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Panaspis annobonensis (Fuhn, 1972) Figure 11C Panaspis africana annobonensis Fuhn, 1972. Type locality: “ Ile Annobon, Golfe de Guinée”. Panaspis annobonensis (Fuhn, 1972): Perret 1973: 605. Leptosiaphos annobonensis (Fuhn, 1972): Haft 1993: 59. Afroablepharus annobonensis (Fuhn, 1972): Jesus et al. 2007. Distribution. Endemic to Annobon (Fuhn 1972; Jesus et al. 2007; Soares et al. 2018; Ceríaco et al. 2018) (Map 12B). Comments. The species level status was supported by Jesus et al. (2007). It is more closely related to P. cabindae than to the other congeneric species from the Gulf of Guinea islands (Soares et al. 2018). Specimens examined. Eighteen specimens. Annobon: 1987 (EBD 27702 –27703); 10 November 1987 (EBD 27701, EBD 27699); Pico de Fuego, July 1978 (EBD 23580 –23583); “lowland”, July 1978 (EBD 23584 –23593)., Published as part of Sánchez-Vialas, Alberto, Calvo-Revuelta, Marta & Riva, Ignacio De La, 2022, Synopsis of the terrestrial Reptiles of Equatorial Guinea, pp. 1-197 in Zootaxa 5202 (1) on page 50, DOI: 10.11646/zootaxa.5202.1.1, http://zenodo.org/record/7285600, {"references":["Fuhn, I. E. (1972) Revision du phylum forestier du genre Panaspis Cope (Reptilia, Scincidae, Lygosominae). Revue Roumaine de Biologie, Serie Zoologie, 17 (4), 257 - 271.","Perret, J. - L. (1973) Contribution a l'etude des Panaspis (Reptilia, Scincidae) d'Afrique occidentale avec la description de deux especes nouvelles. Revue Suisse de Zoologie, 80, 595 - 630. https: // doi. org / 10.5962 / bhl. part. 75958","Haft, J. (1993) Ein Beitrag zur Biologie der Echsen der Insel Sao Tome (Golf von Guinea), mit naherer Betrachtung zur Systematik von Leptosiaphos africana (Gray) (Reptilia: Sauria: Geckonidae et Scincidae). Faunistische Abhandlungen, Dresden, 19 (1 - 16), 59 - 70.","Jesus, J., Harris, D. J. & Brehm, A. (2007) Relationships of Afroablepharus Greer, 1974 skinks from the Gulf of Guinea islands based on mitochondrial and nuclear DNA: Patterns of colonization and comments on taxonomy. Molecular Phylogenetics and Evolution, 45 (3), 904 - 914. https: // doi. org / 10.1016 / j. ympev. 2007.08.011","Soares, L. B., Ceriaco, L. M. P., Marques, M. P., Bastos-Silveira, C., Scheinberg, L. A., Harris, D. J., Brehm, A. & Jesus, J. (2018) Review of the leaf-litter skinks (Scincidae: Panaspis) from the Gulf of Guinea Islands, with the description of a new species. African Journal of Herpetology, 67 (3), 1 - 28. https: // doi. org / 10.1080 / 21564574.2017.1413015","Ceriaco, L. M., Marques, M. P. & Bauer, A. M. (2018) Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of Sao Tome, Principe and Annobon islands. Zootaxa, 4387 (1), 91 - 108. https: // doi. org / 10.11646 / zootaxa. 4387.1.4"]}

Panaspis breviceps (Peters, 1873) Figure 11B Euprepes (Mabuia) breviceps Peters, 1873: 604. Type locality: “ Gabun ”. Gabon. Lygosoma breviceps (Peters, 1873): Boulenger 1887: 300. Mabuia batesi Boulenger, 1900: 449. Type locality: “Benito River” Río Benito, Río Muni, Equatorial Guinea. Lygosoma (Leiolopisma) buchneri Werner, 1909: 62. Type locality: “ Kamerun ”. Lygosoma (Leiolopisma) auricillatum Chabanaud, 1917a: 98. Type locality: “ Congo: Lambaréné ”. Emoia breviceps (Peters, 1873): Loveridge 1936b: 71. Riopa (Panaspis) breviceps (Peters, 1873): Smith 1937: 229 Panaspis breviceps (Peters, 1873): Mittleman 1952: 17–23. Distribution. Central Africa, from Cameroon to Central African Republic and southward to northern Angola. In Equatorial Guinea it has been recorded in Bioko at Basilé (Boulenger 1906), Ureca (Mertens 1964a), and at a beach (likely in Aleñá) across from the Island of Loros (Mertens 1964a), and in Río Muni at Benito River (Boulenger 1900; Perret 1973), and Monte Alén National Park (Lasso et al. 2002), including Monte Mitra forest (Gonwouo & Nsang 2005) (Map 12A). Comments. A hypothesis on the phylogenetic relationships of this species was provided by Schmitz et al. (2005). It is phylogenetically allied to Panaspis cf. togoensis from Cameroon (Schmitz et al. 2005). Specimens examined. Two specimens. Río Muni: San Joaquín de Ndyiacom, 1990 (EBD 31521 _B-9523); imprecise locality (from Nsork, Nvom or Campo River), 1986–1987 (EBD 25051)., Published as part of Sánchez-Vialas, Alberto, Calvo-Revuelta, Marta & Riva, Ignacio De La, 2022, Synopsis of the terrestrial Reptiles of Equatorial Guinea, pp. 1-197 in Zootaxa 5202 (1) on page 47, DOI: 10.11646/zootaxa.5202.1.1, http://zenodo.org/record/7285600, {"references":["Peters, W. C. H. (1873) Uber eine neue Schildkrotenart, Cinosternon effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der Koniglich preussischen Akademie der Wissenschaften zu Berlin, 1873, 603 - 618.","Boulenger, G. A. (1887) Catalogue of the Lizards in the British Museum (Natural History), Vol. 3. The Trustees of the British Museum, London, 575 pp., 40 pls.","Boulenger, G. A. (1900) A list of the batrachians and reptiles of the Gaboon (French Congo), with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1900, 433 - 456.","Werner, F. (1909) Beschreibung neuer Reptilien aus dem Kgl. Naturalienkabinett in Stuttgart. Jahreshefte des Vereins fur vaterlandische Naturkunde in Wurttemberg, 65, 55 - 63.","Chabanaud, P. (1917 a) Enumeration des reptiles non encore etudies de l'Afrique occidentale, appartenant aux collections du Museum, avec la description des especes nouvelles. Bulletin du Museum National d'Histoire Naturelle, Paris, 23, 83 - 105.","Loveridge, A. (1936 b) African reptiles and amphibians in the Field Museum of Natural History. Zoological series Field Museum of Natural History, Chicago, 22 (1), 1 - 122. https: // doi. org / 10.5962 / bhl. title. 3069","Smith, M. A. (1937) A review of the genus Lygosoma (Scincidae: Reptilia) and its allies. Records of the Indian Museum, 39 (3), 213 - 234. https: // doi. org / 10.26515 / rzsi / v 39 / i 3 / 1937 / 162283","Mittleman, M. B. (1952) A generic synopsis of the lizards of the Subfamily Lygosominae. Smithsonian miscellaneous collections, 117, 1 - 35.","Boulenger, G. A. (1906) Report on the reptiles collected by the late L. Fea in West Africa. Annali del Museo Civico di Storia Naturale di Genova, 3 (2), 196 - 216.","Mertens, R. (1964 a) Die Reptilien von Fernando Poo. Bonner zoologische Beitrage, 15, 211 - 238.","Perret, J. - L. (1973) Contribution a l'etude des Panaspis (Reptilia, Scincidae) d'Afrique occidentale avec la description de deux especes nouvelles. Revue Suisse de Zoologie, 80, 595 - 630. https: // doi. org / 10.5962 / bhl. part. 75958","Lasso, C. A., Rial, A. I., Castroviejo, J. & De la Riva, I. (2002) Herpetofauna del Parque Nacional de Monte Alen (Rio Muni, Guinea Ecuatorial). Graellsia, 58, 21 - 34. https: // doi. org / 10.3989 / graellsia. 2002. v 58. i 2.276","Gonwouo, N. L. & Nsang, A. (2005) An assessment of the reptiles and amphibians of the Monte Mitra forest, Monte Alen National Park. In: Sunderland, T. C. H. (Ed.), A Biodiversity Assessment of the Monte Mitra Forest, Monte Alen National Park, Equatorial Guinea. Smithsonian Institution, Washington, D. C., pp. 32 - 49.","Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato (Reptilia: Scincidae) in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863 (1), 1 - 28. https: // doi. org / 10.11646 / zootaxa. 863.1.1"]}

We recovered molecular phylogenetic relationships amongst species of the genusAblepharusin Iran and Iraq. Partial sequences of three mitochondrial genes (cytochrome C oxidase subunit I – COI, 12S rRNA and 16S rRNA) were analysed. In addition, phylogenetic relationships and taxonomic evaluation ofAblepharusspecies in Cyprus, India, Greece, Turkey and Syria were performed using partial sequences of the 16S rRNA gene. Phylogenetic trees and estimated genetic distances showed that theAblepharuspopulations of Iran and Iraq clustered into three distinct clades. One is found in northwest Iran (A. bivittatusin Ardabil, East and West Azerbaijan and Hamedan Provinces). The second clade, formed byA. chernovi, is found only in Uromia. The third and most heterogeneous clade is divided into two subclades, the first includes two lineages ofAblepharusin Khorasan Razavi and Semnan Provinces (A. pannonicus) and in eastern and south-eastern Iran (A. grayanus); the second subclade is distributed in the eastern part of Iraq and west and south-western Iran (Ablepharussp.). Our analyses indicated that splitting ofA. chernoviwithin the genus occurred in the early Miocene [about 22.5 million years ago (Mya)].Ablepharus bivittatusdiverged 15.2 Mya, in the middle Miocene.Ablepharus pannonicusdiverged in the late Miocene (8.4 Mya) andA. grayanusseparated in the late Miocene (6.7 Mya). The lineages of eastern Iraq and south-western Iran (Ablepharussp.) diverged also in the late Miocene (7.0 Mya).

Panaspis cabindae (Bocage, 1866) Material (two specimens): One adult specimen (MHNCUP/REP 266, formerly UP-MHNFCP-017381), collected from Lucálla [=Lucala] [-9.27268º, 15.25206º, 708 m a.s.l.] Kwanza Norte Province, in 1903; one adult specimen (MHNCUP/REP 267, formerly UP-MHNFCP-017380; Fig. S37), collected from Catete [-9.11667º, 13.70000º, 21 m a.s.l.] Bengo Province, in September 1903. Comments: Panaspis cabindae is a species with a distribution extending from southwestern Democratic Republic of the Congo to southwestern Angola (Marques et al. 2018; Ceríaco et al. 2020b). The presented specimens were cited as “ Ablepharus cabindae, Boc ” by Ferreira (1904) and are the only published records for the species from “Lucala” and “Catete” localities. Recently, Ceríaco et al. (2016a, 2018b, 2020b) and Medina et al. (2016) published new records for the species in Angola, including the first accounts for Bié, Luanda, Uíge, Malanje, Lunda Norte and Huíla provinces. This species is one of the most widespread in the country, despite the lack of published records (Ceríaco et al. 2020b)., Published as part of Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton's Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, pp. 1-80 in Zootaxa 5028 (1) on page 23, DOI: 10.11646/zootaxa.5028.1.1, http://zenodo.org/record/5453967, {"references":["Bocage, J. V. B. (1866) Reptiles nouveaux ou peu connus recueillis dans les possessions portugaises de l'Afrique occidentale, qui se trouvent au Museum de Lisbonne. Jornal de Sciencias Mathematicas, Physicas e Naturaes, I, 57 - 78.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501.","Ceriaco, L. M. P., Heinicke, M. P., Parker, K. L., Marques, M. P. & Bauer, A. M. (2020 b) A review of the African snake-eyed skinks (Scincidae: Panaspis) from Angola, with the description of a new species. Zootaxa, 4747 (1), 077 - 112. https: // doi. org / 10.11646 / zootaxa. 4747.1.3","Ferreira, J. B. (1904) Reptis e amphibios de Angola da regiao ao norte do Quanza (Colleccao Newton 1903). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (25), 111 - 117.","Ceriaco, L. M. P., De Sa, S. A. C., Bandeira, S., Valerio, H., Stanley, E. L., Kuhn, A. L., Marques, M. P., Vindum, J. V., Blackburn, D. C. & Bauer, A. M. (2016 a) Herpetological survey of the Iona National Park and Namibe National Park, Angola, with a synoptic list of the amphibians and reptiles of Namibe Province, southwestern Angola. Proceedings of the California Academy of Sciences, Series 4, 63 (2), 15 - 61.","Ceriaco, L. M. P., Marques, M. P., Bandeira, S., Blackburn, D. C. & Bauer, A. M. (2018 b) Herpetological survey of Cangandala National Park with a synoptic list of the amphibians and reptiles of Malanje Province, central Angola. Herpetological Review, 49 (3), 408 - 431.","Medina, M. F., Bauer, A. M., Branch, W. R., Schmitz, A., Conradie, W., Nagy, Z. T., Hibbitts, T. J., Ernst, R., Portik, D. M., Nielsen, S. V., Colston, T. J., Kusamba, C., Behangana, M., Rodel, M. - O. & Greenbaum, E. (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100, 409 - 423. https: // doi. org / 10.1016 / j. ympev. 2016.04.026"]}

Panaspis annettesabinae sp. nov. (Figs. 1���2) lsid: http://zoobank.org:XXX Holotype. TJC264 (ZMNH H2019,2176), unsexed adult, 8km SW of Bedele on the Metu-Bedele Road, Buno Bedele zone, Oromia Region, (-8.4032��, 36.3105�� elevation 1840m; datum WGS 84; Fig. 3) Ethiopia, 9 February 2013 by Timothy J.Colston. Diagnosis. Panaspis annettesabinae can be distinguished from other members of the genus by the following combination of characters: eye in the ���ablepharine��� condition; scales in 24 rows at the midbody; adult coloration of light-colored upper labials lacking round black spots; coppery dorsum flecked with black, separated from dark brown or black lateral coloration by a single row of light-colored scales; and a coppery bronze tail. Description of the holotype. Unsexed adult in good condition. Body form (size and shape) and scalation typical for Panaspis (Medina et al. 2016). Relatively slender, cylindrical body with pentadactyl fore- and hind-limbs. Trunk elongate; limbs do not overlap when adpressed. Mass 1.3g. SVL 42.1 mm, tail broken in preserved specimen; approximately 1.5 times body length in life. Head length 7.5 mm, with modestly acuminate snout (HL 139% HW). Other measurements presented in Table 1. Rostral wider than high, and visible from above. Nasals widely separated behind rostral by frontonasal. Frontonasal acuminate anteriorly, wider than long. Nostrils large, set centrally in the nasals and not bordering the postnasal. Prefrontals in contact with one another, and first supraocular, first supraciliary and frontal separated from frontonasal by supranasals (which are in contact). Two loreals, the posterior margins of the largest loreal border preoculars (3), which is wider than high. Frontal length less than the distance between anterior tip of frontal and tip of snout; frontal in contact with prefrontals anteriorly, six supraoculars (three on each side) and frontoparietals. Frontoparietals fused, in contact with the frontal, two supraoculars (one on each side), parietals and interparietal. Interparietal triangular with visible parietal window in center; parietals 2 times larger than frontoparietals and contacting each other at the anterior point of the interparietal. Parietals in contact at posterior of interparietal. Four large, broad nuchals collectively bordered by a total of nine dorsals. Supraoculars four (per side). Supraciliaries six, third widest, first tallest. Pretemporals two. Tympanum visible, approximately one third the height of the eye. Supralabials seven, V-VII being the suboculars. Ablepharine eye. Infralabials five. Postmental bordering five scales (mental, two primary chin-shields, and one infralabial on each side). Ventral scales smooth. MSR 24, SAD 62, SAV 68. Limbs with five digits; scales on soles of hands and feet smooth. Relative length of digits of manus IV=III>II>V>I relative length of digits of pes IV>III>II>I>V. Finger-IV scales 12. Tail long, robust and tapering smoothly in life, tail broken on preserved specimen. Coloration in life. Dorsum coppery brown, distinct from darker brown lateral coloration separated by a lighter (whitish tan) stripe encompassing a single scale row. Coppery dorsal coloration flecked with black, loosely arranged in 3���5 irregular lines which become less distinct posteriorly, each black fleck representing one-quarter to one-half of a dorsal scale. Darker brown lateral coloration irregularly interspersed with whitish and blackish speckles. Limbs uniformly darker chocolate brown, with occasional whitish flecking. Coppery dorsal coloration extends onto head scales, uniformly merging into tan-colored rostral. Blackish streak from nostril to orbit, dividing coppery dorsal coloration from whitish labials. Anterior tail coloration (proximo-anterior) similar to limbs, a continuation of the coppery dorsal coloration darkened to uniform chocolate brown. Posterior portion of tail (approximately the final one-third) lighter, more metallic copper coloration, possibly old regrowth or breeding plumage. Ventral coloration not recorded in life Coloration in preservative. In preservative, entire specimen with uniformly milky coloration suggestive of imminent ecdysis. Limbs and light brown, noticeably lighter than the body. Venter uniformly whitish beige, as are underside of limbs. Remnants of narrow dorsolateral stripe still visible, as are black flecks on the dorsum. Unbroken portion of tail and examination of limbs reveals clusters of melanophores across each light-colored scale. Darker blotches evident on tips of dorsal cephalic scales which are otherwise grey or light brown. Coloration of lateral surfaces similar to life, though clouded and faded due to preservation. Comparison with other Ethiopian Panaspis. As noted above, only two species are currently considered to occur in Ethiopia: P. tancredi (known from a single specimen from the northern part of the country���most likely near the region of Debarik, approximately 550km from the type locality of P. annettesabinae in markedly different habitat), and scattered records in the south-central and western regions previously assigned to P. ��� wahlbergi ��� s.l. Medina et al. (2016) restricted the nominotypical form to southeastern Africa, and revealed that there are no phylogenetically close congeners of TJC264 (P. annettesabinae), which was the sole member of a long branch in all analyses. In their analyses P. annettesabinae is the sister lineage either to most southern African species or several southeastern lineages, dating at least to the Oligocene ~33Ma. As also noted by Cer��aco et al. (2018), the extreme morphological conservatism and crypsis of these species makes comparison and identification difficult in the absence of molecular data. Thus, appropriate and robustly adequate comparisons are difficult and unclear for P. tancredi. From the only known specimen of P. tancredii (an unsexed animal of unknown reproductive status), P. annettesabinae is distinguished by a larger body size (SVL 42.1mm versus 28), 24 dorsal scale rows at midbody (versus 22), blackish flecking on the dorsum (versus lack thereof), whitish flecking on the lateral surfaces (versus lack thereof), and lack of round black spot on upper and lower labials (versus presence). Largen and Spawls (2010) included photographs of two specimens of P. ��� wahlbergi ��� s.l. from Ethiopia. Neither of these specimens were indicated to have been collected or deposited in a museum for examination. The first (Fig. 267, p. 408) is an adult or subadult from Debre Zeit (Bishoftu) in the central region, and closely resembles the holotype of P. annettesabinae. The second (Fig. 268, p. 408) is a juvenile from Bedele, near the type locality of P. anettesabinae. This specimen also resembles a juvenile version of the holotype, with darker limbs and lateral coloration and less evident dark flecking dorsally and light flecking laterally. A key difference is that the juvenile specimen has a bright blue tail in contrast to the orange tail of the holotype and the Debre Zeit specimen (the latter appears to be regenerated, while the holotype had its original tail in life), though ontogenetic color-change in skink tails (particularly in breeding males) is relatively common. Finally, a series of specimens from Negele Borena in southern Ethiopia represents another new, highly distinct species based on molecular and morphological data (TJC in prep.), suggesting that many of the remaining central and eastern records may belong to this or other new species. TABLE 1. Measurements (in g or mm) and scale counts of the holotype; abbreviations given in the Materials and Methods and definitions following Cer��aco et al. (2018). Distribution. This species is currently known only from the type locality (Fig. 3). Additional fieldwork will be needed to determine the geographic extent of the species��� range. As noted, a juvenile specimen from near the type locality (Largen and Spawls 2010) likely represents an additional record of this species, while an adult or subadult specimen from Debre Zeit may extend the range into central Ethiopia. Habitat and Natural History notes. The habitat at the type locality is a clearing within moist evergreen montane forest (elevation 1840m) in the Oromia Region of southwestern Ethiopia (Fig. 4). This clearing is likely of man-made origin, although natural ���glades��� within this forest type are known. The clearing sits alongside a stream that has been diverted for use by the Bedele Brewing company, which granted TJC access to camp on and survey the area. While P. annettesabinae is likely more common in forest openings, the type specimen was found underneath a tarp in basecamp during camp maintenance, and it should be noted that while pitfall traps were utilized in the immediate area for 21 consecutive days no other specimens of Panaspis were collected. Etymology. The specific epithet ��� annettesabinae ��� honors Annette Sabin of the Sabin family, long-time philanthropic supporters of biodiversity conservation. We propose the English vernacular name ���Sabin���s Snake-Eyed Skink.���, Published as part of Colston, Timothy J., Pyron, R. Alexander & Bauer, Aaron M., 2020, A new species of African Snake-Eyed Skink (Scincidae: Panaspis) from Ethiopia, pp. 190-200 in Zootaxa 4779 (2) on pages 192-195, DOI: 10.11646/zootaxa.4779.2.2, http://zenodo.org/record/3833467, {"references":["Medina, M. F., Bauer, A. M., Branch, W. R., Schmitz, A., Conradie, W., Nagy, Z. T., Hibbitts, T. J., Ernst, R., Portik, D. M., Nielsen, S. V., Colston, T. J., Kusamba, C., Behangana, M. & Rodel, M-O. (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100, 409 - 423. https: // doi. org / 10.1016 / j. ympev. 2016.04.026","Ceriaco, L. M., Branch, W. R. & Bauer, A. M. (2018) A new species of African snake-eyed skink (Scincidae: Panaspis) from central and northern Namibia. Zootaxa, 4527 (1), 140 - 150. https: // doi. org / 10.11646 / zootaxa. 4527.1.12","Largen, M. & Spawls, S. (2010) The amphibians and reptiles of Ethiopia and Eritrea. Edition Chimaira, Frankfurt am Main, 694 pp."]}

A recent molecular phylogenetic revision of the snake-eyed skinks (genus Panaspis Cope, 1868) uncovered extensive cryptic diversity, including several new species from throughout sub-Saharan Africa. Here, we describe one of these from Ethiopia as Panaspis annettesabinae sp. nov. This description is based on a previous molecular phylogeny and morphological, scalation, and coloration data collected from the type specimen. Phylogenetic analyses place the species alone in what we term the P. annettesabinae species group from Ethiopia. This group forms the sister lineage to a large southern African radiation and suggests a potential northern origin for much of the extant diversity of Panaspis. Many new taxa have recently been discovered in the genus and region, and there are several historical records of Panaspis from elsewhere in Ethiopia. Thus, we suggest that the range of this new species (known only from a single specimen at present) may be much larger, and that additional undescribed species may exist in northern sub-Saharan Africa.

Panaspis mocamedensis sp. nov. (Fig. 17–19) With the exception of P. cabindae, the presence of Panaspis species in Namibe province has never been noted. According to our molecular data, some Panaspis from Namibe province represent a distinct lineage of the Panaspis maculicollis species complex, sister to the recently described P. namibiana from neighboring Namibia (Ceríaco et al. 2018a). Based on the molecular and morphological evidence produced in this study, we herein describe this population as a new species, putatively endemic to the province. Holotype. FLMNH 187243 (field number AMB 10652), unsexed adult, from a river bed, 45 km SW of Virei (-16.11958°, 12.83458°, 523 m), Namibe Province, Republic of Angola, collected by Luis M. P. Ceríaco, Ishan Agarwal, Suzana Bandeira and Pedro Baptista on 3 December 2016 (Fig. 17–18). Paratypes. Four specimens, all from the Republic of Angola: CAS 263325, 263326, 263327 (field numbers AMB 10649, 10650, 10651 respectively) all unsexed adults, with the same collecting data as the holotype; CAS 264266 (field number AMB 11401) from Maungo farm (-14.538018º, 12.744793º, 365 m), Namibe Province, Republic of Angola, collected by L.M.P. Ceríaco, Mariana P. Marques, Joyce Janota and Alvaro (Varito) Baptista on 9 August 2018; Diagnosis. Panaspis mocamedensis sp. nov. can be distinguished from other members of the genus by the following combination of characteristics: 1) absence of supranasals; 2) ablepharine eye (as defined by Greer 1974); 3) frontoparietals fused; 4) dorsum coppery-brown, with a dorsolateral light stripe extending approximately to midbody, accompanied by a thinner stripe composed by black dots band starting on the temporals and extending to the base of the tail; 5) presence of rows of light spots on the neck; 6) absence of a white ventrolateral stripe; 7) 23 to 24 midbody scales rows. Description of the holotype. Unsexed subadult in good condition, but with a regenerated tail. Arrangement and relative size of head, body and tail scales typical for Panaspis. Robust, cylindrical body with well-developed pentadactyl limbs. Fore- and hind-limbs do not overlap when adpressed against the body. SVL 32.6 mm, tail regenerated. Head length 6.2 mm, with relatively acuminate snout (HL 172% HW). Other relevant measurements are presented in Table 5. Rostral wider than high, and visible from above. Nasals widely separated behind rostral by frontonasal. Frontonasal acuminate anteriorly, wider than long. Nostrils small, set posteriorly in the nasals bordering the postnasal. Prefrontals in contact with one another, almost pentagonal, contacting loreals, first supraocular, and frontal. Two loreals, the posterior margins of the largest loreal bordering the preocular, which is higher than wide. Frontal length shorter than distance between anterior tip of frontal and tip of snout; frontal in contact with prefrontals anteriorly, the first supraocular, and frontoparietals. Frontoparietals fused, in contact with the frontal, three supraoculars, parietals and interparietal. Frontoparietal plus interparietal length 2.2 times length of the frontal. Interparietal pentagonal with visible parietal foramen in anterior part; parietals about the same width of that of the frontoparietals and contacting each other at the anterior point of the interparietal. Parietals in contact. A pair of large, broad nuchals collectively bordered by a total of eight dorsals. Supraoculars three. Supraciliaries four (left side) and four (right side), third largest. Pretemporals two. Tympanum visible, approximately one seventh the height of the eye. Supralabials six, the fifth and sixth being the suboculars. Ablepharine eye. Infralabials five. Postmental bordering five scales (mental, two primary chin-shields, and one infralabial on each side). Ventral scales smooth. MSR 23, SAD 56, SAV 57. Limbs with five digits; scales on soles of hands and feet smooth. Relative length of digits of manus III>IV>II>VI relative length of digits of pes IV>III>II=V>I. Finger-IV scales 9 (right side), Toe-IV scales 15 (right side). Tail long, robust and tapering smoothly. Both in life and preservative, background color of flanks and upper side of head, neck, dorsum, legs and tail is coppery brown, with a darker dot in the anterior part of each scale. Posteriorly the tail is lighter, with disjunct dark speckles. A beige stripe, less than one scale wide, limited by the dorsal background color above and by a thin stripe of darker brown below (especially evident under the anterior part of eye), extends from the temporals and runs along the insertion of the arms. Areas below the dark brown stripe become lighter, until they reach the ventral side, which is beige. Presence of rows of light spots on the neck. Supralabials light beige anteriorly and posteriorly, with strong coppery brown markings in the middle. This pattern extends to the infralabials. Venter uniformly light-beige, as are the palmar regions of hands and feet. Variation. Variaton in scalation and body measurements of the paratypes of Panaspis mocamedensis sp. nov. is reported in Table 5. The majority of the paratypes agree entirely with the holotype, except regarding the contact between the prefrontals, which are separated in paratypes CAS 263325 and 263326. Comparison with other south and southwestern African Panaspis. As the molecular data provide evidence of the independence of the P. mocamedensis sp. nov. lineage from all other taxa, we here restrict our morphological comparisons to those named congeners occurring in Namibia and Angola. The extreme morphological conservatism of Panaspis limits diagnostic characters for most species (except P. cabindae and P. aff. breviceps) to subtle features, chiefly related to coloration pattern. Comparing P. mocamedensis sp. nov. with P. cabindae and P. aff. breviceps, the newly described species can be easily distinguished by not having supranasals (present in P. cabindae), by having an ablepharine eye (pre-ablepharine eye in P. cabindae and completely movable lower eyelids in P. aff. breviceps), and by having the frontoparietals fused (divided in P. cabindae and in P. aff. breviceps). In comparison with P. wahlbergi, P. mocamedensis sp. nov. can be distinguished by its coloration, namely by the lack of the broad black lateral stripe and bordering ventrolateral white stripe extending from the supralabials through the ear to the groin (males) or terminating behind the shoulder (females). Panaspis mocamedensis sp. nov. can be distinguished from P. maculicollis on the basis of the lack of black patch extending from ear to just beyond the shoulder and by having a smaller count of midbody scale rows (23 to 24 in P. mocamedensis sp. nov. versus typically 26 in P. maculicollis). P. mocamedensis sp. nov. can be distinguished from the newly described P. namibiana on the basis of the presence two or three vertical or diagonal rows of white spots on the neck and by having a smaller count of midbody scale rows (23 to 24 in P. mocamedensis sp. nov. versus usually more than 24 in P. namibiana). Distribution. The newly described species is currently only known from central Namibe Province (Fig. 19). It is likely that the species occurs in other regions of the province where appropriate habitat is available, but further sampling is needed to confirm its presence. Habitat and Natural History notes. The species is terrestrial and is typically found under leaf litter or other debris. It is only known from habitats represented by mosaics of steppes and savannah dominated by Mopane trees and sandy soils (Fig. 20; Grandvaux-Barbosa 1973). Specimens from Virei were captured by day, but CAS 264266 was active under debris well after sunset. Etymology. The specific epithet “ mocamedensis ” refers to the former name of Namibe Province (Moçâmedes), to which the species is apparently endemic. We propose the English vernacular name “Moçamedes Snake-Eyed Skink” and the Portuguese vernacular name of “Lagartixa da Manta-Morta de Moçâmedes”.

Panaspis wahlbergi (Smith, 1849) (Fig. 7, 13–15) Ablepharus Wahlbergii (Bocage 1895: 52) Ablepharus wahlbergii (Boulenger 1905: 111; Loveridge 1957: 219) Afroablepharus wahlbergi (FitzSimons 1937: 269; Greer 1974: 32; Perret 1975: 8; Schmitz et al. 2005: 19; Bates et al. 2014: 257) Panaspis wahlbergi (Branch 1998: 159; Jacobsen & Broadley 2000: 63; Branch & Conradie 2015: 200; Medina et al. 2016: 410; Branch et al. 2019a: 296) Panaspis aff. wahlbergi (Marques et al. 2018: 249; Branch et al. 2019a: 318) Panaspis sp. (Baptista et al. 2018: 400) Panaspis aff. namibiana (Butler et al. 2019: 233) P. wahlbergi was originally described as Cryptoblepharus wahlbergi by Smith (1849) based on specimens from the “Country to the eastward of the Cape Colony,” later restricted to KwaZulu-Natal Province, South Africa, by FitzSimons (1937). Similar to all other members of the genus, the species has a complex taxonomic history, especially regarding its genetic allocation (for a review of its taxonomic and nomenclatural history see Schmitz et al. 2005 and Medina et al. 2016). The first records of this species for Angola were provided by Bocage (1895), who noted specimens collected by J.A.O. de Anchieta from Cahata and Caconda, in the higher elevation areas of Benguela and Huíla provinces, respectively. As he had done with P. cabindae, Bocage (1895) provided a detailed illustration of Angolan P. wahlbergi (Fig. 7). Boulenger (1905) referred to this species specimens collected “between Benguella and Bihé” [between what it is currently understood as Benguela and Bié provinces] by the British explorer William John Ansorge (1850–1913). These are the only published records of the species for Angola, although some other specimens assignable to the species were collected in the country during the twentieth century and deposited in the collections of Portuguese (IICT), North American (CAS, FMNH) and South African (TM) museums. More recently the species has been relocated in the country. Baptista et al. (2018) reported the presence of Panaspis sp. in theTundavala area. Butler et al. (2019) reported one specimen from Bicuar National Park, which they mistakenly labeled as P. aff. namibiana. Other specimens were recently collected by the authors LMPC and MPM in southeastern Bié Province. Both Marques et al. (2018) and Branch et al. (2019) referred to the central Angolan populations of P. wahlbergi as P. aff. wahlbergi, suggesting doubts about its conspecificity with the nominotypical form, and even suggesting that the Angolan population may represent a different taxon (Branch et al. 2019a). However, contrary to what Medina et al. (2016) demonstrated for the eastern African populations, where a high level of cryptic diversity exists, and to the suspicions of Marques et al. (2018) and Branch et al. (2019), our results unambiguously place the central Angolan population as conspecific with nominotypic wahlbergi. The presence in the country of additional undescribed wahlbergi -like lineages cannot be excluded, as Bittencourt-Silva (2019) recently reported the presence of one of these lineages (“ Panaspis sp. Katanga 2” sensu Medina et al. 2016) in northwestern Zambia, not far from the Angolan border. Specimens from Calombe-Luso, Moxico Province, (IICT 268–1959 and 276–1959) could potentially belong to this lineage as well. Further sampling in the eastern regions of the country is needed to address this possibility. Diagnosis. Panaspis wahlbergi can be distinguished from other members of the genus occurring in Angola by the following combination of characteristics: 1) absence of supranasals; 2) ablepharine eye (as defined by Greer 1974); 3) frontoparietals fused; 4) dorsum coppery-brown, with a broad black lateral band; 5) absence of rows of light spots on the neck; 6) usually a presence of a white ventrolateral stripe, from supralabials to the groin; 7) 24 to 26 midbody scales rows (Figs. 13–14). Specimens examined. ANGOLA: Benguela Province: Cubal [-13.03333º, 14.25, 922 m] ( TM 39762); Kwanza-Sul Province: 10 miles SE of Quirimbo [-10.83998º, 14.308124º, 851 m] (CAS 85966); Bié Province: near Mumbué [-13.85417º, 17.47044º, 1631 m] (AMB 11675 –11676); Chitau [-11.43333º, 17.15º, 1510 m] ( FMNH 74295, AMNH 48505); Huíla Province: Caconda [-13.73333º, 15.06667º, 1675 m] ( NHM 1906.8.24.72); 19 miles NE of Lubango [-14.78503º, 13.730408º, 1691 m] (CAS 85956); Lubango [-14.916667º, 13.5º, 1760 m] (FMNH 74295); Bicuar National Park [-15.1034º, 14.8311º, 1243 m] (CAS 263401). Distribution. Medina et al. (2016) restricted to Mozambique and northeastern South Africa, but we here extend that distribution into south-central Angola. Historical records from Zimbabwe and Zambia might also represent true wahlbergi, potentially connecting the Angolan population to those of Mozambique and South Africa. However, this distribution requires further assessment, as other cryptic lineages may also be present, as evidenced by the recent discovery in northwestern Zambia of one of the Katangan cryptic lineages of wahlbergi by Bittencourt-Silva (2019). In Angola, P. wahlbergi is limited to the central and southern areas of the Angolan plateau, from Benguela Province to Bié (Fig. 15), but is most likely that the species also occurs in the neighboring areas of Moxico and Cuando-Cubango provinces. Habitat and Natural History notes. The species is found in Miombo Woodlands, usually under leaf-litter, under rocks, or moving through grass (Fig. 16)., Published as part of Ceríaco, Luis M. P., Heinicke, Matthew P., Parker, Kelly L., Marques, Mariana P. & Bauer, Aaron M., 2020, A review of the African snake-eyed skinks (Scincidae: Panaspis) from Angola, with the description of a new species, pp. 77-112 in Zootaxa 4747 (1) on pages 98-101, DOI: 10.11646/zootaxa.4747.1.3, http://zenodo.org/record/3693431, {"references":["Smith, A. (1849) Appendix. In: Smith, A. (Ed.), Illustrations of the zoology of South Africa, Reptilia, 1849, pp. 1 - 28 + [i].","Bocage, J. V. B. (1895) Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, 203 pp., 20 pls.","Boulenger, G. A. (1905) A list of the batrachians and reptiles collected by Dr. W. J. Ansorge in Angola with descriptions of new species. Annals and Magazine of Natural History, Series 7, 16, 105 - 115, pl. 4. https: // doi. org / 10.1080 / 03745480509443656","FitzSimons, V. F. M. (1937) Notes on the reptiles and amphibians collected and described from South Africa by Andrew Smith. Annals of the Transvaal Museum, 17, 259 - 274.","Greer, A. E. (1974) The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, 31, 1 - 67. https: // doi. org / 10.1071 / ajzs 031","Perret, J. L. (1975) La differentiation dans le genre Panaspis Cope (Reptilia: Scincidae). Bulletin de la Societe Neuchateloise des Sciences Naturelles, 98, 5 - 16.","Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato (Reptilia: Scincidae) in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863 (1), 1 - 28. https: // doi. org / 10.11646 / zootaxa. 863.1.1","Bates, M. F., Branch, W. R., Bauer, A. M., Burguer, M., Marais, J., Alexander, G. J. & De Villiers, M. S. (2014) Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. Suricata 1. South African National Biodiversity Institute, Pretoria, xvii + 459 pp.","Branch, W. R. (1998) Field Guide to Snakes and other Reptiles of Southern Africa. 3 rd Edition. Struik Publishers, Cape Town, 399 pp.","Jacobsen, N. H. G. & Broadley, D. G. (2000) A new species of Panaspis Cope (Reptilia: Scincidae) from southern Africa. African Journal of Herpetology, 49 (1), 61 - 71. https: // doi. org / 10.1080 / 21564574.2000.9650017","Branch, W. R. & Conradie, W. (2015) IV. Herpetofauna da regiao da Lagoa Carumbo. In: Huntley, B. & Francisco, P. (Eds.), Avaliacao Rapida da Biodiversidade da Regiao da Lagoa Carumbo, Lunda Norte-Angola. Ministerio do Ambiente de Angola, Luanda, Angola, pp. 194 - 209.","Medina, M. F., Bauer, A. M., Branch, W. R., Schmitz, A., Conradie, W., Nagy, Z. T., Hibbitts, T. J., Ernst, R., Portik, D. M., Nielsen, S. V., Colston, T. J., Kusamba, C., Behangana, M. & Rodel, M. - O. (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100, 409 - 423. https: // doi. org / 10.1016 / j. ympev. 2016.04.026","Branch, W. R., Pinto. P. V., Baptista, N. & Conradie, W. (2019 a) The Reptiles of Angola: History, Diversity, Endemism and Hotspots. In: Huntley, B, Russo, V., Lages & Ferrand, N. (Eds.) Biodiversity of Angola, Science & Conservation: A Modern Synthesis. Springer, Cham, pp. 283 - 334. https: // doi. org / 10.1007 / 978 - 3 - 030 - 03083 - 4 _ 13","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola-Atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (supplement 2), 1 - 501.","Baptista, N., Antonio, T. & Branch, W. R. (2018) Amphibians and reptiles of the Tundavala region of the Angolan Escarpment. Biodiversity & Ecology, 6, 397 - 403. https: // doi. org / 10.7809 / b-e. 00351","Butler, B. O., Ceriaco, L. M., Marques, M. P., Bandeira, S., Julio, T., Heinicke, M. P. & Bauer, A. M. (2019) Herpetological Survey of Huila Province, Southwest Angola, Including First Records from Bicuar National Park. Herpetological Review, 50, 225 - 240.","Bittencourt-Silva, G. B. (2019) A herpetological survey of western Zambia. Amphibian and Reptile Conservation, 13 (2), 1 - 28."]}

Panaspis maculicollis Jacobsen & Broadley, 2000 Panaspis maculicollis (Conradie et al. 2016: 25; Medina et al. 2016: 414; Ceríaco et al. 2018a: 141; Marques et al. 2018: 248; Branch et al. 2019a: 293, 317) This species was described by Jacobsen & Broadley (2000), based on specimens from northern Limpopo and Mpumalanga provinces of South Africa, northern Botswana, northeastern Namibia, Zimbabwe, southern Zambia and central Mozambique. Medina et al. (2016) presented evidence that the central Namibian populations, as well as those from Mozambique and Limpopo represent undescribed species. The central Namibian population was recent- ly described by Ceríaco et al. (2018), while the other lineages await description. Conradie et al. (2016) presented the first record of the species for Angola, based on a single specimen collected in the southeastern tip of Cuando- Cubango Province. The taxonomic identity of this specimen was confirmed by Medina et al. (2016). Diagnosis. Panaspis maculicollis can be distinguished from other members of the genus occurring in Angola by the following combination of characteristics: 1) absence of supranasals; 2) ablepharine eye (as defined by Greer 1974); 3) frontoparietals fused; 4) dorsum coppery-brown, with a dorsolateral light stripe extending approximately to midbody, and a thin darker band starting on the temporals and extending to midbody; 5) presence of rows of light spots on the neck; 6) absence of a white ventrolateral stripe; 7) 22 to 28 midbody scales rows (usually 26). Additional material. ANGOLA: Cuando-Cubango Province: Cuando Basin [-17.535º, 23.18916º, 983 m] (PEM R 20007). Distribution. The species is known from the Kalahari sand areas of northeastern Namibia, northern Botswana and western Zimbabwe. In Angola, the species enters the extreme southeastern tip of Cuando-Cubango Province, however it is probable that it is more widespread in the province, and similar habitat extends into Moxico Province (Fig. 12). Habitat and Natural History notes. A terrestrial species occupying a variety of habitats from rocky or stony hillsides to open woodland/grassland at altitudes of 220-1200 m (Jacobsen & Broadley 2000). In Angola the species is known from the sparse Miombo woodlands from southeastern Moxico.

The genus Panaspis in Angola is represented by four species, most of them part of taxonomically and nomenclaturally challenging species-complexes. We present a taxonomic revision of the group in the region and describe one new species, Panaspis mocamedensis sp. nov., endemic to the lowland areas of the Namibe province, southwestern Angola. Phylogenetic analysis using a combination of mitochondrial (16S, cytb) and nucleares (RAG1, PDC) markers, as well as morphological and meristic data support the recognition of the new species. In addition, these data support the presence of nominotypical Panaspis cabindae, P. wahlbergi and P. maculicollis in Angola. Reexamination of the Angolan population of P. breviceps was based on morphological analysis, as no molecular data from Angola is available for this species. According to our results, this population likely represents the nominotypical form, but due to its complex taxonomic and nomenclatural history and the lack of molecular data, this population needs to be reconsidered when molecular data become available. The description of a new species and revision of the Angolan Panaspis contributes to a better understanding of the true species richness of the Angolan herpetofauna, as well as to understanding the major biogeographic patterns of the region. A key to Angolan Panaspis species is also presented.

Panaspis cabindae (Bocage, 1866) (Fig. 7–10) Chresonymy (includes only citations unambiguously pertaining to specimens referable to the species, or explicitly stating or depicting its geographic range). Ablepharus cabindae (Bocage 1866a: 64, 1895: 51; Peters 1877: 614; Boulenger 1887: 352; Ferreira 1904: 116; Parker 1936: 136) Ablepharus Cabindae (Bocage 1866b: 45, 1867: 224, 1887: 179, 1897: 196) Ablepharus aeneus (Boulenger 1887: 352) Riopa (Panaspis) cabindae (Smith 1937: 229) Panaspis cabindae (Greer 1974: 29; Perret 1975: 8; Schmitz et al. 2005: 23; Ceríaco et al. 2016a: 57; 2016b: 65; Medina et al. 2016: 411; Ceríaco et al. 2018a: 141; Ceríaco et al. 2018c: 423; Marques et al. 2018: 247; Branch et al. 2019a: 317) This species was described by Bocage (1866) based on three specimens collected by the Portuguese explorer José Alberto de Oliveira de Anchieta (1832–1897) in the Cabinda enclave, northwestern Angola. Cope (1868) described a new genus and species Panaspis aeneus. The description of this species was problematic as Cope was unable to provide a specific type locality (according to the original description Habitat. Probably Swan River, Australia; possibly from South-Western Africa.), and the holotype is currently lost. The usage of the nomen aeneus was short-lived, as the only other reference to aeneus as a valid species was that of Boulenger (1887), when referring to specimens from Angola. The two specimens labelled as aeneus by Boulenger (1887) were collected by the London-born Portuguese colonial officer Joaquim John Monteiro (1833–1878) in the Luanda region. These specimens are still extant in the collections of the BMNH under the accession number 1873.7.28.11 and are unambiguously conspecific with P. cabindae. In his scholarly revision of the herpetofauna of Angola, Bocage (1895) was the first to synonymize aeneus with cabindae, a decision then followed by all subsequent authors (e.g. Fuhn 1972; Perret 1975). In the same work Bocage (1895) provided the first illustration of the species, including detailed views of the head scalation (Fig. 7). Despite the unavailability of aeneus type material, the original description provides enough morphological data to confidently confirm that it is conspecific with cabindae, namely the presence of supranasals, pre-ablepharine eye and frontoparietals separated, diagnostic features of cabindae (see below). Due to this lack of ambiguity, a neotype designation for aeneus is not needed. The same holds true for cabindae itself, as the original type series was destroyed in the fire that engulfed the zoological collections of Museu Bocage, Lisbon, in 1978. The most recent data on the species was provided by Medina et al. (2016) who confirmed, through molecular data, that the species extended from southwestern Democratic Republic of the Congo to southwestern Angola; our sequence data here further fills in the Angolan distribution of the species. Other recent published records include those from Cangandala National Park, Malanje Province, Central Angola reported by Ceríaco et al. (2016b) and Ceríaco et al. (2018c). Marques et al. (2018) presented and mapped all available records of the species for Angola. Despite being a quite common species in Angola (LMPC pers. obs.), not much has been published about the distriution, natural history, ecology or conservation status of this species. Diagnosis. Panaspis cabindae can be distinguished from other members of the genus occurring inAngola and surroundingsregionbyhaving1)presenceofsupranasals;2)pre-ablepharineeye(asdefinedbyGreer1974);3)frontoparietalsseparated;dorsumcoppery-brown,withfourveryvaguelongitudinalstripesextendingapproximatelytomidbody, and a thin darker band starting on the temporals and extending to midbody; 5) absence of rows of light spots on the neck; 6) absence of a white ventrolateral stripe; 7) 23 to 26 midbody scales rows (Figs. 8–9). Specimens examined. ANGOLA: Cabinda Province: Chinchoxo [-5.1º, 12.1º, 45 m] (ZMB 9168, 9276); Zaire Province: Soyo [-6.134903º, 12.368935º, 10 m] (PEM 20256 –20260); Uíge Province: Kimpa Vita University Campus [-7.618614º, 15.065536º, 1122 m] (MTD 48612); Luanda Province: Luanda [-8.83333º, 13.26667º, 81 m] (NHM 1951.1.1.94), Cabiri [-8.91667º, 13.66667º, 30 m] ( AMB 10881, 10883), Kissama National Park [-9.18303°, 13.37063°, 136 m] (FLMNH 187235–241); Bengo Province: Catete [-9.11667º, 13.7º, 21 m] (MHN- CUP 017380), Riverine Forest [-9.11667º, 13.7º, 21 m] PEM R 21594; Benguela Province: Lobito Bay [-12.35º, 13.55º, 7 m] (AMNH R-48501), Hanha [-13.3º, 14.2º, 953 m] (AMNH R-40671, 48502, 48504, 40718); Malanje Province: Duque de Bragança / Kalandula [-9.1º, 15.95º, 1011 m] (ZMB 7775, CAS 263590), Cangandala National Park [-9.84606º, 16.72233º, 1111 m] (CAS 258403–407); Laúca [-9.762749º, 15.143798º, 750 m] (ANG 40-41, AMB 9190, 9209, 9214, 9230, 9250, 9251, 9254, 9255, 9256, 9257, 9258, 9259, 9260, 9261, 9262, 9676, 9677, 9678, 9685, 9686); Lunda Norte Province: Lagoa Carumbo [-7.74422º, 19.95467º, 789 m] (PEM R 19467); Kwanza-Norte Province: Lucalla [-9.4º, 15.03333º, 411 m] (MHNCUP 017381); N’Dallatando [-9.3º, 14.91667º, 782 m] (NHM 1909.10.29.103); Kwanza-Sul Province: Congulu [-10.86667º, 14.28333º, 639 m] (NHM 1936.8.1.622–624, MCZ R- 112222); Bié Province: Cassumbe [-11.08615º, 16.66619º, 1235 m] (AMB 11696, 11717, 11753, 11788); Chitau [-11.43333º, 17.15º, 1510 m] (FMNH 74295); Namibe Provice: Mamué Riparian área [-13.80147º, 13.12056º, 660 m] (CAS 10315, AMB 10316, CAS 263554, 10321, CAS 263550); Capangombe [-15.1º, 13.15º, 553 m] (MHNCUP 017379, NHMW 10236, CAS 264244). REPUBLIC OF THE CONGO: Pointe-Noire [-4.762602º, 11.866835º, 33 m] (MNHN 1966.554). DEMOCRATIC REPUBLIC OF THE CONGO: Bombo-Lumene Reserve Headquarters [-4.42218º, 16.04904º, 619 m] (UTEP Herp-21173); Malela [-5.982438º, 12.621458º, 1 m] (CAS 54820 –821; MCZ R-11286, 14011–14012). Distribution. The species occurs in the southwestern region of Republic of the Congo and the Democratic Republic of the Congo, extending to all parts of northern Angolan and descending south to central and southwestern Angola through the Angolan escarpment (Fig. 10). Habitat and Natural History notes. The species is found in a wide variety of habitats, from some more coastal, Euphorbia and baobab dominated savannahs, in northwestern Angola, to typical Miombo woodland habitats (Fig. 11) in the central plateau of Angola. The species appears to be absent from the more xeric and desertic areas of the southwest of Angola and northwestern Namibia, however it may possibly be found in more humid areas close to the escarpment. The species is active during the day in leaf-litter, but is rarely seen exposed., Published as part of Ceríaco, Luis M. P., Heinicke, Matthew P., Parker, Kelly L., Marques, Mariana P. & Bauer, Aaron M., 2020, A review of the African snake-eyed skinks (Scincidae: Panaspis) from Angola, with the description of a new species, pp. 77-112 in Zootaxa 4747 (1) on pages 90-93, DOI: 10.11646/zootaxa.4747.1.3, http://zenodo.org/record/3693431, {"references":["Bocage, J. V. B. (1866 a) Lista dos reptis das possessoes portuguezas d'Africa ocidental que existem no Museu de Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes, 1, 37 - 56.","Bocage, J. V. B. (1895) Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, 203 pp., 20 pls.","Peters, W. C. H. (1877) Ubersicht der Amphibien aus Chinchoxo (Westafrika), welche von der Afrikanischen Gesellschaft dem Berliner zoologischen Museum ubergeben sind. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1877, 311 - 621, 1 pl.","Boulenger, G. A. (1887) Catalogue of the Lizards in the British Musuem (Natural History). Vol. 3. Lacertid ӕ, Gerrhosaurid ӕ, Scincid ӕ, Anelytropsid ӕ, Dibamid ӕ, Chamaeleontid ӕ. Trustees of the British Museum, London, xii + 575 pp., 40 pls.","Ferreira, J. B. (1904) Reptis e amphibios de Angola da regiao ao norte do Quanza (Colleccao Newton- 1903). Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segunda Serie, 7, 111 - 117.","Parker, H. W. (1936) Dr. Karl Jordan's expedition to South-West Africa and Angola: herpetological collection. Novitates Zoologicae, 40, 115 - 146.","Bocage, J. V. B. (1866 b) Reptiles nouveaux ou peu connus recueillis dans les possessions portugaises de l'Afrique occidentale, qui se trouvent au Museum de Lisbonne. Jornal de Sciences Mathematicas, Physicas e Naturaes, 1, 57 - 78.","Bocage, J. V. B. (1867) Segunda lista dos reptis das possessoes portuguezas d'Africa occidental que existem no Museu de Lisboa. Jornal de Sciences Mathematicas, Physicas e Naturaes, 1, 217 - 228.","Bocage, J. V. B. (1887) Melanges erpetologiques. I. Reptiles et batraciens du Congo. Jornal de Sciences Mathematicas, Physicas e Naturaes, 11, 177 - 192.","Bocage, J. V. B. (1897) Mammiferos, reptis e batrachios d'Africa de que existem exemplares typicos no Museu de Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segunda Serie, 4, 187 - 206.","Smith, M. A. (1937) A review of the genus Lygosoma (Scincidae: Reptilia) and its allies. Records of the Indian Museum, 39 (3), 213 - 234.","Greer, A. E. (1974) The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, 31, 1 - 67. https: // doi. org / 10.1071 / ajzs 031","Perret, J. L. (1975) La differentiation dans le genre Panaspis Cope (Reptilia: Scincidae). Bulletin de la Societe Neuchateloise des Sciences Naturelles, 98, 5 - 16.","Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato (Reptilia: Scincidae) in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863 (1), 1 - 28. https: // doi. org / 10.11646 / zootaxa. 863.1.1","Ceriaco, L. M. P., de Sa, S. A. C., Bandeira, S., Valerio, H., Stanley, E. L., Kuhn, A. L., Marques, M. P., Vindum, J. V., Blackburn, D. C. & Bauer, A. M. (2016 a) Herpetological survey of Iona National Park and Namibe Regional Natural Park, with a synoptic list of the amphibians and reptiles of Namibe Province, southwestern Angola. Proceedings of the California Academy of Sciences, Series 4, 63, 15 - 61.","Ceriaco, L. M. P., Marques, M. P. & Bandeira, S. A. (2016 b) Anfibios e Repteis do Parque Nacional da Canganda. Instituto Nacional da Biodiversidade e Areas de Conservacao, Angola & Museu Nacional de Historia Natural e da Ciencia, Lisboa, 97 pp.","Medina, M. F., Bauer, A. M., Branch, W. R., Schmitz, A., Conradie, W., Nagy, Z. T., Hibbitts, T. J., Ernst, R., Portik, D. M., Nielsen, S. V., Colston, T. J., Kusamba, C., Behangana, M. & Rodel, M. - O. (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100, 409 - 423. https: // doi. org / 10.1016 / j. ympev. 2016.04.026","Ceriaco, L. M. P., Branch, W. R. & Bauer, A. M. (2018 a) A new species of African snake-eyed skink (Scincidae: Panaspis) from central and northern Namibia. Zootaxa, 4527 (1), 140 - 150. https: // doi. org / 10.11646 / zootaxa. 4527.1.12","Ceriaco, L. M. P., Marques, M. P., Bandeira, S., Blackburn, D. C. & Bauer, A. M. (2018 c) Herpetological survey of Cangandala National Park with a synoptic list of the amphibians of Malanje Province, central Angola. Herpetological Review, 49 (3), 408 - 431.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola-Atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (supplement 2), 1 - 501.","Branch, W. R., Pinto. P. V., Baptista, N. & Conradie, W. (2019 a) The Reptiles of Angola: History, Diversity, Endemism and Hotspots. In: Huntley, B, Russo, V., Lages & Ferrand, N. (Eds.) Biodiversity of Angola, Science & Conservation: A Modern Synthesis. Springer, Cham, pp. 283 - 334. https: // doi. org / 10.1007 / 978 - 3 - 030 - 03083 - 4 _ 13","Jacobsen, N. H. G. & Broadley, D. G. (2000) A new species of Panaspis Cope (Reptilia: Scincidae) from southern Africa. African Journal of Herpetology, 49 (1), 61 - 71. https: // doi. org / 10.1080 / 21564574.2000.9650017","Cope, E. D. (1868) Observations on reptiles of the old world. Proceedings of the Academy of Natural Sciences of Philadelphia, 20 (1868), 316 - 323.","Fuhn, I. E. (1972) Revision du phylum forestier du genre Panaspis Cope (Reptilia, Scincidae, Lygosominae). Revue Roumaine de Biologie, serie de Zoologie, 17 (4), 257 - 272."]}

Historically Panaspis wahlbergi (Smith, 1849) has been the only assignable species present in Kenya. Recent studies have shown that it comprises multiple cryptic species and the nominal species is now restricted to southern Africa. Newly collected mitochondrial data (16S rRNA) helped to resolve the status of the Kenyan populations, which revealed the presence of two distant related species. Pairwise distances show average 5.87% differences between the two Kenyan species, and 3.58–5.27% and 8.62–9.15% to nominal P. wahlbergi and P. maculicollis Jacobsen & Broadley, 2000 respectively. Ablepharus massaiensis Angel, 1924 was described from the Maasai plains near Nairobi, but has long been considered a junior synonym of P. wahlbergi. We herein resurrect Panaspis massaiensis comb. nov. as a valid species and describe a new species, Panaspis tsavoensis sp. nov. from the Tsavo Conservation Area in south-eastern Kenya. Morphological examinations of specimens reveal minor differences from each other as well as nominal forms of P. wahlbergi and P. maculicollis. Panaspis massaiensis comb. nov. shares with the P. wahlbergi group a white ventrolateral stripe but can be distinguished by the presence of 26 midbody scale rows versus 24. Panaspis tsavoensis sp. nov. on the other hand, lacks the white ventrolateral stripe, most similar to the P. maculicollis group but differs in that P. maculicollis breeding males have a black patch on the neck with diagonal rows of white spots. Panaspis massaiensis comb. nov. is widespread in the Kenyan and northern Tanzanian highlands, isolated dryland montane forests and rocky hills, while P. tsavoensis sp. nov. occur in the expansive arid lowlands of Tsavo Conservation Area and should be present in similar arid lowlands in northern Kenya as well as in adjacent Tanzania.

Panaspis namibiana Cer��aco, Branch & Bauer sp. nov. (Fig. 1���2) Chresonymy (includes only citations unambiguously pertaining to specimens referable to the new species, or explicitly stating or depicting its geographic range). Ablepharus wahlbergi (Lawrence 1929:26) Ablepharus wahlbergii (FitzSimons 1943:236; Mertens 1955:74, pl. 23, fig. 143; Mertens 1971:64) Panaspis (Afroablepharus) wahlbergi (Visser 1984:59) Panaspis wahlbergii (Branch 1988:134; Bauer et al. 1993:134; Branch 1994:134; Branch 1998:159). Panaspis sp. nov. (Griffin 2003:60). Afroblepharus wahlbergi (Herrmann & Branch 2013:29). Afroablepharus sp. Namibia (Medina et al. 2016:411). Holotype. CAS 206967, unsexed subadult, adjacent to Opuwo Road (D3704), 21 km N of Palmweg / Sesfontein Road (D3706) (18.9628333�� S, 13.758666�� E; WGS-84; Fig. 3), Kunene Region, Republic of Namibia, collected by Aaron M. Bauer, A. C. Lamb, J.L. Wright, P. Moler, W.R. Branch and R.D. Babb on 25 July 1998 (Fig. 1). Paratypes. Ten specimens, all from the Republic of Namibia: MCZ R183767, unsexed subadult, Sesfontein, Paracamp, Kunene Region (19.13222222�� S, 13.5877777�� E; WGS-84), collected by Aaron M. Bauer and Anthony P. Russell on 11 December 2002; MCZ R 190318, unsexed subadult, Kamanjab Rest Camp, Kunene Region (19.629611�� S, 14.682417�� E; WGS-84) collected by Aaron M. Bauer on 29 November 2011; CM 130371 (Fig. 2), unsexed adult, 80 air km E Sesfontein, 60 Air km WNW Kamanjab, Kunene Region (19.332654�� S, 14.369919�� E; WGS-84), collected by Paul Freed et al. on 3 April 1992 (Fig. 4); LACM 77840, unsexed adult, 40 km WNW Grootfontein, Otjozondjupa Region (19.48333�� S, 17.73333�� E; WGS-84), collected by R.L. Bezy on 20 November 1973; LACM 77841, 77842, unsexed adults, same collecting data as LACM 77840; PEM R2123, unsexed adult, Kaoko Otavi, Kunene Region (18.3�� S, 13.7�� E; WGS-84), collected by G. Mclachlan and J. Spence on 14 October 1965; PEM 2124, unsexed subadult, same collecting data as PEM R2123; PEM R10262, unsexed adult, Okahandja, Otjozondjupa Region (21.983333�� S, 16.916667�� E; WGS-84), collected by R. Bradfield on 23 April 1929; PEM R10273, unsexed adult, same collecting data as PEM R10262. Additional material. SMF 46570, Farm Beula, Kunene Region (19.6608�� S, 14.91185 E; WGS-84); SMF 50620, Okahandja, Otjozondjupa Region, (21.983333�� S, 16.916667��E; WGS-84); SMF 66144, Warmquelle near Zesfontein, Kunene Region (19.182�� S, 13.81384�� E; WGS-84); SMF 67050 -02, 60 mi. east of Okahandja, Otjozondjupa Region (21.623�� S, 17.64941�� E; WGS-84); SAM 0 17483, Kaoko Otavi, Kunene Region (18.3��S, 13.7��E; WGS-84); SAM 0 17557, Okorosave, Kunene Region (18.17�� S, 13.8�� E; WGS-84); SAM 0 46548, Tsumeb Townlands (19.23�� S, 17.72�� E; WGS-84), Oshikoto Region; PEM R8845, 2 km south of Grootfontein, Otjozondjupa Region (19.3206�� S, 18.39306�� E, WGS-84); TM 85249, Farm Uisib 422, Otjozondjupa Region (19.5531�� S, 17.21667�� E, WGS-84); SMW 1946, Namutoni, Oshikoto Region (18.80776�� S, 16.940231�� E, WGS-84), SMW 1947, Kaoko Otavi, Kunene Region (18.3�� S, 13.7��E; WGS-84); SMW 1948, Klein Windhoek, Khomas Region (22.578�� S, 17.10083�� E, WGS-84); SMW 1949, Windhoek, Khomas Region (22.5613�� S, 17.07214�� E; WGS-84); SMW 5700, Barab on Palmwag 702, Kunene Region (19.8214�� S, 13.7614�� E; WGS-84); SMW 6471, Warmquelle, Kunene Region (19.182�� S, 13.81384�� E; WGS-84). Diagnosis. Panaspis namibiana sp. nov. can be distinguished from other members of the genus by the following combination of characteristics: 1) absence of supranasals; 2) ablepharine eye (as defined by Greer 1974); 3) frontoparietals fused; 4) dorsum coppery-brown, with a dorsolateral light stripe extending approximately to midbody, and a thin darker band starting on the temporals and extending to midbody; 5) absence of rows of light spots on the neck; 6) absence of a white ventrolateral stripe. Description of the holotype. Unsexed subadult in good condition. Arrangement and relative size of head, body and tail scales typical for Panaspis. Robust, cylindrical body with well-developed pentadactyl limbs. Fore- and hind-limbs do not overlap when adpressed against the body. SVL 29.6 mm, tail length 38.6 mm. Head length 5.9 mm, with relatively acuminate snout (HL 184% HW). Other relevant measurements are presented in Table 1. Rostral wider than high, and visible from above. Nasals widely separated behind rostral by frontonasal. Frontonasal acuminate anteriorly, wider than long. Nostrils small, set posteriorly in the nasals but not bordering the postnasal. Prefrontals in contact with one another (atypical considering the type series), hexagonal, contacting frontonasal, loreals, first supraocular, first supraciliary and frontal. Two loreals, the posterior margins of the largest loreal border preocular, which is wider than high. Frontal length shorter than distance between anterior tip of frontal and tip of snout; frontal in contact with prefrontals anteriorly, two supraoculars (one on each side) and frontoparietals. Frontoparietals fused, in contact with the frontal, three supraoculars, parietals and interparietal. Frontoparietal plus interparietal length 2.2 times length of the frontal. Interparietal quadrangular with visible parietal foramen in anterior part; parietals 1.2 times larger than frontoparietals and contacting each other at the anterior point of the interparietal. Parietals in contact. A pair of large, broad nuchals collectively bordered by a total of nine dorsals. Supraoculars three. Supraciliaries three (left side) and four (right side), third largest. Pretemporals two. Tympanum visible, approximately one fifth the height of the eye. Supralabials six, the fifth and sixth being the suboculars. Ablepharine eye. Infralabials five. Postmental bordering five scales (mental, two primary chin-shields, and one infralabial on each side). Ventral scales smooth. MSR 22, SAD 58, SAV 55. Limbs with five digits; scales on soles of hands and feet smooth. Relative length of digits of manus IV=III>II=V>I relative length of digits of pes IV>III>II>I>V. Finger-IV scales 9 (right side), Toe-IV scales 13 (right side). Tail long, robust and tapering smoothly. In preservative, background color of flanks and upper side of head, neck, dorsum, legs and tail is coppery brown, with a darker dot in the anterior part of each scale. Posteriorly the tail is lighter, with disjunct dark speckles. A beige stripe, less than one scale wide, limited by the dorsal background color above and by a thin stripe of darker brown below (especially evident under the anterior part of eye), extends from the temporals and runs along the dorsolateral surface to the midbody. Areas below the dark brown stripe become lighter, until they reach the ventral side, which is beige. Supralabials light beige anteriorly and posteriorly, with strong coppery brown markings in the middle. This pattern extends to the infralabials. Venter uniformly light-beige, as are the palmar regions of hands and feet. Variation. Variaton in scalation and body measurements of the paratypes of Panaspis namibiana sp. nov. is reported in Table 1. The majority of the paratypes agree entirely with the holotype, except regarding the contact between the prefrontals, which are separated in paratypes CM 130371, LACM 77841, LACM 77842, PEM R10273, and narrowly separated in LACM 77840, PEM R2124 and PEM R 10262. Comparison with other South and Southwestern African Panaspis. As the molecular framework of Medina et al. (2016) provided evidence of the independence of the P. namibiana lineage from all other taxa, we here restrict our morphological comparisons to those named congeners occurring in Namibia, Angola, Botswana and South Africa. The extreme morphological conservatism of Panaspis limits diagnostic characters for most species (except P. cabindae) to subtle features, chiefly related to coloration pattern. Comparing P. namibiana sp. nov. with P. cabindae, the newly described species can be easily distinguished by not having supranasals (present in P. cabindae), by having a ablepharine eye (preablepharine eye in P. cabindae), and by having the frontoparietals fused (divided in P. cabindae). In comparison with P. wahlbergi, P. namibiana sp. nov. can be distinguished by its coloration, namely by the lack of the broad black lateral stripe and bordering ventrolateral white stripe extending from the supralabials through the ear to the groin (males) or terminating behind the shoulder (females). Panaspis namibiana sp. nov. can be distinguished from nominotypical P. maculicollis on the basis of the lack of black patch extending from ear to just beyond the shoulder with two or three vertical or diagonal rows of white spots. Distribution. The newly described species is currently only known from central and northern Namibia. Confirmed localities are in three clusters, one in the northwest above the Western Escarpment from Sesfontein in the south to near Opuwo in the north, one in the Otavi Highlands, and a third in central Namibia in the Khomas Hochland (Fig. 3). It is likely that the species occurs in the intervening regions at higher elevations where appropriate habitat is available, but further sampling is needed to confirm its presence. Habitat and Natural History notes. With the exception of Paracamp, near Sesfontein (~ 650 m), all localities are above 1200 m. The species is terrestrial and is typically found under leaf litter or other debris. Although most specimens were captured by day, MCZ R183767 was active in the leaf litter well after sunset (Fig. 4). Specimen CAS MCZ MCZ CM LACM LACM LACM PEM PEM PEM PEM 206967 R-183767 R- 190318 130371 77840 77841 77842 R 2123 R 2124 R 10262 R 10273 (holotype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) Locality Opuwo Paracamp, Kamanjab 80 km E 40 km 40 km 40 km Kaoko Otavi Kaoko Otavi Okahandja Okahandja road, 21 km Sesfontein Sesfontein, WNW WNW WNW N of 60 km Grootfontein Grootfontein Grootfontein Palmweg- WNW Sesfontein Kamanjab road SVL 29.6 30.8 30.1 35.6 39.8 34.8 33.0 35.4 29.3 42.6 38.7 TL 38.6 Broken Broken Broken Broken Broken Broken Broken Broken 50.0 55.7 HL 5.9 5.6 5.9 5.7 5.5 5.5 6.0 5.7 5.5 5.8 5.6 HH 2.5 2.5 2.0 2.2 2.2 2.4 2.3 2.6 2.3 2.6 3.2 HW 3.2 3.6 3.9 3.1 3.2 3.7 3.8 3.5 3.2 3.6 3.8 LA 1.8 2.5 2.3 1.7 1.7 2.0 2.0 2.5 2.1 3.1 3.3 LFA 1.6 2.2 1.8 2.3 2.3 2.5 2.8 2.3 2.1 2.6 2.8 LL 3.5 3.5 3.3 2.5 2.6 2.6 2.9 3.2 3.1 3.5 3.8 LFL 2.8 3.3 2.5 4.1 4.0 4.5 4.0 3.1 3.0 3.3 3.5 LD 18.3 19.0 18.4 22.1 25.0 20.4 20.1 20.7 17.3 24.8 23.0 ED 1.2 1.2 1.1 1.1 1.1 0.9 1.0 1.1 1.0 1.2 1.2 ET 2.7 2.3 2.4 2.2 2.2 2.2 2.3 2.4 2.3 2.3 2.6 ES 2.3 2.3 2.2 2.3 2.3 2.3 2.2 2.7 2.4 2.3 2.5 MSR 26 25 24 25 25 25 26 24 24 24 22 SAV 64 65 63 64 67 64 65 67 56 66 59 SAD 60 64 60 63 65 63 64 68 57 68 58 LUFT 15 16 14 15 14 15 16 13 14 15 - LUFF 11 11 10 12 10 10 10 11 9 9 8 SC (Left/ 4/4 4/4 4/4 5/5 4/4 4/4 4/5 4/4 4/4 5/5 4/4 Right) SL 6 6 6 6 6 6 6 6 7 6 6/7 CP In contact In contact In contact In contact In contact In contact In contact In contact In contact In contact In contact CPF In contact In contact In contact Separated Narrowly Separated Separated Broad Narrowly Narrowly Separated separated contact separated separated Etymology. The specific epithet ��� namibiana ��� refers to the Republic of Namibia, to which the species is apparently endemic, and is applied here as a substantive in apposition. We propose the English vernacular name ���Namibian Snake-Eyed Skink.���, Published as part of Cer��aco, Luis M. P., Branch, William R. & Bauer, Aaron M., 2018, A new species of African snake-eyed skink (Scincidae: Panaspis) from central and northern Namibia, pp. 140-150 in Zootaxa 4527 (1) on pages 142-147, DOI: 10.11646/zootaxa.4527.1.12, http://zenodo.org/record/2611932, {"references":["Lawrence, R. F. (1929) The reptiles of S. W. A. Journal of the South West Africa Scientific Society, 2, 13 - 27.","FitzSimons, V. F. (1943) The lizards of South Africa. Memoirs of the Transvaal Museum, 1, i-xv + 1 - 528, 24 pls., map.","Mertens, R. (1955) Die Amphibien und Reptilien Sudwestafrikas, aus den Ergebnissen einer im Jahre 1952 ausgefuhrten Reise. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 490, 1 - 172.","Mertens, R. (1971) Die Herpetofauna Sudwest-Afrikas. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 529, 1 - 110.","Visser, J. (1984) Gladde akkedisse. Magiese ' springslang' maar net ' n woelige akkedis! Landbouweekblad, 12 Oktober 1984, 58 - 59, 61.","Branch, W. R. (1988) Field Guide to Snakes and other Reptiles of Southern Africa. Struik Publishers, Cape Town, 328 pp.","Bauer, A. M., Branch, W. R. & Haacke, W. D. (1993) The herpetofauna of the Kamanjab area and adjacent Damaraland, Namibia. Madoqua, 18, 117 - 145.","Branch, W. R. (1994) Field Guide to Snakes and other Reptiles of Southern Africa. 2 nd Edition. Struik Publishers, Cape Town, 328 pp.","Branch, W. R. (1998) Field Guide to Snakes and other Reptiles of Southern Africa. 3 rd Edition. Struik Publishers, Cape Town, 399 pp.","Griffin, M. (2003) Annotated Checklist and Provisional National Conservation Status of Namibian Reptiles. Biodiversity Inventory. Namibia Scientific Society, Windhoek, [2] + 169 pp.","Herrmann, H. W. & Branch, W. R. (2013) Fifty years of herpetological research in the Namib Desert and Namibia, with an updated and annotated species checklist. Journal of Arid Environments, 93, 94 - 115. https: // doi. org / 10.1016 / j. jaridenv. 2012.05.003","Medina, M. F., Bauer, A. M., Branch, W. R., Schmitz, A., Conradie, W., Nagy, Z. T., Hibbitts, T. J., Ernst, R., Portik, D. M., Nielsen, S. V., Colston, T. J., Kusamba, C., Behangana, M. & Rodel, M. - O. (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100, 409 - 423. https: // doi. org / 10.1016 / j. ympev. 2016.04.026","Greer, A. E. (1974) The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, 31, 1 - 67. https: // doi. org / 10.1071 / ajzs 031"]}

A recent molecular revision of the snake-eyed skinks of the genus Panaspis Cope, 1868 uncovered extensive cryptic diversity within the P. wahlbergi (Smith, 1849) and P. maculicollis Jacobsen & Broadley, 2000 species complexes. We here describe an unnamed central and northern Namibian lineage of the P. maculicollis group as a new species. We base the description on previously published molecular evidence and newly collected morphological, meristic and coloration data. Members of the group share a conservative morphology and the newly described species is very similar to P. maculicollis, from which it differs mostly on the basis of its coloration. This description raises the number of Panaspis recognized to 17 species and reinforces the already strong evidence that northern Namibia is a hotspot of reptile diversity.

Panaspis namibiana Ceríaco, Branch & Bauer sp. nov. (Fig. 1–2) Chresonymy (includes only citations unambiguously pertaining to specimens referable to the new species, or explicitly stating or depicting its geographic range). Ablepharus wahlbergi (Lawrence 1929:26) Ablepharus wahlbergii (FitzSimons 1943:236; Mertens 1955:74, pl. 23, fig. 143; Mertens 1971:64) Panaspis (Afroablepharus) wahlbergi (Visser 1984:59) Panaspis wahlbergii (Branch 1988:134; Bauer et al. 1993:134; Branch 1994:134; Branch 1998:159). Panaspis sp. nov. (Griffin 2003:60). Afroblepharus wahlbergi (Herrmann & Branch 2013:29). Afroablepharus sp. Namibia (Medina et al. 2016:411). Holotype. CAS 206967, unsexed subadult, adjacent to Opuwo Road (D3704), 21 km N of Palmweg / Sesfontein Road (D3706) (18.9628333° S, 13.758666° E; WGS-84; Fig. 3), Kunene Region, Republic of Namibia, collected by Aaron M. Bauer, A. C. Lamb, J.L. Wright, P. Moler, W.R. Branch and R.D. Babb on 25 July 1998 (Fig. 1). Paratypes. Ten specimens, all from the Republic of Namibia: MCZ R183767, unsexed subadult, Sesfontein, Paracamp, Kunene Region (19.13222222° S, 13.5877777° E; WGS-84), collected by Aaron M. Bauer and Anthony P. Russell on 11 December 2002; MCZ R 190318, unsexed subadult, Kamanjab Rest Camp, Kunene Region (19.629611° S, 14.682417° E; WGS-84) collected by Aaron M. Bauer on 29 November 2011; CM 130371 (Fig. 2), unsexed adult, 80 air km E Sesfontein, 60 Air km WNW Kamanjab, Kunene Region (19.332654° S, 14.369919° E; WGS-84), collected by Paul Freed et al. on 3 April 1992 (Fig. 4); LACM 77840, unsexed adult, 40 km WNW Grootfontein, Otjozondjupa Region (19.48333° S, 17.73333° E; WGS-84), collected by R.L. Bezy on 20 November 1973; LACM 77841, 77842, unsexed adults, same collecting data as LACM 77840; PEM R2123, unsexed adult, Kaoko Otavi, Kunene Region (18.3° S, 13.7° E; WGS-84), collected by G. Mclachlan and J. Spence on 14 October 1965; PEM 2124, unsexed subadult, same collecting data as PEM R2123; PEM R10262, unsexed adult, Okahandja, Otjozondjupa Region (21.983333° S, 16.916667° E; WGS-84), collected by R. Bradfield on 23 April 1929; PEM R10273, unsexed adult, same collecting data as PEM R10262. Additional material. SMF 46570, Farm Beula, Kunene Region (19.6608° S, 14.91185 E; WGS-84); SMF 50620, Okahandja, Otjozondjupa Region, (21.983333° S, 16.916667°E; WGS-84); SMF 66144, Warmquelle near Zesfontein, Kunene Region (19.182° S, 13.81384° E; WGS-84); SMF 67050 -02, 60 mi. east of Okahandja, Otjozondjupa Region (21.623° S, 17.64941° E; WGS-84); SAM 0 17483, Kaoko Otavi, Kunene Region (18.3°S, 13.7°E; WGS-84); SAM 0 17557, Okorosave, Kunene Region (18.17° S, 13.8° E; WGS-84); SAM 0 46548, Tsumeb Townlands (19.23° S, 17.72° E; WGS-84), Oshikoto Region; PEM R8845, 2 km south of Grootfontein, Otjozondjupa Region (19.3206° S, 18.39306° E, WGS-84); TM 85249, Farm Uisib 422, Otjozondjupa Region (19.5531° S, 17.21667° E, WGS-84); SMW 1946, Namutoni, Oshikoto Region (18.80776° S, 16.940231° E, WGS-84), SMW 1947, Kaoko Otavi, Kunene Region (18.3° S, 13.7°E; WGS-84); SMW 1948, Klein Windhoek, Khomas Region (22.578° S, 17.10083° E, WGS-84); SMW 1949, Windhoek, Khomas Region (22.5613° S, 17.07214° E; WGS-84); SMW 5700, Barab on Palmwag 702, Kunene Region (19.8214° S, 13.7614° E; WGS-84); SMW 6471, Warmquelle, Kunene Region (19.182° S, 13.81384° E; WGS-84). Diagnosis. Panaspis namibiana sp. nov. can be distinguished from other members of the genus by the following combination of characteristics: 1) absence of supranasals; 2) ablepharine eye (as defined by Greer 1974); 3) frontoparietals fused; 4) dorsum coppery-brown, with a dorsolateral light stripe extending approximately to midbody, and a thin darker band starting on the temporals and extending to midbody; 5) absence of rows of light spots on the neck; 6) absence of a white ventrolateral stripe. Description of the holotype. Unsexed subadult in good condition. Arrangement and relative size of head, body and tail scales typical for Panaspis. Robust, cylindrical body with well-developed pentadactyl limbs. Fore- and hind-limbs do not overlap when adpressed against the body. SVL 29.6 mm, tail length 38.6 mm. Head length 5.9 mm, with relatively acuminate snout (HL 184% HW). Other relevant measurements are presented in Table 1. Rostral wider than high, and visible from above. Nasals widely separated behind rostral by frontonasal. Frontonasal acuminate anteriorly, wider than long. Nostrils small, set posteriorly in the nasals but not bordering the postnasal. Prefrontals in contact with one another (atypical considering the type series), hexagonal, contacting frontonasal, loreals, first supraocular, first supraciliary and frontal. Two loreals, the posterior margins of the largest loreal border preocular, which is wider than high. Frontal length shorter than distance between anterior tip of frontal and tip of snout; frontal in contact with prefrontals anteriorly, two supraoculars (one on each side) and frontoparietals. Frontoparietals fused, in contact with the frontal, three supraoculars, parietals and interparietal. Frontoparietal plus interparietal length 2.2 times length of the frontal. Interparietal quadrangular with visible parietal foramen in anterior part; parietals 1.2 times larger than frontoparietals and contacting each other at the anterior point of the interparietal. Parietals in contact. A pair of large, broad nuchals collectively bordered by a total of nine dorsals. Supraoculars three. Supraciliaries three (left side) and four (right side), third largest. Pretemporals two. Tympanum visible, approximately one fifth the height of the eye. Supralabials six, the fifth and sixth being the suboculars. Ablepharine eye. Infralabials five. Postmental bordering five scales (mental, two primary chin-shields, and one infralabial on each side). Ventral scales smooth. MSR 22, SAD 58, SAV 55. Limbs with five digits; scales on soles of hands and feet smooth. Relative length of digits of manus IV=III>II=V>I relative length of digits of pes IV>III>II>I>V. Finger-IV scales 9 (right side), Toe-IV scales 13 (right side). Tail long, robust and tapering smoothly. In preservative, background color of flanks and upper side of head, neck, dorsum, legs and tail is coppery brown, with a darker dot in the anterior part of each scale. Posteriorly the tail is lighter, with disjunct dark speckles. A beige stripe, less than one scale wide, limited by the dorsal background color above and by a thin stripe of darker brown below (especially evident under the anterior part of eye), extends from the temporals and runs along the dorsolateral surface to the midbody. Areas below the dark brown stripe become lighter, until they reach the ventral side, which is beige. Supralabials light beige anteriorly and posteriorly, with strong coppery brown markings in the middle. This pattern extends to the infralabials. Venter uniformly light-beige, as are the palmar regions of hands and feet. Variation. Variaton in scalation and body measurements of the paratypes of Panaspis namibiana sp. nov. is reported in Table 1. The majority of the paratypes agree entirely with the holotype, except regarding the contact between the prefrontals, which are separated in paratypes CM 130371, LACM 77841, LACM 77842, PEM R10273, and narrowly separated in LACM 77840, PEM R2124 and PEM R 10262. Comparison with other South and Southwestern African Panaspis. As the molecular framework of Medina et al. (2016) provided evidence of the independence of the P. namibiana lineage from all other taxa, we here restrict our morphological comparisons to those named congeners occurring in Namibia, Angola, Botswana and South Africa. The extreme morphological conservatism of Panaspis limits diagnostic characters for most species (except P. cabindae) to subtle features, chiefly related to coloration pattern. Comparing P. namibiana sp. nov. with P. cabindae, the newly described species can be easily distinguished by not having supranasals (present in P. cabindae), by having a ablepharine eye (preablepharine eye in P. cabindae), and by having the frontoparietals fused (divided in P. cabindae). In comparison with P. wahlbergi, P. namibiana sp. nov. can be distinguished by its coloration, namely by the lack of the broad black lateral stripe and bordering ventrolateral white stripe extending from the supralabials through the ear to the groin (males) or terminating behind the shoulder (females). Panaspis namibiana sp. nov. can be distinguished from nominotypical P. maculicollis on the basis of the lack of black patch extending from ear to just beyond the shoulder with two or three vertical or diagonal rows of white spots. Distribution. The newly described species is currently only known from central and northern Namibia. Confirmed localities are in three clusters, one in the northwest above the Western Escarpment from Sesfontein in the south to near Opuwo in the north, one in the Otavi Highlands, and a third in central Namibia in the Khomas Hochland (Fig. 3). It is likely that the species occurs in the intervening regions at higher elevations where appropriate habitat is available, but further sampling is needed to confirm its presence. Habitat and Natural History notes. With the exception of Paracamp, near Sesfontein (~ 650 m), all localities are above 1200 m. The species is terrestrial and is typically found under leaf litter or other debris. Although most specimens were captured by day, MCZ R183767 was active in the leaf litter well after sunset (Fig. 4). Specimen CAS MCZ MCZ CM LACM LACM LACM PEM PEM PEM PEM 206967 R-183767 R- 190318 130371 77840 77841 77842 R 2123 R 2124 R 10262 R 10273 (holotype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) Locality Opuwo Paracamp, Kamanjab 80 km E 40 km 40 km 40 km Kaoko Otavi Kaoko Otavi Okahandja Okahandja road, 21 km Sesfontein Sesfontein, WNW WNW WNW N of 60 km Grootfontein Grootfontein Grootfontein Palmweg- WNW Sesfontein Kamanjab road SVL 29.6 30.8 30.1 35.6 39.8 34.8 33.0 35.4 29.3 42.6 38.7 TL 38.6 Broken Broken Broken Broken Broken Broken Broken Broken 50.0 55.7 HL 5.9 5.6 5.9 5.7 5.5 5.5 6.0 5.7 5.5 5.8 5.6 HH 2.5 2.5 2.0 2.2 2.2 2.4 2.3 2.6 2.3 2.6 3.2 HW 3.2 3.6 3.9 3.1 3.2 3.7 3.8 3.5 3.2 3.6 3.8 LA 1.8 2.5 2.3 1.7 1.7 2.0 2.0 2.5 2.1 3.1 3.3 LFA 1.6 2.2 1.8 2.3 2.3 2.5 2.8 2.3 2.1 2.6 2.8 LL 3.5 3.5 3.3 2.5 2.6 2.6 2.9 3.2 3.1 3.5 3.8 LFL 2.8 3.3 2.5 4.1 4.0 4.5 4.0 3.1 3.0 3.3 3.5 LD 18.3 19.0 18.4 22.1 25.0 20.4 20.1 20.7 17.3 24.8 23.0 ED 1.2 1.2 1.1 1.1 1.1 0.9 1.0 1.1 1.0 1.2 1.2 ET 2.7 2.3 2.4 2.2 2.2 2.2 2.3 2.4 2.3 2.3 2.6 ES 2.3 2.3 2.2 2.3 2.3 2.3 2.2 2.7 2.4 2.3 2.5 MSR 26 25 24 25 25 25 26 24 24 24 22 SAV 64 65 63 64 67 64 65 67 56 66 59 SAD 60 64 60 63 65 63 64 68 57 68 58 LUFT 15 16 14 15 14 15 16 13 14 15 - LUFF 11 11 10 12 10 10 10 11 9 9 8 SC (Left/ 4/4 4/4 4/4 5/5 4/4 4/4 4/5 4/4 4/4 5/5 4/4 Right) SL 6 6 6 6 6 6 6 6 7 6 6/7 CP In contact In contact In contact In contact In contact In contact In contact In contact In contact In contact In contact CPF In contact In contact In contact Separated Narrowly Separated Separated Broad Narrowly Narrowly Separated separated contact separated separated Etymology. The specific epithet “ namibiana ” refers to the Republic of Namibia, to which the species is apparently endemic, and is applied here as a substantive in apposition. We propose the English vernacular name “Namibian Snake-Eyed Skink.&rdquo

A small skink of the genus Leptosiaphos from the Adamaoua Massif in Central-Northern Province of Cameroon is described. The new species is most similar to the East African Leptosiaphos kilimensis from which it differs in coloration and in having 22 scales around midbody. The genus Panaspis Cope, 1869, sensu lato has been thoroughly studied morphologically, espe- cially in west Africa (Fuhn, 1969, 1970, 1972; Perret, 1973, 1975, 1982). Four subgenera have been described within Panaspis (Afroablepharus Greer, 1974; Lacertaspis Perret, 1975; Leptosiaphos Schmidt, 1943; Panaspis Cope, 1869). Several of these subgenera have also been elevated to full genera (Broadley, 1989), but this arrangement has not been followed by all subsequent authors (e.g., Bohme, 1975; Bohme and Schmitz, 1996; LeBre- ton, 1999). Because the last complete revision of the subgenera within Panaspis sensu lato (Broad- ley, 1989) regarded Leptosiaphos as a full genus, we herein follow this classification scheme. Perret (1973) made a comprehensive revision of Panaspis sensu lato known to occur in Cameroon. He recognized 12 forms as full species, which he divided tentatively into three groups according to their general morphology: sepsinoid, with an elongated body, short and robust limbs, not overlapping when adpressed to body, a muscular tail enlarged at its base (amieti, fuhni, gemmiventris, lepesmei, pauliani, vigintiser- ierum); lacertiform, with slender body, very long limbs, fingers and toes with an increased number of subdigital lamellae, limbs which overlap when adpressed to body, a long and thin tail, more then twice the snout—vent length (SVL), not enlarged at its base (reichenowi, rohdei); and mabuiform, with an increased number of scales around midbody and supranasals present or absent (africana, breviceps, kitsoni, nimbaensis (sic)). He later (Perret, 1975) referred these groups to three subgenera, respectively Leptosiaphos Schmidt, 1943, Lacertaspis subg. nov. and Panaspis Cope

The genus Panaspis in southern Africa is reviewed and a new species, P. maculicollis is described. This species differs from P wahlbergii (A. Smith) in lacking a white ventrolateral stripe, while breeding males have a black patch on the neck with diagonal rows of white spots and the vermillion ventral coloration is restricted to the chin and throat instead of covering the entire ventral surface. It is also smaller in size, with an average snout-vent length of 35 mm, compared with 40 mm in P. wahlbergii. The two species have been recorded in sympatry in the north-eastern Kruger National Park. Panaspis wahlbergii inhabits the moist savannas to the east and south of the subcontinent, whereas the distribution of P. maculicollis seems to be centred on the Kalahari sand areas of northern Botswana and western Zimbabwe, extending eastwards via the Zambezi and Limpopo valleys into western Mozambique and the Northern Province of South Africa.

Partial sequences of three mitochondrial DNA genes, 12S rDNA, 16S rDNA and cytochrome b, and one nuclear gene, c-mos, were used to assess the phylogenetic relationships of species belonging to the genus Afroablepharus from the volcanic islands of the Gulf of Guinea (West Africa) and neighboring continental Africa. Additionally, partial sequences of cytochrome b were used to compare levels of sequence divergence within populations. The three forms from Sao Tome, Principe and Annobon (one per island) are genetically distinct, with high levels of divergence, supporting the recognition of a distinct species in each island. Populations within each island contain very low levels of genetic diversity. These three forms form a monophyletic group suggesting a single initial colonization followed by radiation to the other islands, possibly from Sao Tome to Principe and Annobon. This is contrary to what was found in other reptiles from these islands such as Mabuya (sensu lato) and Hemidactylus, which colonized the islands multiple times. Assuming a molecular clock for cytochrome b of about 2% divergence per million years (usually applied to Sauria), the lineage on Annobon island exceeds the age of the island, thus casting further doubt on this widely used divergence estimate. Partial sequences of c-mos showed no variation within islands. Five to seven sites were variable among islands, which is a high value further supporting the treatment of each island form as a distinct species.

The scincid taxa connected to the genus Panaspis sensu lato have undergone various taxonomic changes based mainly on morphological studies in the last century, but their true relationships are still far from clear. In the present study we applied Bayesian, maximum parsimony, and maximum likelihood methods of phylogenetic inference to mitochondrial 12S rRNA and 16S rRNA gene fragments to examine the degree of genetic variation within the genus. We found considerable genetic differentiation between the different proposed subgenera. Similarly, we could show that two species considered to be part of the former subgenus Leptosiaphos are in reality genuine members of the genus Lacertaspis. Based on our results we propose the recognition of the four former subgenera as full genera, and we identify several additional, possibly subgeneric, lineages within the different newly recognized full genera.

It is generally admitted that the some 38 species referred to Ablepharus Lichtenstein, 1823 (Sauria, Scincidae) belong to the same evolutionary level. Smith (1935), followed by Parker (1936), De Witte (1936), Mitchell and Storr (in. litt.) pointed out the “polyphyletic” origin of the “ablepharine” skinks, and Greer (1967) presented arguments on the basis of the skull morphology. The study of nearly all the skinks assigned to the genus Ablepharus is in agreement with the above statements. A tentative new generic arrangement is presented, based on the assumption of several parallel evolutionary series, starting from closely related although different lines, showing convergent morphological features under the action of a similar way of life (burrowing and cryptic). The “ablepharine” skinks assemblage includes different evolutionary levels, which have to be referred to distinct genera. Only bivittatus, pannonicus, deserti and kitaibelii are assigned to Ablepharus Lichtenstein; boutonii belongs to Cryptoblepharus Wiegmann. The African “ablepharine” skinks, together with several other species having a transparent window in the lower lid, are included in Panaspis Cope. The Australian species commonly ascribed to the genus Ablepharus, should be considered as belonging to following generic taxa: 1. Morethia Gray, 1844, including the species lineo-ocellata, butleri, taeniopleura; 2. Pseudemoia Fuhn, 1967, monotypic (spenceri); 3. Cryptoblepharus Wiegmann, 1834, monotypic (boutonii), but the species is polytypic. 4. Notoscincus n. g., with the species ornatus and wotjulum; 5. Menetia Gray, 1844, monotypic (greyi); 6. Species incertae sedis, burnetti (polytypic); 7. Proablepharus n. g., including tenuis (polytypic); 8. Species incertae sedis, with the species davisi and kinghorni; 9. Lerista Bell, 1833, including elegans, distinguenda, lineata, muelleri, timida and all the species assigned to Rhodona and Nodorha (Greer, 1967).

Leaf-litter skinks of the genus Panaspis are currently represented in the oceanic islands of the Gulf of Guinea by two species, Panaspis africana and P. annobonensis. Here we describe a third speci...

The populations of three sympatric scincid lizards, Mabuya buettneri, Mabuya maculilabris and Panaspis nimbaensis, have been studied for five consecutive years in the Lamto savanna (Ivory Coast). The seasonal abundance of the three species differ greatly, even though they all increase during the rainy season (April to November). The differences stem mainly from differences in the hatching dates of the three species. Six months are required for these skinks to reach a mature size and reproduce. The average yearly reproductive capacity of the females of M. buettneri, M. maculilabris and P. nimbaensis is, respectively, 13, 30 and 13 eggs produced in 1.5, 5.5 and 5.0 clutches per year. Crude mortality of eggs and hatchlings is high, and survivorship to maturity is low: 18, 8 and 25% of eggs produced, respectively. After sexual maturity the death rate for females is lower than that of males in all species, but it is fairly high for both sexes: the best mean life expectancy after maturity is about 9 months for female M. maculilabris and the lowest is 1.3 months for male