Trapelus boehmei, Wagner & Melville & Wilms & Schmitz, 2011

Wagner, Philipp, Melville, Jane, Wilms, Thomas M. & Schmitz, Andreas, 2011, Opening a box of cryptic taxa - the first review of the North African desert lizards in the Trapelus mutabilis Merrem, 1820 complex (Squamata: Agamidae) with descriptions of new taxa, Zoological Journal of the Linnean Society 163 (3), pp. 884-912 : 900-904

TRAPELUS BOEHMEI SP. NOV.

Holotype: ZFMK 49751 : Morocco, between Akka and Icht; leg. W. Bischoff & U. Joger, 27.v.1988 ( Fig. 9).

Paratypes: ZFMK 49752–754 : Morocco, between Akka and Icht ; leg. W. Bischoff & U. Joger, 27.v.1988 ; ZFMK 49664 : Algeria, Colomb-Bechar ; leg. W. Bischoff & U. Joger, 13.v.1988 ; MNHN 2006.0343 : Mauritania, about 220 km south of Nouadhibou on the main road to Nouakchott; leg. I. Ineich, 21.vii.2005; ZMB 52277 : Morocco, 40 km north of Zagora ; leg. M. Barts, iv.1993 ; ZSM 207/1993: Morocco, 10 km south-east of Goulmima ; adult female; leg. H. H. Schleich, vi.1993 ; ZSM 225/1993: Morocco, 25 km west of Tissint , adult male; leg. H. H. Schleich, vi.1993 ; ZSM 688/1979/1-5: Morocco, Ksar es Souk (W Bou Denib), adult males, leg. E. Linsenmair, 26.vi.1967 .

Diagnosis: A large Trapelus species with the typical ear opening found in the genus: sunken tympanum and spiny scales above the ear opening. Trapelus boehmei sp. nov. has a relatively long head tapering abruptly at the nose, giving it a stout appearance. Body scalation is a matrix of small, feebly keeled and homogeneous scales intermixed with larger keeled scales, which usually differ in coloration from the matrix scales in breeding coloration of adult males; sometimes vertebral matrix scales are larger than the lateral matrix scales and nearly as large as the intermixed scales. Males have bluish coloration on the throat and body when in nuptial coloration, and a small gular pouch. The new species differs from all described African taxa in Trapelus by its unique coloration, body proportions, and scale morphology.

Trapelus boehmei sp. nov. differs from currently valid African species of Trapelus (some selected variable characters comparing the new species with T. mutabilis s.l. are given in Table 8):

– Trapelus m. mutabilis (type locality: Egypt) in possessing higher scale counts, larger size, and a relatively shorter tail (average of ratio TL/SVL 1.39 in T. mutabilis instead of 1.19 in T. boehmei sp. nov.; for details see Table 7), and DNA sequences.

– Trapelus m. pallidus (type locality: southern Egypt) in having homogeneous scalation on the upper hindlimb and base of the tail, and in having a greater proportion of enlarged body scales and all dorsal scales keeled instead of only the enlarged ones, and DNA sequences.

– Trapelus m. poppeki ssp. nov. (type locality: Libya: east of Tarbu ) in not possessing enlarged vertebral scales, a lower count of scale rows around midbody and a smaller size .

– Trapelus schmitzi (type locality: Ennedi Mts., Chad) in not possessing dorsal scales equal in size, with only few intermixed larger scales.

– Trapelus savignii (type locality: Egypt) in having smooth ventral scales and a small, instead of large, gular pouch, and DNA sequences.

– Trapelus tournevillei (type locality: Ouargla, Algeria) in having a shorter head, a small gular pouch, different coloration (e.g. no longitudinal lines on the belly) and smooth ventral scales, instead of keeled ventral scales.

Trapelus boehmei sp. nov. differs from the following synonyms ( A. aspera is a probably valid taxon but clearly different to the herein-described new species):

– Agama inermis Reuss, 1833 (type locality: southern Egypt) in possessing a lower proportion of slightly enlarged scales, only one row of precloacal scales, and a nonvisible tympanum.

– Agama gularis Reuss, 1833 (type locality: southern Egypt) in possessing heterogeneous scales and a dorsal crest.

– Agama latastii Boulenger, 1885 (type locality: Egypt) in lacking equal sized, rhomboidal dorsal scales.

– Agama aspera F. Werner, 1893 (type locality: Algerian Sahara, between Kef-el-Dhor and Chegga; Biskra-Bordj-Saada; Zab-el-Zig south of El Meranyer) in possessing no spiny scales and a heterogeneous dorsal scalation.

Description of the holotype: Habitus: stout; tail only moderately longer than the body; limbs and digits relatively long. Measurements. Snout-vent length 107.1 mm; tail length 135.9 mm; head length 27.7 mm; head height 15.4 mm; head width 24.7 mm; length of left forelimb 49.3 mm; length of left hindlimb 70.6 mm. Scalation. Nostril sits on the canthus rostralis, piercing the posterior portion of a large, flat nasal scale; nostril is directed obliquely upwards. Head scales are heterogeneous in size and shape: supraocular scales smooth; parietal scale small, more or less rectangular, surrounded by three enlarged scales and four scales of equal size; pineal organ visible, piercing the middle of the scale. Posterior to the parietal scale: scales originating from both sides of the parietal midline have imbrications directed laterally, with their free anterior margins having sensory pits. The eyelids have mucronate scales that form a ring. The ear opening is of medium size, about half of the size of the eye, having a superior margin with five spiny, mucronate scales on both sides; the tympanum is sunken but visible. A rudimentary nuchal crest is present, consisting of six spiny, mucronate scales. Gular scales are flat, smooth, with the *In two rows.

†In two to three rows.

Min, minimum; Max, maximum.

x = average (value in bold).

posterior margins being enlarged and slightly imbricate; gular scales are equal in size but becoming smaller on the gular fold. The gular pouch is small. Ventral scales are smooth, slightly imbricate, and equal in size. Dorsal scales are heterogeneous (larger scales interspersed amongst a matrix of smaller scales), being smooth to keeled, slightly imbricate, and partly mucronate, with the base of the scales being thickened; the intermixed larger scales are two to three times larger than the other scales, keeled, mucronate, and strongly raised. Scales on tail strongly keeled, mucronate, and not arranged in whorls. The tail is cylindrical and relatively short – only 27% longer than the snout-vent length. There are two rows of 18 (ten anterior, eight posterior) precloacal pores. Forelimbs have strongly keeled scales on the upperside, becoming smooth to the underside, homogeneous in size; digits are long with long claws, fourth digit longest, digital length increasing 4-3-2-5-1, plantar scales and subdigital lamellae are strongly keeled. Hindlimbs have strongly keeled and slightly mucronate scales on the upperside, becoming smooth and nonmucronate to the underside, scales on the upperside are homogeneous in size, becoming smaller on the underside; digits are very long with long claws, fourth digit longest, approximately 25% longer than the third digit, digital length increasing 4-3-2-5-1, plantar scales and subdigital lamellae strongly keeled. The hindlimbs are relatively long, reaching the ear with the tip of the longest digit. Coloration in alcohol. Body is pale bluish-grey above, larger scales dirty-white, giving the impression of speckled coloration. Tail dirty-white but the concretive annulations of dark bands are visible. Lateral scales and margins of belly are bluish, undersides of the limbs, tail, chest and centre of the belly dirtywhite. Throat bluish, with slightly striated margins.

Coloration in life ( Fig. 10 ): Nuptial (seasonal) coloration of males resembles the nuptial coloration of T. flavimaculatus from Arabia. Males get a more or less complete blue and additionally white speckled body and a uniform blue throat while head and belly remain in normal coloration; the tail becomes orange (see Schleich et al., 1996: plate 21, fig. 61). Pregnant and normal coloration in females is unknown.

Variations: The adult male paratypes are more or less concordant with the characters outlined for the holotype, but show more distinctly barred tails and striped throats. Some voucher specimens show enlarged vertebral matrix scales, more or less as large as the intermixed scales. Other material. One small male ( SVL 84.3 mm) from Algeria ( ZFMK 49664 ) mostly shows the coloration described above, but only with a striped throat instead of the uniform blue one of the type series and lacking the uniform brilliant blue coloration with white dots. This coloration seems to represent the nonbreeding coloration .

Relations: We compared again the molecular uncorrected 16S pairwise sequence divergences between the North African taxa of Trapelus and the newly described taxon: T. boehmei sp. nov. – T. m. mutabilis = 3.23–3.37%; T. m. mutabilis – T. m. pallidus = 1.41%; T. boehmei sp. nov. – T. m. pallidus = 2.63–2.76%; T. boehmei sp. nov. – T. boehmei sp. nov. = 0.22%; T. boehmei sp. nov. – T. savignii = 6.65–7.01%; T. m. mutabilis – T. savignii = 5.65%; T. m. pallidus – T. savignii = 6.45% (see Table 5). Here again the molecular analyses clearly support the morphological results as they corroborate the full distinctiveness of the new species from the other Trapelus species ( Fig. 4 ).

Habitat: Detailed habitat preferences of this species remain unclear. The image shown in Schleich et al. (1996) refers to a stony desert or semi-desert with little vegetation. Additionally, the image by Geniez et al. (2004) shows a bare stony soil, lacking vegetation. Trapelus mutabilis favours open, dry country with precipitation under 250 mm ( Schleich et al., 1996), for example coastal plains, semi-desert, erosion terraces, and steppes up to 1500 m. In such habitats, individuals are known to stay close to shelter, such as rocks or small bushes. Similar habitat preferences of T. boehmei sp. nov. can be assumed.

Etymology: This new species is dedicated to Prof. Dr Wolfgang Böhme, former Curator of Herpetology and former Deputy Director at the Zoologisches Forschungsmuseum A. Koenig, for his considerable and invaluable contributions to African herpetology.

Distribution: Trapelus boehmei sp. nov. probably has a broad distribution in north-western Africa. The holotype and some paratypes were collected between Akka and Icht in Morocco, and the images presented by Schleich et al. (1996: no. 61, pl. 21) and Geniez et al. (2004; no. 94) show specimens from Erfoud and Oued Mird near Ikhf-n-Ouaroun, respectively. Other Moroccan vouchers are known from 40 km north of Zagora (ZSM 52277), 10 km south-east of Goulmima (ZSM 207/1993), 25 km west of Tissint (ZSM 225/ 1993) and Ksar es Souk (ZSM 688/1979/1-5). An additional voucher in the ZFMK collection is from Colomb Béchar ( Algeria) and can be confidently assigned to T. boehmei sp. nov. (ZFMK 49664). One of the paratypes was collected 220 km south of Nouadhibou on the main road to Nouakchott in northern Mauritania. All these specimens indicate that the new species has a distribution from Algeria through Morocco to Mauritania ( Fig. 11 ). Therefore, it is most probable that T. boehmei sp. nov. is a widespread species in northwestern Africa.

Figures

Figure 4. Cladogram of the tree recovered by analysis based on 512 bp of the 16S mtDNA gene. Values above nodes are neighbour-joining bootstrap replicates (20 000 replicates; values below 50% not shown); values below nodes are Bayesian posterior probabilities (values below 0.80 not shown). Red dots (grey in print) and values on the nodes indicate the estimated age of the clade.

Figure 10. Live male of Trapelus boehmei sp. nov. of the ZFMK-type series from the type locality.

Figure 11. Distribution of Trapelus boehmei sp. nov. Morocco: 1 = between Akka and Icht (ZFMK 49751–754); 2 = Oued Mird near Ikhf-n-Ouaroun (Geniez et al. 2004); 3 = Erfoud (Schleich et al., 1996); 6 = 10 km south-east of Goulmima (ZSM 207/1993), and Ksar es Souk (ZSM 688/1979/1-5); 7 = 40 km north of Zagora (ZMB 52277); 8 = 25 km west of Tissint (ZSM 225/1993). Algeria: 4 = Colomb Béchar (ZFMK 49664). Mauritania: 5 = 220 km south of Nouadhibou (MNHN 2006.0343).