Acanthodactylus montanus, Miralles & Geniez & Beddek & Aranda & Brito & Leblois & Crochet, 2020

Miralles, Aurélien, Geniez, Philippe, Beddek, Menad, Aranda, Daniel Mendez, Brito, José Carlos, Leblois, Raphaël & Crochet, Pierre-André, 2020, Morphology and multilocus phylogeny of the Spiny-footed Lizard (Acanthodactylus erythrurus) complex reveal two new mountain species from the Moroccan Atlas, Zootaxa 4747 (2), pp. 302-326: 316-322

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Acanthodactylus montanus

sp. nov.

Acanthodactylus montanus   sp. nov.

(Appendixes I, III; Figs. 6 View FIGURE 6 , 8 View FIGURE 8 , 9 View FIGURE 9 )

Holotype. Muséum national d’Histoire naturelle (Paris) MNHN-RA-2018.0026 (formerly BEV.11794, tissue sample in the BEV tissue collection, code T5771), adult male, collected on May 12 th 2012, 800 m after the Tizi n’Tichka pass towards Marrakech, southwestern High Atlas, Morocco, 31.2879°N, 7.3824°W (WGS84), 2,176 m a.s.l. by Pierre-André Crochet and Raphaël Leblois ( Fig. 8A View FIGURE 8 ). GoogleMaps  

Paratypes. Nine individuals, BEV.11791 to 11793, 11795 to 11800 (tissue sample codes T5768–5770, 5772– 5777), collected at the type locality at the same date by the same collectors.

Additional material. Eight specimens from El Azib n-Iriri, Jbel Siroua, 30.7470°N, 7.6093°W, 2,319 m a.s.l., collected on April 23 rd 2005 by José C. Brito: BEV.10459–10460 (field number JCB.A544 & A547, tissue samples T11829 View Materials & T11834 View Materials ) and tissue samples + photos T11826 View Materials (JCB.A541), T11827 View Materials (A542)   , T11828 View Materials (A543)   , T11830 View Materials (A545)   , T11831 View Materials (A546)   and T11833 View Materials   (A458).

We identified the following specimens as A. montanus   on the basis of morphology, distribution and ecology: BEV.4973 from 5 km south of Askaoun, Jbel Siroua, 30.69°N, 7.77°W, collected on October 9 th 1974 by Gérard Délye; BEV.4974 from 3 km west of Askaoun, Jbel Siroua, 30.74°N, 7.81°W, collected on October 8 th 1974 by Gérard Délye; BEV.4975 from Tisfaldat mountain, Jbel Siroua, 30.68°N, 7.58°W, 2,460 m a.s.l., collected on March 13 th 1994 by Tahar Slimani.

Lastly, a series of specimens exhibit morphological features of A. montanus   but were apparently collected at a lower elevation than other specimens (around 1,800 m a.s.l.) and slightly south of the main massif of the Jbel Siroua. They are tentatively identified as A. montanus   here (white square in Fig. 10 View FIGURE 10 ) but this identification will require further data and possibly genetic confirmation before it can be firmly established: BEV.4932–4941 and 14898 from 40 km past Tazenakht along road to Taroudant, approx. 30.455°N, 7.558°W, around 1,800 m a.s.l., collected on April 5 th 1960 by Jacques Bons, Georges Pasteur and Bernard Girot.

Etymology. The specific epithet montanus   is a Latin adjective meaning “of mountains, belonging to mountains”, the high mountains of the Western High Atlas being the unique habitat where this species has been found until now.

Diagnosis. A new species of the Acanthodactylus erythrurus   species-group (small flat or carinated dorsal scales; three series of scales around the fingers; three entire supraoculars; 8–10, sometimes 12 straight longitudinal row of ventrals; slightly pectinate toes; undertail and underside of the hind limbs red or reddish in juveniles, subadults and young adults) from the south-western part of the High Atlas and from the Jbel Siroua above 2,000 m a.s.l., characterized by the combination of the following characters: (1) head scalation usually of the “ bellii   ” type (subocular in large contact with the upper lip, wedged between the 4 th and 5 th supralabials), one internasal plate, no scales inserted between the prefrontals, but BEV. T11828 View Materials has no contact between the left supralabial and the lip; (2) a low number of dorsal scales (47–59 longitudinal rows around the body, mean 54.0), femoral pores (17–23, mean 19.9) on each side and subdigital lamellae (19–22, mean 19.9) on the 4 th toe; (3) one or one and a half row of supraciliary granules on each side with a very reduced number (15–46, mean 25.7) of scales and granules around the 2 nd and 3 rd supraoculars on each side; (4) temporals smooth or slightly keeled; (5) dorsal scales smooth or slightly keeled on the neck and progressively turning to distinctly tectiform or sometimes keeled on the back; (6) stocky proportions, especially in adults, with a relatively short tail (57.9–65.9% of the total length, mean 60.8%, almost always longer for all the other Acanthodactylus   of the erythrurus   group except lacrymae   ); (7) base of the tail of adult males very thick; (8) in juveniles, underparts of the tail orange red to intense coral red, slightly paler under the base of the tail, even paler under the rear of the thighs, this colour usually not reaching the rows of femoral pores; (9) some marginal ventral plates yellow of yellowish in adult and subadult males and females, sometimes in juveniles; (10) a series of small round ocelli on the flanks (each one containing 4–9 brightly coloured scales), usually yellow greenish in adults, which tend to disappear completely in adult females.

Body coloration similar to the other members of the Acanthodactylus erythrurus   group. Juveniles are very dark (black or blackish) with three pale continuous lines on each side of the body (one on lower flanks, one at the junction between the dorsum and the flanks and one on the dorsum joining with the opposite one on the tail base) and a dark vertebral area dissected by a paler narrow vertebral area, frequently split on the nape by dark spots or a dark line and disappearing before the base of the tail; one or two series of pale ocelli are found in the dark interspaces between these stripes (juveniles of the other lineages are usually paler and lack the bifurcation of the pale central vertebral area on the nape). Adults are medium brown with faint pale lines on the dorsum and upper flanks (corresponding to the juvenile pattern) and a better marked line on the lower flanks; irregular dark marks are present on the back between the pale lines, sometimes invading the area on the centre of the dorsum. The flanks typically appear darker due to the large dark areas surrounding each pale ocellus. The pileus presents a complex pattern of pale and dark reticulations in juvenile; in adults it is of the same colour as the back with isolated irregular dark spots.

Very similar to the allopatric Acanthodactylus lacrymae   and single individuals are not always possible to separate but differs on average by the following characters: (1) in juveniles, parietals are less clearly striped but exhibit a more reticulated pattern; (2) orange red coloration on the undertail fading less toward tail base and more extensive around the cloacal slit; (3) supralabials usually with diffuse dark vertical stripes (uniformly pale creamy, even in juveniles, in A. lacrymae   ); (4) contrast between the lower border of the flanks and tail side and under surface of the belly and tail usually less marked in adults, especially along the sides of the tail; (5) temporal less flat and dorsal scales slightly killed in adults (smooth in A. lacrymae   except sometimes on the rear of the back); (6) some specimens have one and a half row of supraciliary granules on each side (always one row in A. lacrymae   ) and with a larger number of scales and granules around the 2 nd and 3 rd supraoculars (maximum 46 against maximum 29 in A. lacrymae   ).

Description of the holotype. An adult male ( Fig. 8A View FIGURE 8 ) measuring 60.0 mm of snout-vent length, 14.3 mm of pileus length, 7.7 mm of pileus width, 8.4 mm of head height, 19.5 mm of forelimb length, 31.0 mm of hindlimb length, with missing tail tip, and having the following scalation features: 57 longitudinal rows of dorsal scales at mid-body which are smooth on the neck but carinated on the back, 25 gular scales along a line from the contact between the fourth pair of maxillary scales to the collar, 26 transversal rows of ventral plates from the collar to the anal plate, 21 and 20 femoral pores on the left and right sides respectively, 20 subdigital lamellae beneath the fourth toe, one and a half row of supraciliary granules on each side, with 41 (left) and 46 (right) scales and granules around the 2 nd and 3 rd supraoculars, one entire internasal, no scale inserted between the two prefrontals, two supralabials in contact with the subocular, the latter one in large contact with the upper lip on each side. Base of the tail very thick.

Coloration in life: general ground colour light rufous brown with 3 longitudinal light grey stripes on each side; a series of irregular transverse black marks on the sides of the dorsum between the dorsolateral and dorsal pale lines and dark marks on the edge of the vertebral area between the two dorsal pale lines; a series of 7–8 small pale greenish-yellow ocelli, each surrounded a large black spot that are connected by black coloration, are present along the flanks; pileus roughly of the same colour as the dorsum but more intensely rufous, with very few dark marks; underparts pure white except for the tail and thighs (whitish with pale cream/salmon tinge) and some grey marks on the outer ventrals, 5 spots of yellow-green scales on the outer ventrals/lower dorsals; a poorly contrasted and fragmented dark longitudinal stripe runs along each side of the tail; tail base invaded by yellow-green spots of the same colour as the ones on the flanks and outer belly.

Distribution. Known only from high altitudes (between 2,000 m and 2,500 m a.s.l. but precise elevation known for a very small number of localities only) in the southwestern High Atlas (Tizi n’Tichka) and in the Jbel Siroua massif, 60–70 km southwest of the Tzin-n-Tichka area ( Fig. 10 View FIGURE 10 ). Based on observations of animals with a bellii   -like subocular scalation in the western High Atlas ( Bons and Geniez 1995) and on habitat availability, the distribution probably extends rather continuously from just east of Tizi n’Tichka to Jbel Siroua but maybe not much further east. For example, no member of the Acanthodactylus erythrurus   complex has ever been found at Oukaimeden in spite of the availability of similar habitats and of numerous visits by many herpetologists. If specimens of doubtful identity from between Tazenakht and Taroundant prove to be A. montanus   (white square in Fig. 10 View FIGURE 10 ) the distribution would extend to the northernmost parts of the eastern Anti-Atlas.

Natural history. At the Tizi n’Tichka locality, the habitat was very similar to the habitats occupied by A. lacrymae   further east: earthy slopes with bare gravel ground or short grass interspaced with dense low bushes and some stones. Population density was rather high and the animals were relatively easy to catch (compared to many populations of the IM clade) due to their more confident behaviour and slower escape. The animals were active in sunny and calm weather and their behaviour was typical of the A. erythrurus   complex, actively foraging in the open and retreating to the shelter of bushes when disturbed.