Phrynocephalus lutensis, Kamali, Kamran & Anderson, Steven C., 2015

Kamali, Kamran & Anderson, Steven C., 2015, A New Iranian Phrynocephalus (Reptilia: Squamata: Agamidae) from the hottest place on earth and a key to the genus Phrynocephalus in southwestern Asia and Arabia, Zootaxa 3904 (2), pp. 249-260 : 250-256

Phrynocephalus lutensis sp. nov.

Lut Desert Toad Headed Agama (Figs. 3–4)

(There are no synonyms, as the species has not been seen previously or at least never recorded in the literature).

Holotype. Adult male, ZISP 28014 (Zoological Institute, St. Petersburg). Rig-e Yalan, Dasht-e Lut (Mega Dune, Lut Desert), Kerman Province, near the junction of Kerman, South Khorasan and Sistan & Baluchistan Provinces. 30° 27’ N, 59° 21’ E. 963m. Collected by Mehrdad Ghazvinian 11 March 2014.

Paratypes. Adult male, ZISP 28015; Adult female, ZISP 28016, Same data as holotype.

Diagnosis. No upraised swollen scales on dorsum; no cutaneous fold at angle of mouth; no fringe of scales on posterior border of thigh and sides of base of tail; side of head and neck without projecting fringe-like scales; prominent fold on lateral sides of body, separating dorsal and ventral scales; dorsal scales subequal, smooth in general appearance, with upturned mucros when viewed from the side under magnification, homogeneous, not keeled; no enlarged scales along flanks; nasals not in contact and separated by 1–3 small scales; tail 106–119 percent of snout-vent length; no spinose scales on neck or back of head; both sides of third and fourth toes and one side of second toe strongly fringed; suborbital scales small and equal in size and never elongate; nostril not visible in entirety when head viewed from side; supralabial scales separated from eye by 3–4 rows of scales; about 30 lamellae under fourth toe; width of space between nostrils less than one half of distance between nostril and preocular ridge; about 30 scales from eye to eye across head. Snout-vent length (mm) 65–75, tail 69–89 (tail = 106–119% body length).

Comparisons. It can be distinguished from the morphologically similar species, Phrynocephalus luteoguttatus Boulenger 1887 by having homogeneous dorsal scales, by having 1–3 scales between nasals, by lacking long, flat, upturned fringe-like scales on sides of back of head and neck, by having laterally upturned (about 90 degree upward) and mucronate dorsal scales, by smaller number of scales between eyes (25 versus 30), by higher percentage of tail length to snout-vent length (106–119 versus 85–102), and distinctly longer body size (longest SVL of 75 mm and longest tail length of 89 mm versus longest SVL of 46 mm and longest tail length of 43 mm respectively). From the morphologically similar species, P. euptilopus , which has scattered enlarged flat dorsal scales amongst its uniform small scales, it differs in that the latter species has red, orange, and black dorsal spots, top of head green in life; nasal shields usually in contact, and spinose scales on head and neck ( Smith 1935; Minton 1966). It can be distinguished from P. cl ar k o r u m Anderson & Leviton 1967 by lacking single very elongate suborbital scale, by having 1–3 small scales between nasals, by having strongly fringed 3rd toe, by lower percentage of tail length to SVL (106–119 versus 124–153), by lacking the dorsolateral straight-edged black margined white stripe from behind eye down body onto tail, and by lacking the series of elongate oval brown spots down vertebral line, linked by light chain. It can be distinguished from P. helioscopus ( Pallas 1771) by having homogeneous and not keeled dorsal scales, by having large and strongly fringed scales on both sides of third and fourth toes and one side of second toe, and by lacking two kidney-shaped prescapular pink or red patches margined by light blue. It can be distinguished from P. interscapularis Lichtenstein 1856 by lacking a row of enlarged tubercular scales on posterior margin of thigh and sides of tail base that form short fringes, by higher number of scale rows between supralabials and eye (3 versus 2), and by distinct body size (longest SVL of 75 mm and longest tail length of 89 mm versus longest SVL of 38.2 mm and longest tail length of 42.5 mm, respectively) and the higher percentage of tail length to snout-vent length (106–119 percent versus 97–111 percent). From P. sogdianus Chernov 1948 it is distinguished by having the position of nostrils well above an imaginary line joining anterior corners of eyes when viewed from in front, the greater number of internasals, usually 4 rows (as opposed to 2 or 3) of horizontal scales above supralabials counted below the anterior part of eye, no enlarged, diagonally keeled and elongated scale on anterior temporal region. It can be distinguished from P. maculatus Anderson 1872 and P. arabicus Anderson 1894 by its distinctive color pattern, black third or more of distal end of tail, by its more vertical snout, by lacking considerably larger scales on vertebral region, by presence of strongly fringed scales on both sides of third and fourth toes and one side of second toe and by lower percentage of tail length to SVL (106–119 versus 140–158). It can be distinguished from P. mystaceus (Pallas 1776) by lacking large, fringed, cutaneous fold at angle of mouth. It can be distinguished from P. ornatus Boulenger 1887 by lacking large dorsal scales, by presence of 1–3 scales between nasals, by having more suborbital scales (5 versus 2–3), by lower percentage of tail length to SVL (106–119 versus 119–132), and by lacking two rows of light orange or pinkish spots, dark gray, festooned dorsolateral stripe, and by having tail dark-colored at its distal area instead of having four or five dark black or brown bars. It can be distinguished from P. persicus De Filippi 1863 by lower number of scales between nasals (1- 3 versus 3-5), by small and homogeneous dorsal scales, never tubercular or nail-like, by having strongly fringed scales on both sides of third and fourth toes and one side of second toe, by having nostrils not visible in their entirety when head viewed from side, and by lacking two kidney-shaped prescapular pink or red patches margined by light blue. It can be distinguished from P. raddei Boettger 1890 by higher number of subdigital lamellae under fourth toe (30 versus 21–24), by lacking clusters of dorsal scales with upraised posterior margins and prominently keeled scales on midline of back and by lacking half-moon shaped red patch on each scapular region. It is distinguishable from P. scutellatus ( Olivier 1807) by having homogeneous dorsal scales, by having 1–3 scales between nasals, by lower percentage of tail length to SVL (106–119 versus 118–157) and by lacking intense dark (black) crossbars on tail. See Arnold (1999) for additional characters of differentiation among species of Phrynocephalus .

Description of holotype. Overall appearance: A mature male Phrynocephalus of moderate size; outline of body viewed from above robust and rounded; Scales of back of more or less uniform size, juxtaposed, not keeled; dorsal scales of thigh weakly keeled, more so proximally and laterally, juxtaposed, larger toward knee, with metallic yellow sheen; in lateral view it has a bluish green sheen; top of head dark reddish orange, sides of head lighter orange (at least at the time of year collected); dorsum of body and tail yellowish gray with lighter, bluish white dots, loosely arranged in longitudinal and lateral rows; tail and body with darker gray, indistinct, interrupted crossbars, most prominent on sides; dorsal anterior half of tail brighter yellow, distal half of tail black; shank with slightly imbricate scales, more so distally and laterally, with metallic yellow sheen; slight fringe of scales on lower sides of body and thigh; third and fourth toes strongly fringed, dorsal scales of tail strongly keeled. Ventral surfaces of body, legs and tail pale aquamarine blue, this color extending onto throat nearly to the jawline; ventral half of tail black; ventral scales of tail and body keeled, those of tail strongly so, subimbricate. Snout-vent length 75 mm. Head: Scales of head swollen, supralabials 15 (on right side of head); sublabials 15 (on right side of head); 3–4 rows of scales between lower edge of orbit and supralabials; 12 long pointed scales at upper edge of lower eyelids forming fringe, those of upper eyelids not forming projecting fringe; 7 enlarged supraorbital scales, scales covering tympanic area relatively small, grading into larger scales of temporal region, where the yellowish scales of the lower lateral head meet the darker orange-red scales of the upper lateral and top of head; preocular scales relatively small ( Fig. 4 i). Nasal scales separated by one upper scale, 2 central and 3 lower scales, nostrils covered by small scales forming valves ( Fig. 4 m); snout vertical, nasals cannot be seen from dorsal aspect. Top of head covered with swollen scales of various sizes, but larger than those of limbs, tail, and body; translucent parietal scale encircled with a ring of subcircular scales; dark orange-red color begins at back of skull ahead of fold on neck and extends forward onto snout 2 rows below nasals. Largest scales run from occipital along crown of head to nasals, scales on either side between these large scales and supraorbitals smaller, subequal; lower eyelid with 11 prominent denticles ( Fig. 4 i); about 25 scales across top of head between supraorbitals at narrowest point. Ventral head and throat covered with smooth subimbricate scales, smaller than ventral body scales; yellowish scales extend onto edges of lower jaw; aquamarine blue scales cover remainder, darkest along midline; Prominent fold across throat,covered FIGURE 3. Body photos of Phrynocephalus lutensis sp. nov. All photos by K. Kamali. a. Phrynocephalus lutensis sp. nov. ♂ holotype; b. Phrynocephalus lutensis sp. nov. ♂ holotype. Ventral view; c. Phrynocephalus lutensis sp. nov. ♀ paratype. Dorsal view; d. Phrynocephalus lutensis sp. nov. ♀ paratype. Ventral view; e. Phrynocephalus lutensis sp. nov. ♂ holotype. Dorsal trunk scales, anterior to the right; f. Phrynocephalus lutensis sp. nov. ♂ holotype. Dorsal view of hind limb and proximal tail. g. Phrynocephalus lutensis sp. nov. ♀ holotype. Ventral view of right hind foot and toes. h. Phrynocephalus lutensis sp. nov. ♂ holotype. Dorsal trunk scales under magnification, viewed from the side, anterior to the right.

j. Phynocephalus lutensis sp. nov. ♂ holotype. Top of head.

k. Phrynocephalus lutensis sp. nov. ♂ paratype. Ventral head

l. Phrynocephalus lutensis sp. nov. ♀ paratype of Ventral head

m. Phrynocephalus lutensis sp. nov. ♂ holotype. Front of head.

with very small scales, smaller than those of ventral surface of lower jaw ( Fig. 4 k, 4l); very small, granular scales on sides of throat behind level of angle of mouth. Body: A prominent fold along sides of body, separating dorsal and ventral scales; dorsal scales of body juxtaposed, approximately uniform in size, swollen in centers, not keeled, but viewed from the side under magnification, each scale with sharp mucros extending upwards from center of scale at approximately 90°, approximately 66–68 across widest part of back (no two observers likely to count exactly the same number) (Figs. 3e, 3h); a prominent fold along length of body separating dorsal scales and much smaller, sharply keeled lateral scales; venter with smooth, quadrangular scales of uniform size (Fig. 3b–3d), approximately 38 across widest part of abdomen. Tail: Proximal dorsal tail the same color as dorsal body scales, these scales roughly the same size as dorsal body scales, but more quadrangular, juxtaposed, the center of scales with upward-directed sharp points (Fig. 3f); distal half of tail black, scales subimbricate, prominently keeled, with sharper, more prominent points as are those of sides of entire length of tail; scales of dorsal and ventral tail arranged in rings of four rows of scales, presumably equivalent to each caudal vertebra, but far less distinct as compared with e.g., Laudakia ; proximal ventral tail same color as ventral body and limbs, distal part black, ventral surface of anterior part of tail dirty white with aquamarine blue sheen, scales similar in appearance to dorsal tail scales; tail length 89 mm, 119% of snout-vent length (Fig. 3b–3d). Limbs: Dorsal and ventral surfaces of limbs similar in color to those of body, but dorsal limbs with metallic sheen; scales of thigh and shank all prominently keeled, those on outer surface of thigh giving the appearance of forming a slight fringe (Fig. 3f). Second and third toes and fingers with long lateral fringes on both edges; 30 lamellae under fourth (longest) toe. (Fig. 3g).

Paratypes. Female (ZISP 28016) (Figs 3c, 3d, 3g & 4l). Similar to holotype in overall general appearance, legs and tail less intensely aquamarine blue in color, venter and ventral surface of anterior part of tail white with no aquamarine blue sheen; dorsal scales 65–66, ventral scales 37–38, supralabials 14, lower labials 15, upper head scales 27, snout-vent length 65 mm, tail length 69 mm (106% of snout-vent length).

Male (ZISP 28015). Similar in size and general overall appearance, legs and tail pale aquamarine blue, venter and ventral surface of anterior part of tail dirty white with aquamarine blue sheen, Dorsal scales 68–70, ventral scales 38–40, upper head scales 28, supralabial scales 14, lower labials 16, tail length 89 mm, snout-vent length 75 mm, ratio of tail length to snout-vent length 119%.

Habitat ( Fig. 2 ). The substratum on and in which the species is found is wind-blown sand, to which the morphology, and especially the fringed digits and nostril valves, are well adapted. The immediate habitat lies at the south-easternmost edge of the Dasht-e Lut, a semi-enclosed basin, surrounded by mountains, similar to such basins throughout southern Central Asia. The area in which this species lives is called Rig-e Yalan (Mega Dune). The elevation of the area from which the specimens were taken was about 700–1000 meters above sea level with high elevated dunes with height of about 400 meters in some parts. Within the Dasht-e Lut, wherein Mega Dune lies, is the hottest point in the world. The surface temperature in warm seasons goes up to 70.7°C (as recorded by NASA) and in cold seasons and at night it decreases to -10°C (reported by collector). At the time that specimens were taken the lowest and highest daily air temperatures were 15° and 30°C, respectively.

Thus far, nothing is known of the species’ behavior or ecology.

Figures

FIGURE 2. Habitat (type locality) of Phrynocephalus lutensis sp. nov.. Dasht-e Lut (Lut Desert), Kerman Province, Iran. Photo M. Ghazvinian.

FIGURE 4. Phrynocephalus lutensis sp. nov. Photos of head. All photos by K. Kamali i. Phynocephalus lutensis sp. nov. ♂ holotype. Lateral head.