Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012

Parrinha, Diogo, Marques, Mariana P., Heinicke, Matthew P., Khalid, Farkhanda, Parker, Kelly L., Tolley, Krystal A., Childers, Jackie L., Conradie, Werner, Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, A revision of Angolan species in the genus Pedioplanis Fitzinger (Squamata: Lacertidae), with the description of a new species, Zootaxa 5032 (1), pp. 1-46 : 25-28

Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012

( Figs. 6–7 )

Eremias undata undata: Laurent (1964: 60) [part]

Pedioplanis undata: Haacke (2008: 6)

Pedioplanis sp. 2 : Conradie et al. (2012b: 95)

Pedioplanis haackei: Conradie et al. (2012b: 101) ; Ceríaco et al. (2016: 37 ; 2018: 113) [part]; Marques et al. (2018: 223; 2019b: 504); Branch et al. (2019b: 92; 2019c: 296)

Pedioplanis sp. : Ceríaco et al. (2020: 401) [part]

This species was formally described by Conradie et al. (2012b) based on specimens collected between Lake Arco and Espinheira, in Namibe Province. It is possible that Bocage’s (1895) records of Eremias namaquensis from “Capangombe” and “l’intérieur de Mossamedes” included representatives of this species, but the loss of this material precludes any confident conclusion. Specimens from Moçâmedes and Munhino in Namibe Province, and Quilengues in Huíla Province, collected by António de Barros Machado in 1949, were later reported by Laurent (1964) as Eremias undata undata and are still deposited in the collections of the Museu Regional do Dundo. Additional specimens were collected in Namibe Province by Wulf Haacke in the 1970s, deposited in the collections of the Ditsong National Museum of Natural History and later reported as Pedioplanis undata ( Haacke 2008) . New material reported by Ceríaco et al. (2016) and in this paper considerably expand the known distribution area of Pedioplanis haackei . A specimen collected by Conradie et al. (2012b) near the bottom of the Leba escarpment (PEM R18460*, field number MBUR 02132) was provisionally treated by the authors as a separate lineage, Pedioplanis sp. 2 . Our analysis, however, shows that this specimen belongs to a northern clade of P. haackei , and we interpret this as an indication of intraspecific genetic structure rather than a distinct taxon.

Diagnosis: A small Pedioplanis with an average SVL of 45 mm (max 59 mm), with a tail length roughly two and a half times the SVL ( Fig. 6 ). It can be distinguished from other species of the genus in Angola and neighboring regions by the following combination of characters: (1) lower eyelid with two enlarged transparent scales, often with a row of smaller ones below; (2) five (rarely four or six) supralabials anterior to the subocular and two to three posteriorly; (3) two (rarely one or three) rows of small granules between the supraoculars and supraciliaries; (4) a group of 7–28 (> 12 in 75% of specimens) small granules anterior to the supraoculars; (5) ventral scales in ten (rarely nine or 11) longitudinal rows; (6) variable color pattern, usually with three dark dorsal stripes faded posteriorly. Among Angolan Pedioplanis , this species seems to be the most variable in terms of color pattern. The dorsum is greyish brown anteriorly and pale brown to brick red posteriorly, usually with dark stripes starting at the back of the head and fading between midbody and two-thirds along the back. The vertebral stripe splits or widens at the neck, or may be entirely split into two thinner stripes, with a greyish interspace. Dorsolateral stripes are usually wider and more distinct than the vertebral one. On the flanks there is a dark lateral stripe that starts behind the eye and usually fades posteriorly, often faint and reticulated, with a series of yellow to blueish circles running along its lower edge. However, the extension and distinctiveness of dorsal stripes is quite variable, being completely absent in some specimens, and extending nearly to the base of the tail in others. Hind limbs and tail reddish-brown to brick red, with the hindlimbs sometimes covered above by faint pale circles. White ventrally, sometimes reddish at the base of the tail.

Comparison with other Pedioplanis species. Pedioplanis haackei is readily distinguished from P. burchelli , P. laticeps , P. breviceps , P. namaquensis and P. husabensis by the presence of two enlarged transparent scales on the lower eyelid (versus eight or more opaque to semi-transparent scales in other species); from P. lineoocellata by the presence of an enlarged tympanic shield (versus no enlarged tympanic shield in P. lineoocellata ); from P. inornata , P. rubens , P. gaerdesi and P. branchi by the presence of dorsal stripes (versus no stripes in P. inornata , P. rubens , P. gaerdesi and P. branchi ); from P. undata by color pattern (bold anterior dorsal striping that fades posteriorly in P. haackei versus dorsal striping bold or not, may be reduced with pale longitudinal elements or even a single middorsal stripe restricted to the nape in P. undata ) and a greater maximum number of granules anterior to the supraoculars (7–28 in P. haackei versus 8–13 in P. undata ); and from P. mayeri by color pattern (dorsal stripes usually faded posteriorly in P. haackei versus bold and distinct all the way to the tail in P. mayeri ), and a greater maximum number of granules anterior to the supraocular (7–28 in P. haackei versus 9–16 in P. mayeri ). It can be further distinguished from P. undata and P. mayeri by geographic location ( P. haackei restricted to Angola versus P. mayeri and P. undata restricted to Namibia). With respect to Angolan congeners, P. haackei is distinguished from P. benguelensis by color pattern (posterior dorsum usually reddish with faded stripes in P. haackei versus bold stripes all the way to the tail in P. benguelensis ); it is distinguished from P. huntleyi by a smaller average SVL (mean 45 mm in P. haackei versus 54 mm in P. huntleyi ), a higher number of granules anterior to the supraoculars (usually> 11 in P. haackei versus <12 in P. huntleyi ) and two rows of granules between supraoculars and supraciliaries (versus one in P. huntleyi ). It is distinguished from an undescribed species by the presence of two transparent scales on the lower eyelid (versus one, see description below).

Holotype: PEM R 18465*, an adult male collected along the road to Tambor , Namibe Province [-15.87606°, 12.20583°, 196 m], by William R. Branch, G. John Measey, Werner Conradie and Krystal A. Tolley on 19 January 2009.

Additional material. 121 specimens: Benguela Province: 2 km from Lucira-Benguela road, towards Cape Santa Maria [-13.51167°, 12.62833°, 285 m] ( PEM R 24087) ; Huíla Province: 13 km N of Quilengues , on road to Benguela [-13.97247°, 14.04717°, 936 m] ( PEM R 21661*) ; Namibe Province: 10 km S of Lake Arco [- 15.83044°, 12.14125°, 121 m] ( PEM R 18470, 18471 *, 18472*) ; on the road to Benguela, 2 km N of turning to Lucira [-13.90528°, 12.54417°, 312 m] ( AG 98 *) ; road to Bentiaba, 52 km N of junction with Lubango-Namibe road [-14.65806°, 12.52717°, 586 m] ( PEM R 21662*, 21663*) ; 5 km along fence leading to Baptista farm [-16.05847°, 12.42597°, 343 m] ( PEM R 18468) ; Humpata-Namibe road, 7.8 km from bottom of Leba pass [- 15.04467°, 13.15947°, 642 m] ( PEM R 18460*, 21664*) ; along road to Tambor [-15.88778°, 12.36417°, 300 m] ( PEM R 18464*) , [-15.87606°, 12.20583°, 196 m] ( PEM R18466–18467 ) ; base of Serra da Neve, on dirt road to Quilengues [-13.81594°, 13.32644°, 892 m] ( MHNC-UP / REP 645 ) ; base of Serra da Neve , on dirt road to the top of the mountain [-13.83424°, 13.27669°, 803 m] ( MHNC-UP / REP 634-635 , CAS 266123 ) ; Catara River valley [-13.62417°, 12.83556°, 454 m] ( PEM R 24057*, 24058, 24059, 24060, 24061*, 24062*) ; Caraculo [-15.01651°, 12.64251°, 487 m] ( CAS 264769 *) ; Chipumpo-Virulundo [-16.19125°, 12.85595°, 539 m] ( CAS 264805 *) ; Curoca-Omahua [-15.81918°, 12.14273°, 100 m] ( CAS 264774 *, 264775*, 264776), [-15.95026°, 12.49531°, 394 m] ( CAS 264777 *–264779*) ; dry river valley, approximately 10 km S of turning to Lucira [-13.96389°, 12.53222°, 137 m] ( PEM R 24223, 24224 ) ; Lucira Dam [-13.87889°, 12.56556°, 339 m] ( PEM R 24103*, 24104) ; Mucungo farm [-14.78111°, 12.48983°, 296 m] ( MHNCUP / REP 633 ) , [-14.77890°, 12.48745°, 305 m] ( CAS 266248 ) ; Munhino [-14.92000°, 13.00000°, 393 m] ( MD 1918 b) ; near Virulundo [-16.28523°, 12.94192°, 718 m] ( CAS 264806 *– 264808*, 264810*) ; near Curoca River [-16.24468°, 12.34068°, 265 m] ( CAS 264781 *) ; off road 20 km N of Omauha Lodge [-16.07414°, 12.43328°, 352 m] ( PEM R 18474*) ; Omauha-Virei [-16.14506°, 12.59460°, 538 m] ( CAS 264791 *) ; Omauha Lodge [-16.19858°, 12.40073°, 340 m] ( CAS 264780 *, 264784*–264790*, MHNC-UP / REP 644 , INBAC / AMB 10416*), [-16.20033°, 12.40003°, 343 m] ( CAS 254835 *) ; Omauha Lodge, N of entrance to Iona National Park [-15.99694°, 12.40694°, 301 m] ( PEM R 18473) ; Red Canyons near Lake Arco [-15.74597°, 12.13989°, 81 m] ( PEM R 18461*, 18462, 18463) ; road from Lake Arco to Espinheira [-15.91356°, 12.39519°, 319 m] ( PEM R 18475*) ; road to Tambor [-15.88119°, 12.22267°, 192 m] ( PEM R 18469*) ; sandstone hill near Piambo [-14.65833°, 12.36944°, 365 m] ( PEM R 22041) ; vicinity of N’Dolondolo [-13.80866°, 13.13521°, 731 m] ( CAS 264757 *, 264765*) ; Virei-Virulundo [-16.31018°, 12.79597°, 471 m] ( CAS 264794 *, 264801*, 264803*) ; 10 km E of Munhino [-14.87821°, 13.13195°, 566 m] ( TM 46726 ) ; 12 km N of Furnas [-15.02513°, 12.18828°, 69 m] ( TM 40570 ) ; 12 km N of Tambor [-15.96780°, 12.39581°, 262 m] ( TM 40496 ) ; 23 km W of Virei [-15.66886°, 12.74089°, 397 m] ( TM 41025 ) ; 30 km N of Tambor [-15.89287°, 12.37609°, 312 m] ( TM 40489–40491 ) ; 35 km S of Moçâmedes [-15.48000°, 12.21000°, 150 m] ( MD 1946 ) ; 3 km E of Chicambi village [-13.89556°, 12.78222°, 612 m] ( AG 69 *–71*) ; 5 km (by road) NW of Pico Azevedo [-15.47547°, 12.46219°, 408 m] ( CAS 254763 *, 254764*), [-15.47600°, 12.46150°, 399 m] ( CAS 254767 *) ; 60 km (by road) E of Tômbua [-15.87760°, 12.21668°, 189 m] ( TM 40451–40453 ) ; Assunção [-14.86667°, 13.10000°, 505 m] ( TM 40181 ) ; 17 km W of Chicamba village [-13.91722°, 12.68111°, 531 m] ( AG 84 *) ; Inamangando farm [-14.05114°, 12.42733°, 24 m] ( TM 41157 , 41158 ) ; Iona National Park , N of Tambor [-15.99539°, 12.40647°, 306 m] ( CAS 254840 *) ; Iona National Park, Curoca river crossing [-16.30000°, 12.43333°, 198 m] ( TM 40580 ) ; Lungo [-14.66667°, 13.25000°, 679 m] ( TM 24397 , 24398 , 24400 ) ; Moçâmedes [-15.16667°, 12.16667°, 6 m] ( TM 22844–22845 ) ; Pediva Hot Springs , S side of the river [-16.29381°, 12.56033°, 235 m] ( CAS 254860 *) ; Pico Azevedo [-15.53400°, 12.49197°, 359 m] ( CAS 254939 ) ; Namibe Nature Reserve [-15.77292°, 12.33269°, 262 m] ( CAS 254951 *, 254953*) ; Saiona River [-15.40000°, 13.20000°, 534 m] ( TM 40983–40985 ) ; Tambor [-16.13556°, 12.42972°, 379 m] ( TM 40501 , 40502 ) , [-16.06710°, 12.42970°, 348 m] ( INBAC / AMB 10393*) .

The following specimens were tentatively assigned to this species based on a combination of morphological characters and geographic location (10 specimens): Benguela Province: 2 km turning on Cape Santa Maria from Lucira-Benguela coastal road [-13.51167°, 12.62833°, 285 m] ( PEM R 24086, 24088 ) ; Namibe Province: 13.6 km S of Mucungo Farm [-14.85528°, 12.42889°, 400 m] ( PEM R 24153) ; 1 km SE of Mucungo Farm [-14.78667°, 12.49611°, 303 m] ( PEM R24133–24135 ) ; 8.8 km S of Mucungo Farm [-14.80167°, 12.41917°, 579 m] ( PEM R 24160) ; Catara River valley [-13.62278°, 12.80750°, 454 m] ( PEM R 27734) ; Lucira [-13.86667°, 12.53333°, 51 m] ( TM 24430 , 41190 ) .

Distribution and habitat. Pedioplanis haackei occurs over most of Namibe Province north of -16º latitude, extending marginally into southern Benguela Province, and northwestern Huíla Province below 1000 m ( Fig. 7 ). It is absent from the more xeric areas of southwestern Namibe Province, as well as southeastern Namibe Province, where it is replaced by P. huntleyi , with both species occurring in sympatry near Virulundo and Pediva Hot Springs. Pedioplanis haackei is sympatric with P. benguelensis in coastal areas, and with the undescribed species in several localities along most of its range. This species inhabits a vast area covering different habitat types, from more xeric areas to areas dominated by Mopane woodlands ( Grandvaux-Barbosa 1970; Fig. 5 ).

Figures

FIGURE 5. Typical habitats of Angolan Pedioplanis. A. Dry coastal savanna with sparse vegetation at Chimalavera Natural Regional Park, Benguela Province (P. benguelensis). Photo by Luis M.P. Ceríaco. B. Rocky outcrops near Omauha Lodge, Namibe Province (P. haackei). Photo by Ishan Agarwal. C. Mopane dominated shrubland near Capangombe, Namibe Province (P. serodioi sp. nov.). Photo by Luis M.P. Ceríaco. D. Mopane shrubland near Oncocua, Cunene Province (P. huntleyi). Photo by Luis M.P. Ceríaco.

FIGURE 6. Live specimens of Pedioplanis haackei. Above: CAS 266123 from the base of Serra da Neve, Namibe Province, with typical coloration. Photo by Luis M.P. Ceríaco. Below: CAS 264795 from Virei-Virulundo, Namibe Province, with indistinct dorsal stripes. Photo by Ishan Agarwal.

FIGURE 7. Geographic distribution of Pedioplanis haackei based on museum vouchers. Localities represented in the phylogenetic analysis are indicated with a black center. White star denotes the type locality. Major cities are represented by a black square.