Platysaurus attenboroughi, Whiting, Martin J., Branch, William R., Pepper, Mitzy & Keogh, J. Scott, 2015

Whiting, Martin J., Branch, William R., Pepper, Mitzy & Keogh, J. Scott, 2015, A new species of spectacularly coloured flat lizard Platysaurus (Squamata: Cordylidae: Platysaurinae) from southern Africa, Zootaxa 3986 (2), pp. 173-192 : 185-190

Platysaurus attenboroughi sp. nov.

English: Attenborough’s Flat Lizard

Afrikaans: Attenborough se Platakkedis

Synonymy. Platysaurus capensis (part). FitzSimons, 1935: 535; 1943: 473; Loveridge, 1944: 97; Rose, 1950: 155; 1962: 156; Broadley, 1978: 157; Branch, 1998: 165; Van Wyk & Mouton, 1996: 117; Whiting, 2014: 214.

Type material. Holotype. TM 85806 (MJW 936), adult male collected by Bryan Maritz, Nick Tye and Chris Barichievy, May 2008 ( Table 2 , Figure 4–5 ). Type locality: Fish River Canyon, Karasburg District, southern Namibia (27˚52' 21.7S, 17˚31' 15.7E; quarter-degree unit, 2717Dc). Allotype. TM 85807 (MJW 937), adult female collected by Bryan Maritz, Nick Tye and Chris Barichievy, May 2008; same locality data as holotype. Paratypes (2 specimens): TM 85805 (MJW 935), adult male, TM 85804 (MJW 934), adult female, both collected by Bryan Maritz, Nick Tye and Chris Barichievy, May 2008, Fish River Canyon, Karasburg District, southern Namibia (27˚50' 03.7S, 17˚32' 47.5E; quarter-degree unit, 2717Dc) ( Table 2 ).

Diagnosis. A medium-sized Platysaurus (Table 3, Figure 4 ) distinguished from all congeners, except P. capensis and P. broadleyi , in that the scales on the side of the neck are indistinguishable from those on the dorsum. It can be distinguished from P. broadleyi as follows: the breeding male has blue forelimbs (anterior surface) while that of P. broadleyi may be orange, yellow, or a combination; the male has a light blue throat ( Fig. 4 , 5 ) compared to the dark blue of P. broadleyi (although this may be highly variable and could be related to male fighting ability (Whiting et al., 2006); and the male also has an extensive blue belly with a small orange lower abdominal patch and sometimes with an irregular black abdominal patch (centre), while P. broadleyi has a darker (deep blue-black) abdomen with the lower abdomen usually orange (but may also be yellow or a mix). It also differs from P.

broadleyi in some features of scalation, particularly the number of collar scales (mean P. attenboroughi sp. nov. 7.93 ± 1.06, P. broadleyi 9.22 ± 1.04), transverse dorsal scale rows (mean P. attenboroughi sp. nov. 85.23 ± 4.32, P. broadleyi 104.96 ± 5.38) and the smaller upper forelimb scales (mean P. attenboroughi sp. nov. 17.17 ± 2.09, P. broadleyi 25.82 ± 2.31; Table 3). Adult male coloration in P. attenboroughi sp. nov. is further distinguished from P. capensis by having the dorsum more extensively covered with white spots ( Figure 4 ), and with a reduced fine vertebral stripe that only partially extends on to the hindbody, and with reduced (or absent) broad, dark paravertebral stripes; adult female coloration is more vaguely patterned than in either P. capensis or P. broadleyi , lacking the bold dark and pale paravertebral stripes and with scattered pale spots. It also differs from P. capensis in features of scalation (summarized in Table 3), particularly the greater number of upper forelimb scale rows (mean P. attenboroughi sp. nov. 17.17 ± 2.09, P. capensis 14.62 ± 1.04) and number of subdigital lammelae beneath the 4th toe (mean P. attenboroughi sp. nov. 19.10 ± 1.11, P. capensis 17.92 ± 0.64).

Description. Holotype. TM 85806, an adult male with original tail (lacking 2–3 mm from tip, removed for DNA analysis). Head strongly depressed, much longer than broad (head length: tip of snout to anterior border of ear-opening, HL): 16.6 mm; head width (HW): 12.9 mm; HL/HW: 1.27). Large supranasals in broad contact behind rostral; nostril directed slightly backwards and piercing a very small nasal that contacts the rostral, first supralabial, a small postnasal and the large supranasal. Frontonasal hexagonal, as broad as long and in good contact on sides with loreal. Prefrontals in median contact. Frontal longer (3.32 mm) than broad (max. width: 2.2 mm), much wider in front than behind. A pair of frontoparietals, each in contact laterally with middle pair of supraoculars. Interparietal small, diamond-shaped, set in the middle of the two pairs of parietals, the anterior pair the smallest, and with a conspicuous pineal pore. Occipital absent, two slightly enlarged triangular granules in its position. Four supraoculars, the first large and triangular and contacting the pentagonal preocular, the last the smallest. Four supraciliaries, anterior elongate, middle pair largest. Lower eyelid with a semi-transparent disc faintly divided into a number of vertical septa. Three elongate upper temporals bordering parietals on each side, middle one largest and longitudinally elongate, twice the length of posterior one. Two additional rows of enlarged scales present in dorsal temporal region, upper row (five right, four left) vertically elongate and more than twice the size of lower row. Ventral temporal region covered with five rows of irregular granules, slightly larger than those along backbone. A small postnasal; an elongate loreal and a preocular, the former much smaller than the latter. Four suboculars, the second elongate and extensively bordering the lip below. Rostral pentagonal, broader than deep. Six supralabials, five anterior to subocular. Mental subpentagonal. Seven infralabials, 5–7 very small and elongate and sandwiched between lip and very large fourth sublabial; five, all slightly larger than infralabials, the fourth by far the largest, the fifth the smallest. A longitudinal median series of four enlarged quadrangular or polygonal gular scales, bordered by scales that become smaller (subgranular and rounded) particularly laterally on the neck, and increase in size towards the collar, which consists of eight enlarged scales; in 25 rows between the angle of the jaws. Dorsal scales smooth, small, flat and rounded, without minute granules at junctions, and largest along backbone and on flanks, smallest dorso-laterally and minute on sides, in 89–91 transverse rows across midbody. Ventrals square or a little broader than long medially, arranged in 39 mainly regular transverse and 20 longitudinal series. Eight preanal plates, median pair largest and much larger than ventrals, reducing laterally towards the groin. Limbs long and slender, length of tibia subequal to head length. Upper forelimb and thighs with subequal granules above. Forearm and tibia with enlarged keeled scales; a row of nine large transverse plates on underside of tibia, largest at midcalf. Toes long, clawed and with smooth subdigital lamellae (17 on 4th toe on both sides). A series of 16 femoral pores on lower surface of right thigh, 13 on left, with 2–3 rows of modified generation gland scales (32 right thigh, 30 left) anterior to these pores. Tail depressed, tapering, with regular whorls of elongate, quadrangular scales, strongly keeled above laterally, smooth below. Measurements for the holotype and other type material are given in Table 2 .

Description. Allotype. TM 85807, an adult female with original tail (lacking 2–3 mm from tip, removed for DNA analysis). Scalation as for holotype except for: head length 16.4 mm, head width 11.4 mm; HL/HW: 1.44); large supranasals in only narrow contact behind rostral; a small triangular occipital present at middle border of posterior parietals. Middle row of temporals comprised of six vertically elongate scales on each side of head; gulars in 26 rows between the angle of the jaws, and seven scales in collar; dorsal scales in 90 transverse rows across midbody; ventrals arranged in 38 mainly regular transverse and 18 longitudinal series; femoral pores minute, 17 on each thigh; thighs without modified scales with generation glands; subdigital lamellae under 4th toe 18/19.

Coloration. Holotype: In life: Above dull grey-brown dorsally, with a single narrow pale stripe along the backbone, running from the base of tail to crown of head, where it shallowly divides; scattered pale spots, 3–5 granules across, speckled body and flanks, with largest aligned in a longitudinal row on the upper flank which is confluent with a pair of pale lateral stripes on head that arise above each eye and travel backwards across upper temporal region; forebody flanks and front of forelimbsh bright blue, which extends through neck and onto upper lip; upper surface of tail and flanks anterior to groin orange-brown; proximal tail and upper surface of hindlimbs light brown at tail tip; upper hindbody and upper surface of tail grey-tan, darker along midline, with vague, paler spots laterally. Ventral surface of neck and first three-quarters of belly brilliant blue, paler under head, and with black ‘badge’ with irregular outline in centre of belly; lower surface of forelimbs ivory-pink; rear of belly, underside of tail and lower surface of hindlimbs light orange. In preservative (70% ethanol): faded, most orange and blue coloration lost; conspicuous central black belly ‘badge’ inconspicuous; thin dorsal stripe and pale spots only vaguely visible; upper and lower surface of head back to forelimb insertion dark grey, lighter towards hindlimbs; tail and lower surface of hindlimbs pinkish-ivory, darker on upper surface of limbs.

Museum Number Subdigital lamellae (4th toe) 17/17 18/19 18/16 19/18

* tail original, but 2–3mm removed for DNA analysis

** tail tip regenerated and 2-3mm removed for DNA analysis

Allotype: In life: above grey brown, with a thin vertebral stripe from top of head to tail base, bordered on each side from eye to groin by a pale dorso-lateral ‘stripe’ comprising a paler background underlying a longitudinal series of pale spots; back, flanks and upper surface of hindlimbs speckled with pale spots, 2–4 granules wide; top of head dark brown lighter with three pale stripes; upper lip and lower temporal region tan-cream; tail dirty gold, darker along midline, paler below; belly pale brown, lacking a central black ‘badge’. In preservative (70% ethanol): as above but much paler above and below; vertebral stripe almost inconspicuous and pale spots less visible.

Distribution. Along the lower Orange River from Goodhouse to the Richtersveld, extending north into Namibia and recorded from the Hunsberg, Huamsib and Ploegberg mountains and the Fish River Canyon (Figure 1).

Habitat and climate. Platysaurus attenboroughi sp. nov. occurs in the arid-subtropical region of the Northern Cape Province of South Africa and southern Namibia and specifically within the Gariep Desert Bioregion ( Mucina & Rutherford, 2006). This is an arid area characterized by low and erratic summer rainfall. Summers are typically hot and dry. Like all flat lizards, they are dependent on rock (mostly granite in this area) and take refuge in narrow rock fissures where they can escape suboptimal temperatures and predators. These areas are largely devoid of significant vegetation bar the occasional fig tree ( Ficus ) or succulent. For more detailed descriptions of climate, vegetation and topography see Mucina & Rutherford (2006).

TABLE 3. Summary statistics (n = sample size, mean ± 1 SD, range) for SVL (snout-vent-length) and meristic characters: gulars: scales transversely between posterior sublabials, UL: upper labials, Col: scales in collar, LRV: longitudinal rows of ventrals, TRV: transverse rows of ventrals, TD: transverse rows of dorsals, Forearm: upper forelimb scale rows, 4th toe: number of subdigital lamellae beneath 4th toe, and FP: number of femoral pores on left thigh.

Character P. attenboroughi sp. nov. P. broadleyi P. capensis

N 87 45 13

SVL 71.75 ± 6.92 70.18 ± 7.21 68.92 ± 11.00

36-86 40-80 37–82

Gulars 25.5 ± 0.3 27.5 ± 0.4 23.8 ± 1.4

22–30 24–32 21–27

UL 4.95 ± 0.24 4.9 ± 0.27 5 ± 0.20 4–5.5 4–5 4.5–5.5

Col 7.93 ± 1.06 9.22 ± 1.04 7.69 ± 0.63

5–10 7–11 6–8

LRV 19.79 ± 1.08 20.73 ± 1.0 19.69 ± 0.63

18–22 20–22 18–20

TRV 41.84 ± 2.43 43.32 ± 1.86 40.42 ± 1.08

36–48 41–47 38–42

TD 85.23 ± 4.32 104.96 ± 5.38 83.67 ± 3.45

78–94 92–117 77–86

Forearm 17.17 ± 2.09 25.82 ± 2.31 14.62 ± 1.04

13–23 21–30 13–16

4th toe 19.10 ± 1.11 20.6 ± 1.03 17.92 ± 0.64

17–22 18–22 17–19

FP 16.73 ± 1.33 16.68 ± 0.83 16.73 ± 1.33

14–19 15–18 14–18.5 Natural history and behaviour. In South Africa, it is mainly restricted to the Richtersveld region ( Bauer & Branch, 2001; Whiting, 2014). There it is widespread and common in boulder-strewn areas and on broad rock faces, often far from river courses (e.g. Tierhoek). It does not form the high density populations recorded for P. broadleyi (MJW unpubl. data). All Platysaurus have a fixed clutch of two eggs ( Broadley, 1978). The reproductive cycles of P. capensis , P. broadleyi and P. attenboroughi sp. nov. were collectively studied when these were considered a single species ( Van Wyk & Mouton, 1996). The minimum size at sexual maturity is 64 mm (sex not specified); eggs are likely laid in summer (November–December) ( Van Wyk & Mouton, 1996). While we know very little about the diet of P. attenboroughi sp. nov., the closely related P. broadleyi is an omnivore and lives in similar habitat. The marked sexual dichromatism suggests a classic sexual selection system in which males compete heavily for females, as is the case in P. broadleyi ( Whiting et al., 2003; Whiting et al., 2006). Males do have UV-reflective throats, which suggests a role of this colour signal in either settling contests (as in P. broadleyi ) or in mate choice, although this remains to be tested. In the two males we measured, their throats had violet and blue and less pure UV than we would typically see in adult male P. broadleyi ( Figure 5 )(Whiting et al., 2006). Measurement of spectral reflectance in additional individuals is necessary for a proper comparison.

Etymology. We name this new species in honour of Sir David F. Attenborough (Fig. 3), in recognition of his immense contribution to the public understanding and appreciation of animals, plants, ecosystems and nature in general. David Attenborough made flat lizards, specifically the closely related Platysaurus broadleyi , famous in the BBC documentary series Life in Cold Blood.

Conservation status. The conservation status of the Cape Flat Lizard (P. c a pe n s i s sensu lato) was assessed as of Least Concern (Whiting 2014). Our division of this taxon into two species therefore requires a re-assessment of their conservation status. Both have relatively restricted distributions, with that of P. c a pe n s i s now restricted to only seven quarter-degree grid squares (Figure 1). Although the Kamiesberg region is not formally protected, it does form part of an envisaged expansion of the Namaqua National Park (San Parks 2013). The species has also been recorded from Carolusberg, Springbok, within or in close proximity to the Goegap Nature Reserve. Platysaurus attenboroughi is recorded from numerous localities within the Richtersveld National Park, and the Fish River Canyon forms part of the Ai-Ais Hot Springs Game Park in Namibia. Together these two conserved areas form the /Ai/Ais-Richtersveld Transfrontier Park, and thus protect a large proportion of the species’ range. The extensive granite outcrops of both the Kamiesberg region and the /Ai/Ais-Richtersveld Transfrontier Park form the main habitat for both species, and are subject to little existing habitat threat. We conclude that currently both P capensis and P. attenboroughi are not of conservation concern (Least Concern).

In summary, we describe a new species of flat lizard ( Platysaurus attenboroughi ) that was formerly confused with P. capensis . Platysaurus attenboroughi sp. nov. occurs on rock from the region of the Orange River west of Goodhouse, including the Richtersveld of the Northern Cape Province of South Africa and into Namibia as far as the Fish River Canyon. This poorly known species warrants detailed future study.

Figures

FIGURE 4. a.) Platysaurus attenboroughi sp. nov., male (paratype, TM 85805), Fish River Canyon; b.) P. attenboroughi sp. nov., female (allotype, TM 85807), Fish River Canyon; c.) P. attenboroughi sp. nov., male (holotype, TM 85806), venter, Fish River Canyon; d – e.) P. attenboroughi sp. nov., male (holotype, TM 85806), Fish River Canyon; f.) P. attenboroughi sp. nov., male, Richtersveld; g.) Platysaurus capensis, male, Kamiesberg near Kamieskroon; h.) P. capensis, female, Kamiesberg near Kamieskroon; and i.) P. attenboroughi sp. nov., habitat, Fish River Canyon (Photograph by N. Tye). All other photos by M. J. Whiting.

FIGURE 5. a.) Spectral reflectance curves objectively measuring colour by body region in the holotype (TM 85806) in life. b.) Spectral reflectance of throat region for two males: (MJW 935 / TM 85805, MJW 936 / TM 85806 = holotype) and two females (MJW 934 / TM 85804, MJW 937 / TM 85807). Males have more distinct UV in their throats than females although it is still less conspicuous than what has been documented for the closely related P. broadleyi (Whiting et al., 2006).

Tables

TABLE 2. Measurements (mm) for the type series of Platysaurus attenboroughi sp. nov.

	Holotype TM 85806	Allotype TM 85807	Paratype TM 85805	Paratype TM 85804
Sex Snout-vent length Tail length	M 86.6 121 (t) *	F 78.5 124 (t) *	M 77.5 114 (r) **	F 74.0 107 (r) **
Total length Head length Head width	207.6 (t) * 16.6 12.2	202.5 (t) * 16.4 11.4	191.5 (t) ** 16.8 11.9	181.0 (t) ** 16.5 11.2
Head height Snout length Eye diameter	6.0 6.7 4.8	5.4 6.3 4.4	6.1 6.4 4.5	5.7 6.6 4.8
Inter-limb length Elbow to wrist Knee to heel	36.4 12.9 16.0	40.1 11.3 15.0	35.0 12.5 16.2	35.8 10.2 16.3
Gulars Ventrals (long.) Ventrals (trans.)	25 39 20	26 38 18	27 39 20	26 37 18
Femoral pores Dorsals (trans.) Collar scales	16/13 89-91 8	17/17 90 7	17/18 88-91 8	16/16 89=90 10