Pedioplanis serodioi, Parrinha & Marques & Heinicke & Khalid & Parker & Tolley & Childers & Conradie & Bauer & Ceríaco, 2021

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 : 32-38

Pedioplanis serodioi sp. nov.

( Figs. 10–13 )

Eremias benguelensis: Boulenger (1918: 5 ; 1921: 287); Parker (1936: 134); Mertens (1954: 177)

Eremias benguellensis: Monard (1937: 72) ; Mertens (1971: 59)

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

Mesalina benguelensis: Szczerbak (1975: 24)

Pedioplanis benguelensis: Arnold (1989: 213) ; Mayer (1989: 135)

Pedioplanis benguellensis: Arnold (1991: 785) ; Branch (1998: 173); Conradie et al. (2012b: 95); Ceríaco et al. (2016: 37) ; Marques et al. (2018: 222) [part]

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

The first records of this species were probably those provided by Bocage (1895) as Eremias namaquensis from “Capangombe par M. d’Anchieta” and “l’intérieur de Mossamedes par MM. Capello et Ivens”. However, earlier records as Eremias benguelensis from “Benguella” ( Bocage 1867a, 1867b) could have also represented this taxon. Nevertheless, the loss of these specimens precludes any further inferences regarding their taxonomic identity. The only exception is a specimen collected by Anchieta in Capangombe (MHNCUP/REP 0231, Fig. 11 ) and still deposited in the collections of MHNC-UP, which represents the first confirmed record of this species.

Boulenger’s (1918, 1921) diagnosis of “ benguelensis ” was characterized by the presence of a single transparent scale on the lower eyelid, which was present in his specimens from “Huxe” [= Uche, Benguela Province] and “Ponang Kuma” [= Dongoena, Cunene Province], collected by Ansorge. Boulenger’s (1918, 1921) diagnosis was followed by most authors (e.g., Monard 1937; Szczerbak 1975; Mayer 1989; Branch 1998) and Parker (1936) reported a specimen (BMNH 1936.8.1.514) from “Catengue”, Benguela Province, collected during the Karl Jordan’s expedition to South-West Africa and Angola and deposited in the collections of the Natural History Museum, London. Laurent (1964) reported additional Angolan material under the name Eremias undata undata , including among his records two specimens with a single transparent scale on the lower eyelid, collected by Barros Machado in Munhino, Namibe Province (MD 1918a), and Quilengues, Huíla Province (MD 1970-1).

Following Boulenger’s (1918, 1921) diagnosis and the lack of topotypic comparative material, Conradie et al. (2012b) conservatively assigned specimens collected in central and southeastern Namibe Province to P. benguelensis , providing the first molecular data for this species. Three additional specimens conforming to Boulenger’s (1918, 1921) diagnosis of P. benguelensis were collected by Ceríaco et al. (2016) near Caraculo, Namibe Province. The collection of true P. benguelensis from the coastal areas of Benguela Province allowed for a reassessment of Angolan populations assigned to this species. Our analysis revealed species-level phylogenetic distance as well as consistent morphological differences from all named species of Pedioplanis . Based on the evidence produced in this study, we here describe a new species of Angolan Pedioplanis characterized by the presence of a single transparent scale on the lower eyelid (see P. benguelensis account for discussion on the name allocation decision).

Diagnosis: Pedioplanis serodioi sp. nov. is a rather small Pedioplanis , with an average SVL of 44 mm (max 50 mm) and a tail length roughly twice the SVL ( Fig. 10 ). It can be distinguished from other Pedioplanis species in Angola and neighboring regions by the following combination of characters: (1) lower eyelid with a single blackedged, enlarged transparent scale; (2) four (rarely three, five or six) supralabials anterior to the subocular and two (rarely three) posteriorly; (3) One (sometimes two) row of small granules between the supraoculars and supraciliaries; (4) a group of 6–24 (<15 in 75% of specimens) small granules preceding the supraoculars; (5) ventral scales in ten longitudinal rows; (6) a pair of irregularly edged dorsolateral stripes or series of spots and a faint or absent vertebral stripe. The dorsum is greyish brown to pale grey, often with extensive black speckling throughout. There may be a thin, dark vertebral stripe, but this is usually faint and indistinct, or reduced to irregular black streaks. A pair of irregularly edged, black dorsolateral stripes extend from the back of the head to the base of the tail, often broken into series of black spots on a reddish-brown background. A pale line separates the dorsolateral stripes from a lateral stripe that starts behind the eye and extends to the base of the tail, often faint and reticulated, with a series of yellow spots running along its lower edge. There is often a reddish stripe extending from the posterior labials to the hindlimb insertion, up to four scales wide, continuous or broken into series of streaks. The hindlimbs are often covered above by pale circles surrounded by black pigmentation. Venter white.

Comparison with other Pedioplanis species. Pedioplanis serodioi sp. nov. is distinguished from P. gaerdesi by the number of supralabials anterior to the subocular (usually four in P. serodioi sp. nov. versus five to six in P. gaerdesi ) and color pattern (dark dorsolateral stripes or series of spots in P. serodioi sp. nov. versus no dorsal stripes in P. gaerdesi ); and from all the remaining species of the genus by consistently possessing a single transparent scale on the lower eyelid (versus two or more scales in other species). Regarding Angolan congeners, it can be further distinguished from P. benguelensis by a lower number of supralabials anterior to the subocular (usually four in P. serodioi sp. nov. versus five in P. benguelensis ), one row of granules between supraoculars and supraciliaries (versus two in P. benguelensis ), fewer granules anterior to the supraoculars (usually <15 in P. serodioi sp. nov. versus> 13 in P. benguelensis ) and color pattern (dorsolateral stripes often broken and vertebral stripe indistinct in P. serodioi sp. nov. versus all stripes distinct and well-defined in P. benguelensis ); from P. haackei by the presence of one row of granules between supraoculars and supraciliaries (versus two in P. haackei ), fewer granules anterior to the supraoculars (usually <15 in P. serodioi sp. nov. versus> 11 in P. haackei ) and color pattern (dorsolateral stripes often broken and vertebral stripe indistinct in P. serodioi sp. nov. versus dorsal stripes continuous and faded posteriorly in P. haackei ); and from P. huntleyi by a smaller SVL (average 43 mm in P. serodioi sp. nov. versus 54 mm in P. huntleyi ), a lower number of supralabials anterior to the subocular (four in P. serodioi sp. nov. versus five in P. huntleyi ) and color pattern (dorsolateral stripes often broken and vertebral stripe indistinct in P. serodioi sp. nov. versus dorsal stripes continuous and faded posteriorly in P. huntleyi ).

Holotype. CAS 254906 (field number JVV 8598 , Fig. 12 ), adult male collected on the north side of the road from Namibe to Lubango, road marker 59, 1.8 km West (by road) of Caraculo [-15.01888°, 12.64014°, 491 m], Namibe Province, by Luis M.P. Ceríaco, E.L. Stanley, Arianna L. Kuhn, Jens V. Vindum, Sango de Sá , Suzana Bandeira and Hilária Valério on 6 December 2013.

Paratypes. 11 specimens: CAS 254909 * (field number JVV 8601 ), with the same collecting data as the holotype . CAS 264752 * (field number AMB 10181) from Bibala Crossroad [-14.74398°, 13.32852°, 863 m], collected by Luis M.P. Ceríaco, Suzanna Bandeira and Ishan Agarwal on 18 November 2016 . CAS 264767 * and CAS 264768 * (field numbers AMB 10364 and 10365, respectively) from Bibala Crossroad [-14.74398°, 13.32852°, 863 m], collected by Luis M.P. Ceríaco, Suzanna Bandeira and Ishan Agarwal on 23 November 2016 . CAS 264754 * (field number AMB 10183) from Camacuio [-14.11365°, 13.24320°, 670 m], collected by Luis M.P. Ceríaco, Suzanna Bandeira and Ishan Agarwal on 18 November 2016 . CAS 264766 * (field number AMB 10334) from N’Dolondolo [-13.81328°, 13.13618°, 681 m], collected by Luis M.P. Ceríaco, Suzanna Bandeira and Ishan Agarwal on 22 November 2016 . MHNCUP / REP 0231 , collected by José d’Anchieta in Capangombe [-14.93309°, 12.96477°, 422 m] . PEM R 18449* and 18450* (field numbers MBUR 02146 and 02147, respectively), collected by Werner Conradie, William R. Branch, Krystal A. Tolley, G. John Measey among rocks and boulders by roadside 50 km E of Namibe [- 15.01661°, 12.55650°, 516 m] on 18 January 2009 . PEM R 18453* (field number MBUR 02165 ), collected by Dirk Bellstedt among rocks and boulders by roadside 50 km E of Namibe [-15.01958°, 12.52458°, 550 m] on 18 January 2009 . PEM R 18459* (field number WC09-33) from east of Moimba [-16.71833°, 13.12194°, 834 m], collected by Werner Conradie on 24 January 2009 .

Additional material. 70 specimens: Benguela Province: Chimalavera Nature Reserve, vicinity of Main Camp [-12.83377°, 13.16991°, 293 m] ( INBAC / AMB 9967 *) ; Catengue [-13.06222°, 13.75611°, 586 m] ( PEM R 27699, 27700 ) ; 7 km east of Catengue, on road to Cubal [-12.99942°, 13.79856°, 814 m] ( PEM R 21659*, 21660) ; 30 km SSW of Benguela [-12.74389°, 13.15167°, 55 m] ( PEM R 22071, 22072 ) ; 2 km turning on Cape Santa Maria from Lucira-Benguela coastal road [-13.51167°, 12.62833°, 285 m] ( PEM R 24089) ; Maconjo, near Benguela-Catengue road [-12.85472°, 13.39472°, 399 m] ( PEM R 27701) ; 15 km E of Lobito [-12.37056°, 13.72833°, 252 m] ( PEM R 27702) ; Namibe Province: Capangombe [-15.09738°, 13.13909°, 542 m] ( CAS 264240 ) ; base of Serra da Neve , on dirt road to the top of the mountain [-13.83424°, 13.27669°, 803 m] ( MHNC-UP / REP 636 , CAS 266122 ) ; base of Serra da Neve, on dirt road to Quilengues [-13.81594°, 13.32644°, 892 m] ( MHNC-UP / REP 637 , 638 ) ; base of Serra da Neve, Malowe village [-13.83549°, 13.27547°, 798 m] ( MHNC-UP / REP 639 , 646 ) ; Munhino [-14.96667°, 12.96667°, 393 m] ( MD 1918 a) ; 50 km E of Moçâmedes on main road to Leba [-15.01558°, 12.55503°, 516 m] ( PEM R 21653*, 21654*, 21655*, 21656*, 21657*) ; Camacuio [-14.11365°, 13.24320°, 670 m] ( CAS 264755 *) ; 10 km W of Lola, road northwest to Camacuio [-14.29028°, 13.53056°, 802 m] ( PEM R 24006*, 24007*, 24008, 24009, 24010*, 24011, 24012) ; 30 km W of Lola, road northwest to Camacuio [-14.27583°, 13.45806°, 791 m] ( PEM R 24032*–24035*) ; just before Camacuio [-14.14333°, 13.27556°, 712 m] ( PEM R 24050*–24052*) ; 50 km E of Namibe by roadside [-15.01661°, 12.55650°, 516 m] ( PEM R 18451, 18452 ) ; approximately 20 km S of Bentiaba [-14.40278°, 12.44972°, 426 m] ( PEM R 24220) ; approximately 20 km SW of Camacuio [-14.21472°, 13.40556°, 803 m] ( PEM R 27733*) ; road between Namibe and Humpata 47km E of Namibe [-15.01942°, 12.52881°, 527 m] ( PEM R 18454*, 18455–18458) ; road north of Bibala towards Lola [-14.41611°, 13.56472°, 920 m] ( PEM R24026– 24027 ) ; Bibala Crossroad [-14.74398°, 13.32852°, 863 m] ( CAS 264753 ) ; Namibe-Lubango road, 1.8 km W (by road) of Caraculo [-15.01888°, 12.64014°, 491 m] ( CAS 254896 *) ; Caraculo [-15.01667°, 12.66667°, 463 m] ( TM 40211 ) ; 14 km NE of Caraculo [-14.91304°, 12.73356°, 446 m] ( TM 40210 , 40254–40262 ) ; Leba Pass bottom [- 15.04108°, 13.19029°, 696 m] ( TM 46756 ) ; Lungo [-14.66667°, 13.25°, 679 m] ( TM 24399 ) ; 10 km E of Munhino [-14.87821°, 13.13195°, 566 m] ( TM 46727–46728 ) ; Pico do Azevedo [-15.55000°, 12.51667°, 347 m] ( TM 41071 , 41072 ) ; Huíla Province: 15 km N of Quilengues [-13.94208°, 14.04557°, 927 m] ( MD 1970-1 ) ; between Chingoroi and Quilengues [-13.87278°, 13.96528°, 926 m] ( PEM R 22068) .

Description of the Holotype. The holotype is an adult male with a complete original tail ( Fig. 12 ). SVL 48 mm; TL 104 mm. Body relatively stout (SVL/HL 4.29), with hindlimbs longer than forelimbs and tail 2.16 times as long as the SVL. Moderately sized head (HL/SVL 0.23), distinct from the neck. Other relevant measurements are presented in Table 5. Rostral wider than high, visible from above. Nostril pierced between three scales; supranasals slightly swollen and in broad contact with one another behind rostral; infranasal in contact with rostral, first supralabial and anterior loreal; postnasal small and subquadrangular, placed between supranasal, infranasal, anterior loreal and frontonasal. Frontonasal hexagonal, as long as it is wide. Prefrontals in broad contact with each other, the loreals, frontonasal and frontal. Two loreals, posterior largest. Frontal longer than wide, narrower posteriorly, in contact with prefrontals anteriorly, supraoculars laterally and frontoparietals posteriorly. Paired frontoparietals in broad median contact, touching frontal and posterior supraocular anteriorly, and the parietals and interparietal posteriorly. Interparietal longer than broad, rounded anteriorly, its posterior margin much narrower than the anterior. Occipital small and slightly wider than it is long, posterior margin more than twice as wide as anterior, in broad contact with interparietal. Parietals longer than wide, in contact with frontoparietals, interparietal and occipital. Two rounded supraoculars in contact with each other and the frontal, preceded by a group of 14 (right side) and 13 (left side) small granules, those in contact with the frontal and prefrontal largest. One row of small granules between anterior supraocular and supraciliaries, increasing to two rows posterior to supraocular suture. Supraciliaries six, the first longest. Temporal scales small and granular. One narrow and elongated tympanic shield at anterodorsal edge of the ear opening. Subocular bordering lip, upper margin much wider than lower. Five (right side) and four (left side) supralabials anterior to subocular and two posterior to subocular. Lower eyelid with a single black-edged, enlarged transparent scale. Infralabials six. Mental wider than long, in contact with first infralabials and first pair of chin shields. Four pairs of chin shields, first three in median contact and fourth largest. Gular scales 26, in a straight line between symphysis of the chin shields and median collar plate. Collar free, comprising six enlarged plates. Ventral scales smooth, in nine longitudinal and 27 transverse rows; outer scales smaller than others; first complete transverse row posterior to collar notably longer than others. Precloacal scales irregular and subequal, with a single scale larger than the other in the center. Femoral pores 13 on each leg. Lamellae under fourth toe 23. Dorsal scales small and granular, larger towards the ventral scales. Upper forelimb covered above by large hexagonal plates; forearm covered above by slightly imbricate and keeled scales, larger than dorsal scales, and below by enlarged plates. Hindlimbs covered above by slightly imbricate and keeled scales, larger than dorsal scales, and below by enlarged plates. Scales on tail diagonally keeled, except for those on ventral side of basal portion, which are smooth.

Coloration (in preservative). Dorsum pale greyish with some black speckling, especially on the limbs and tail; hindlimbs covered above by pale circles surrounded by black pigmentation. There is a barely distinct, greyish vertebral stripe that splits at the neck, as well as a pair of dorsolateral series of black spots extending to the base of the tail. Lateral stripe somewhat faint and reticulated, with a series of pale circles running along its lower edge. Tail greyish above, with some black speckling and dark keeled scales. White ventrally.

Variation in the type series. Variation in scalation and measurements among the type series of Pedioplanis serodioi sp. nov. is reported in Table 5. Except for minor differences in scalation and color pattern, all paratypes generally agree with the holotype. The prefrontals are separated by a small azygous scale in MHNCUP / REP 0231 , CAS 254909 , CAS 264766 , PEM R 18453 and PEM R 18459, while in CAS 264768 and CAS 264752 there are two longitudinally aligned scales between the prefrontals. Unlike the holotype, dorsolateral stripes are continuous, but irregularly outlined, in CAS 254909 , CAS 264752 , CAS 264768 , CAS 264766 , PEM R 18453, and indistinct in MHNCUP / REP 0231 . The remaining paratypes agree with the holotype in having broken (at least partially) dorsolateral stripes .

Distribution and habitat. Pedioplanis serodioi sp. nov. has a broad distribution in the low elevation areas of southwestern Angola, from central Benguela Province to western Cunene Province, except for the more xeric areas of southwestern Namibe Province ( Fig. 13 ). Voucher specimens from northern Namibia deposited in the Ditsong National Museum of Natural History (TM 38789–95 from Okjivakandu; TM 33292 and 38903 from Opuwo; TM 38868, 38870–71 from Otjiwise) are provisionally referred to Pedioplanis cf. serodioi sp. nov. based on the presence of a single transparent scale on the lower eyelid and dorsal coloration, but the identity of these populations is pending genetic confirmation (see Discussion). The current gaps in the distribution of P. serodioi sp. nov. are likely an artifact of incomplete sampling, and further surveys in southern Angola and adjacent northern Namibia shall improve our knowledge of the distribution range of this species. At 1121 m above sea level, the specimens collected by Ansorge in “Ponang Kuma” [= Dongoena, Cunene Province] ( Boulenger 1921) represent the highest elevation recorded for any Angolan Pedioplanis . It occurs sympatrically with all other Angolan congeners in several localities across 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 ).

Etymology. The species is named after the Angolan scholar João Manuel Serôdio de Almeida (1943–present), professor of the Biology Department of the Faculty of Sciences of Agostinho Neto University, Luanda, Angola. João Serôdio has had a pivotal role in scientific research and biodiversity conservation in Angola, as manager of several conservation areas, Vice-Minister for the Environment (1997–2000) and by training several generations of Angolan biologists. Prof. Serôdio has been a strong supporter and advocate of the present herpetological research in Angola. The name is formed in the genitive masculine singular. We propose the English common name of Serôdio’s Sand Lizard, and the Portuguese common name of Lagartixa da Areia de Serôdio.

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 10. Live specimens of Pedioplanis serodioi sp. nov.. Above: PEM R24010*, male found 10 km west of Lola, on the road to Camacuio, Namibe Province. Photo by William R. Branch. Below: female paratype (CAS 264752) from Bibala Crossroad, Namibe Province. Photo by Ishan Agarwal.

FIGURE 11. Paratype of Pedioplanis serodioi sp. nov. (MHNCUP/REP 0231), collected by Anchieta from Capangombe, Namibe Province. Photos by Bruna Santos.

FIGURE 12. Holotype of Pedioplanis serodioi sp. nov. (CAS 254906*, field number JVV 8598). Photos by Luis M.P. Ceríaco.

FIGURE 13. Geographic distribution of Pedioplanis serodioi sp. nov. 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.