FE1F.taxon	description	We retain for the time being the subspecies audouini for the Saharan specimens, characterized by a lower number of more strongly keeled dorsal scales (73 on average in scutellatus (Bons & Girot, 1964) against 53 on average, and not more than 68 in audouini [own data]) and a smaller size (Boulenger, 1921; Bons & Girot, 1964). A revision of the material included in Acanthodactylus scutellatus is needed before any firm conclusion can be drawn. The limits of the range of A. s. scutellatus and A. s. audouini follow Bons & Girot (1964). There is a slight morphological variation among populations of Acanthodactylus scutellatus audouini in the Sahara. In Tunisia, most of the individuals have a grey throat and a reddish tail, whereas this coloration is exhibited, more or less markedly, by only a small number of individuals from southern Sahara. Remarks. Victor Audouin published the description of Lacerta scutellata in 1827 and not in 1809 (Salvador, 1982; according to his birth date, V. Audouin was only 12 years old in 1809 [R. Bour, pers. com.]) or 1829 (Arnold, 1983; Brygoo, 1988).	en	Crochet, Pierre-André, Geniez, Philippe, Ineich, Ivan (2003): A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications. Zoological Journal of the Linnean Society 137 (1): 117-155, DOI: 10.1046/j.1096-3642.2003.00044.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2003.00044.x
03C1AF1C8314FFF6FF543BD815D0F992.taxon	description	Chresonyms. Acanthodactylus longipes Boulenger, 1918: Bons & Girot, 1964: 324; Salvador, 1982: 132; Arnold, 1983: 324. Distribution (Fig. 19). Most of the Sahara from coastal Mauritania (this study; A. Foucart, pers. com.), southern Morocco (Tafilalet, Iriki) (Geniez & Soto, 1994), northern Mali, northern Niger, northern Chad, Algerian Sahara, Tunisia, Libya (Salvador, 1982; this study), to Egypt (Baha El Din, 1994). Diagnosis. Medium-sized species (reaching 61 mm snout-vent length, mean: 52.4). The subocular in contact with three or (rarely) four supralabials in 100 % of the individuals (Table 3) distinguishes Acanthodactylus longipes from A. aureus and A. taghitensis. Dorsal scales small, very numerous (range: 55 – 77, mean: 66.4; Table 2), elongate, smooth except in the vertebral area where they can be weakly to moderately keeled (code 3 or less in 97 % of the individuals, no individual reaching code 5; Table 3, see Fig. 16 d). Two rows or more of supraciliary granules in 86 % of the individuals (Table 3, see Fig. 20 b). Large number of longitudinal rows of ventral plates (15 or more in 95 % of the individuals, range: 13 – 19, mean: 16.1; Table 2) arranged in oblique rows. Large number of femoral pores (range 17 – 28, mean = 21.5; Table 2). Coloration distinctive: flanks have a mottled pattern which usually tends to disappear towards the vertebral area (Fig. 20 a). Females can have small, regularly disposed spots. Red spots can occur on the dorsum. Pileus weakly vermiculated with red. The combination of dorsal pattern and scale structure gives to the skin of A. longipes a fragile and translucent aspect. This species is further characterized by an elongate and pointed snout compared to A. scutellatus audouini, A. dumerili and A. senegalensis. See A. senegalensis for additional differences from that species. Several scalation characters separate A. longipes from the broadly sympatric A. dumerili and A. s. audouini. A strongly fragmented fourth supraocular is found in 73 % of the A. longipes individuals but in 14 % of the A. dumerili and 37 % of the A. s. audouini. Granules are often inserted between the parietal plates in A. longipes, which is exceptional in A. s. audouini and A. dumerili. The number of longitudinal rows of ventrals is often nearly diagnostic: most A. longipes (81 % of the specimens) have 16 ventrals rows or more, which is extremely rare in A. s. audouini (1 % of the specimens) or A. dumerili (less than 1 % of the specimens). Acanthodactylus longipes is the only species to possess dorsal scales small, elongate and smooth except in the vertebral area. The dorsal scales in the vertebral area are only weekly keeled, a further distinction from most A. s. audouini and A. dumerili: 84 % of the A. longipes individuals have CARE code 2 or (rarely) 1, whereas less than 2 % of the A. dumerili specimens have CARE code 2 or 1. The typical dorsal pattern is also characteristic. It should be noted, however, that some A. longipes females present a uniformly pinkish coloration with small dark spots, making them similar to A. dumerili or A. s. audouini, although their pattern is less contrasting than in these species. In conclusion, although no single character is fully diagnostic between A. longipes and the sympatric species, a combination of several scalation and coloration features will enable to identify the vast majority of specimens. In ambiguous cases, the elongate and pointed shape of the snout and the structure of the dorsal scales are often useful, although occasional individuals remain impossible to identify safely. It should be stressed than other characters proposed by earlier authors (number of supralabials in contact with the subocular, length of hindlegs) proved to be useless. Geographical variation. The subspecies panousei, described from south-eastern Morocco, does not seem to be valid (Salvador, 1982; own results). In some populations from the eastern part of the species distribution, specimens tend to have more strongly carinate dorsal scales.	en	Crochet, Pierre-André, Geniez, Philippe, Ineich, Ivan (2003): A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications. Zoological Journal of the Linnean Society 137 (1): 117-155, DOI: 10.1046/j.1096-3642.2003.00044.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2003.00044.x
03C1AF1C8314FFF5FCA43D8314B1F9B1.taxon	distribution	Distribution (Fig. 21). Saharan oceanic fringe from Agadir (Morocco) to the Cap Blanc Peninsula (Western Sahara / Mauritania). The species is further encountered at two points in Mauritania (Salvador, 1982) and three stations of the Senegal coast (Böhme, 1978; Salvador, 1982). It reaches as far as 200 km inland in Western Sahara (pers. obs.). The animals from Fderik (Mauritania) are referable to Acanthodactylus taghitensis. The Atar locality (specimen BMNH 1982.294) is wrongly positioned (far inland) in Salvador (1982) and Arnold (1983). It is in fact situated near Nouadhibou, i. e. near the coast. A. aureus is the most common lacertid on the whole oceanic side of Western Sahara. Diagnosis. A species of variable size according to populations (reaching a maximum snout-vent length of 65 mm, mean: 53.65). Only two supralabials in contact with the subocular in 99 % of the specimens, as a result of the fusion of the third and fourth (rarely the fourth and fifth) supralabials (Table 3). Dorsal scales relatively large, not very numerous (range: 38 – 59, mean: 46.7; Table 2), slightly larger on the dorsum than flanks, pyramidal, and obtusely carinate (CARE code 3 or 4 in 99 % of the individuals; Table 3). Fewer than two rows of supraciliary granules in 97 % of the individuals (Table 3). Generally 14 or (less often) 13 longitudinal rows of ventrals (in 77 % of the specimens, range: 12 – 17; mean: 13.9; Table 2). Femoral pores rather numerous (range: 19 – 26, mean: 21.2; Table 2). Dorsal coloration distinctive, comprising dark rectangular blotches longitudinally aligned over light longitudinal lines. In old males, however, the dark blotches and the light lines can produce a mottled or even reticulate pattern. A golden yellow tone appears on the dorsum of males during reproduction. Further characterized by a concave forehead following a short but strongly acuminate snout. Supralabials sometimes uniformly greyish or purplish-blue. Easily separated from A. scutellatus, A. senegalensis, A. dumerili and A. longipes by the peculiar conformation of the supralabial scales bordering the subocular. The exceptional individuals lacking this feature can be recognized by the typical habitus of the species, resulting from its distinctive colour pattern and peculiar head profile. For distinction from A. taghitensis, see that species. Geographical variation. None documented.	en	Crochet, Pierre-André, Geniez, Philippe, Ineich, Ivan (2003): A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications. Zoological Journal of the Linnean Society 137 (1): 117-155, DOI: 10.1046/j.1096-3642.2003.00044.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2003.00044.x
