Laudakia stellio salehi Werner, 2006

Lachman, Esther, Carmely, Hilla & Werner, Yehudah L., 2006, Subspeciation befogged by the ‘‘ Seligmann effect’ ’: the case of Laudakia stellio (Reptilia: Sauria: Agamidae) in southern Sinai, Egypt, Journal of Natural History 40 (19 - 20), pp. 1259-1284 : 1270-1278

Laudakia stellio salehi Werner n. ssp.

( Figures 5 , 6)

Lacerta stellio (part) Linnaeus 1758, p 202. Terra typica: Delos, Aegypto, Africa. Terra typica restricta ( Mertens and Mueller 1928; Almog et al. 2005): Delos Island, Cyclades. Stellio antiquorum (part) Eichwald 1831, p 187.

Uromastix horrida (part) Wagler, 1832 in Michahelles 1833, p 902.

Agama stellio (part) Boulenger 1885, p 368.

Agama stellio brachydactyla Haas 1951, p 1052 . Terra typica: Foot of Jebel Lussan, near the Israel –Sinai frontier, SSW of Beer-Sheba.

Stellio stellio (part) Moody 1980.

Agama (Stellio) stellio brachydactyla (part) Arnold 1986, p 415.

Laudakia stellio brachydactyla (part) Leviton et al., 1992, p 14.

Material examined

Holotype: HUJ-R 10732, male, ‘‘ Sinai : 3 km from Sheikh Harun towards Watiya Pass’ ’ [ Israel Grid 051/778; 28 ° 359300N, 33 ° 599E], alt. 1650 m, probably summer 1968, probably coll. Abraham Haim. Paratypes (N 562): listed in the Appendix.

Diagnosis

A subspecies of Laudakia stellio most similar to L. s. brachydactyla but differing from it in longer toes with more subdigitalia (usually.18.5 as average of left and right fourth toes); longer tail (males,.144 PERCRA; females, almost always.124 PERCRA); and in the array of enlarged dorsal scales ( Figure 5 ). While in L. s. brachydactyla the left and right transverse rows of juxtaposed enlarged scales are bridged medially by the broad continuous vertebral band of fairly similar enlarged scales, in L. s. salehi the left and right transverse rows of enlarged scales tend to be separated at the vertebral line by small scales, especially in males. In the dorsal coloration the light markings tend to be numerous narrow transverse bars, rather than four or five bold blotches in L. s. brachydactyla.

Description of the holotype

Adult male. RA, 131 mm; head length, 30.0 mm (22.9 PERCRA); head width, 30.7 mm (23.4 PERCRA); head index, 97.7; head depth, 16.4 mm (12.5 PERCRA); head flatness, 182.9; forelimb length, 69 mm (52.7 PERCRA); hindlimb length, 96 mm (73.3 PERCRA); interlimb ratio, 1.39; fourth toe length, 31 mm (23.7 PERCRA); tail, 190 mm (145 PERCRA).

Supralabialia (L/ R), 12/12; dorsalia pattern, 10; dorsalia size index, 2.47; dorsal enlarged scales juxtaposed in orderly transverse rows, 18 between axilla and groin, each row subdivided into left and right halves by a band of small scales (smaller than the ones between successive tubercle rows) running along the vertebral line. Ventralia size index, 3.62; mid-abdominal band of glandular scales moderately well defined; preanal glandular scales, 60; subdigitalia, 19/19.

Coloration (in alcohol): head uniformly grey (no markings around eyes or ears or on throat). Dorsally, body and limbs dark grey; transverse tubercle rows light cream, creating cross-banding; no dorsal light blotches. Limbs grey. Underparts cream-grey; throat somewhat darker. Tail dorsally with 16 grey cross-bands, poorly demarcated from the cream intervals; ventrally yellowish cream in anterior third, thereafter, gradually, the dorsal grey cross-bands form rings.

Variation

The mensural and meristic variation of the focal type specimens are presented in Tables V and VI. The type series (N 563) varies in the array of dorsal transverse rows of juxtaposed enlarged scales. A variable number of anterior rows tend to be replaced by scattered tubercles or tubercle clusters that may or may not be arranged in transverse rows. When the set is sufficiently complete for counting, which occurs mainly in males, there are 16–19 (usually 18–19) rows between axilla and groin.

Coloration (in alcohol, based on specimens in which coloration was reasonably well preserved, N 557) varies greatly. Dorsally, the coincidence of light cross-banding with the full set of transverse rows of enlarged scales as in the holotype occurs only in 4/ 22 paratypes. In others only every other such scale row is light-coloured (4/ 22 paratypes, females), possibly asymmetrically alternating over left and right half-rows (2/22), or the narrow light cross-bands are joined by a light vertebral band (4/22, males) that sometimes shows a few blotch-like accentuated areas (3/22); additional variants occur. Ventrally the gular area is spotless as in the holotype in 23/ 49 paratypes. Otherwise the grey throat carries darker markings of varying density and shape—tiny dots, dashes, rarely even marbling (20/ 49) or, exceptionally, lighter markings (6/49). The belly is spotless as in the holotype in only 4/ 21 paratypes, usually it is dotted with single dark scales of varying intensity and density (17/21). The limbs are unmarked, dark, in only one paratype; mostly they are crossbanded dark and light (12/21) or only the forelimbs are (9/21). The number of dark crossbands on the complete tail ranges from 12 to 17 (usually 14 or 15) but the last few may be represented by a long black tail tip.

The live coloration (from diapositives of three males and two females, from assorted localities) is more vivid ( Figure 6). In the males, the general background of granules is grey. Mid-dorsally, the light-coloured enlarged scales are orange and the remaining enlarged scales are metallic-bluish lighter grey. On the flanks, clusters of tubercles are light creamyellow. On the head, the eye and ear are framed orange to varying extent. Light orange may be dominant in the cross-banded hindlimbs and anterior part of the tail. The females may be somewhat less colourful, especially on the head, and seem to lack the bluish component of those enlarged dorsal scales that are metallic-grey (the others being orange). In one captive female photographed comparatively on a warm and on a cool day, the grey ground colour was much darker on the cool day.

stellio salehi, with separate presentation of original-tailed individuals.

mountain massif constituting the Irano-Turanian region enclave. Individual localities were at altitudes ranging form 400 to 2100 m, mostly. 1000 m (one at 50 m). Outside this massif the subspecies seems to occur sparsely ( Figure 4 ) and this scarcity of records is probably not merely due to biased collecting effort, as shown by maps for other lizard species ( Werner and Sivan 1994; Segoli et al. 2002). Bodenheimer (1937) regarded L. stellio in Sinai as ‘‘known so far from the higher elevations of the Sinai massif only’’. Saleh (1997), too, commented, ‘‘In southern Sinai, apparently restricted to higher altitude where it usually prefers vertical walls and runs into crevices when disturbed’’.

Danin (1987) characterized the magmatic massive of south Sinai: area 1300 km 2; altitude 1500–2640 m; mean annual precipitation 70–100 mm (some of this, snow on high mountains); mean annual temperature 9–15 ° C; commonest plant—the shrub Artemisia herba-alba .

In our material the two smallest juveniles were 42 mm RA (HUJ-R 11196, 5 September 1968, Sinai: Wadi Naseb Tarfa) and 44 mm RA (HUJ-R 10726, 3 September 1968, Sinai: St Catherine’s Monastery area). Presumably these hatched earlier in the same summer .

Etymology

Laudakia stellio salehi is named for Professor Mostafa A. Saleh, Department of Zoology , Al Azhar University, Cairo, Egypt , who recognized the distinction of this population and partly characterized it morphologically and ecologically ( Saleh 1997). Moreover, ‘‘saleh’’ means righteous, which seems appropriate for a lizard from Mt Sinai.

Comparison between subspecies

Saleh (1997) already noted that ‘‘Individuals from the high mountains of south Sinai … have darker coloration and longer tails and their enlarged mid-dorsal scales are only feebly keeled’’. Here we present, beyond the geographical coherence of the PCA scattergrams, the descriptive statistics of characters that significantly differed between the subspecies, within Tables V (males) and VI (females), specifying the levels of significance. In one character, tail length, there was no overlap of the range of variation between the two subspecies in the males, and almost none in the females. Although the phenomena of directional asymmetry and of sexual dimorphism differed little between the two subspecies, the assemblages of correlated characters, including asymmetry, differed between them, as reported above, accentuating the distinction of these subspecies as biological entities.

Moreover, the two subspecies differ in qualitative terms: foremost, the array of enlarged keeled dorsal scales. While in L. s. brachydactyla the left and right transverse rows of enlarged scales merge medially with a broad continuous vertebral band of fairly similar enlarged scales, in L. s. salehi these left and right rows tend to be separated by small scales that at least in most males form a continuous narrow vertebral band. The dorsal coloration, variable as described above, never displays the bold four to five yellow or orange blotches that adorn typical L. s. brachydactyla ( Werner 1995, p 55, Figure 2 ; Saleh 1997, Plate 27; Amitai and Bouskila 2001, p 131). The gular coloration, of possible social role, is in L. s. salehi usually uniform or with dark markings, while in L. s. brachydactyla it is usually with light spots (10/ 13 specimens), rarely uniform or with dark spots (2/13; 1/13). In L. s. vulgaris the throat is usually dark with lighter spots, at least anteriorly (36/42), rarely uniform (5/42) or marbled (1/42). The subspecifically distinct scale and colour patterns appear already in juveniles of 44 mm RA, possibly earlier.

Figures

Figure 2. Map of the study area (from Werner 1988) delimited towards north by a broken line and subdivided by a second broken line into northern ‘‘Negev’’ and southern ‘‘Sinai’’, with localities of the focal specimens (solid circles) and human settlements for orientation (open squares), explained in Werner (1987, 1988).

Figure 4. Variation in ‘‘Laudakia stellio brachydactyla’’ sensu lato from the Negev (Israel) and Sinai (Egypt). Bimodal distribution of one character, fourth toe length, one mode due to regenerated-tailed specimens, the other due to original-tailed specimens (N513).

Figure 5. Central-dorsal scale pattern distinguishing Laudakia stellio salehi (A) from L. s. brachydactyla (B), from photographs of the holotypes. While in L. s. brachydactyla there is a broad mid-dorsal band of large scales, in L. s. salehi the left and right transverse rows of enlarged scales are separated by a narrow mid-dorsal stripe of small scales. Scale bars: 5 mm.