(2024.05.08) Trachylepis - Nowe Gatunki

THE TRACHYLEPIS (SQUAMATA: SCINCIDAE)
OF ANGOLA: AN INTEGRATIVE TAXONOMIC REVIEW

WITH THE DESCRIPTION OF SEVEN NEW SPECIES
LUIS M.P. CERÍACO

Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO), InBIO Laboratório
Associado, Campus de Vairão, Universidade do Porto;
BIOPOLIS Program in Genomics, Biodiversity and Land Planning, Campus de Vairão Departamento
de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro
Museu Nacional de História Natural e da Ciência, Lisboa
Section of Amphibians and Reptiles, Carnegie Museum of Natural History, Pittsburgh
MARIANA P. MARQUES
Section of Amphibians and Reptiles, Carnegie Museum of Natural History Pittsburgh;
Museu Nacional de História Natural e da Ciência, Lisboa
CIBIO-InBIO, Universidade do Porto
BIOPOLIS Program in Genomics, Biodiversity and Land Planning
Departamento de Biologia, Faculdade de Ciências da Universidade do Porto
DIOGO PARRINHA
CIBIO-InBIO, Universidade do Porto
BIOPOLIS Program in Genomics, Biodiversity and Land Planning
Departamento de Biologia, Faculdade de Ciências da Universidade do Porto
ARTHUR TIUTENKO
Friedrich-Alexander-Universität Erlangen-Nürnberg
JEFFREY L. WEINELL
Department of Herpetology, American Museum of Natural History
BRETT O. BUTLER
Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México
Museo de Zoología “Alfonso L. Herrera,” Departamento de Biología Evolutiva, Facultad de Ciencias,
Universidad Nacional Autónoma de México
AARON M. BAUER
Department of Biology and Center for Biodiversity and Ecosystem Stewardship,
Villanova University, Villanova, PA
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
Number 465, 153 pp., 111 figures, 10 tables, 6 plates
Issued February 20, 2024
Copyright © American Museum of Natural History 2024 ISSN 0003-0090

CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Specimen Collection and Sampling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
DNA Sequences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Phylogenetic Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Morphological Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Additional Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
DNA Sequences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Phylogenetic Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Species Concept . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
Taxonomic accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Trachylepis albilabris (Hallowell, 1857)—Guinea Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Trachylepis albopunctata (Bocage, 1867)—Angolan Variable Skink . . . . . . . . . . . . . . . . . . . . . . 19
Trachylepis ansorgii (Boulenger, 1907)—Ansorge’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
Trachylepis attenboroughi, sp. nov.—Attenborough’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Trachylepis bayonii (Bocage, 1872)—Bayão’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Trachylepis binotata (Bocage, 1867)—Ovambo Tree Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Trachylepis bocagii (Boulenger, 1887)—Bocage’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Trachylepis bouri, sp. nov.—Bour’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Trachylepis chimbana (Boulenger, 1887)—Chimba Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Trachylepis damarana (Peters, 1870)—Damaraland Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Trachylepis hilariae, sp. nov.—Hilária’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Trachylepis hoeschi (Mertens, 1954)—Hoesch’s Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Trachylepis huilensis (Laurent, 1964)—Huíla Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Trachylepis laevis (Boulenger, 1907)—Angolan Blue-Tailed Skink . . . . . . . . . . . . . . . . . . . . . . . 77
Trachylepis maculilabris (Gray, 1845)—Speckled-Lipped Skink . . . . . . . . . . . . . . . . . . . . . . . . . 82
Trachylepis notabilis (Peters, 1879)—Notable Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Trachylepis occidentalis (Peters, 1867)—Western Three-Striped Skink . . . . . . . . . . . . . . . . . . . 91
Trachylepis ovahelelo, sp. nov.—Ovahelelo Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Trachylepis punctulata (Bocage, 1872)—Speckled Sand Skink . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Trachylepis raymondlaurenti Marques et al., 2019—Laurent’s Long-Tailed Skink . . . . . . . . . 104
Trachylepis sulcata (Peters, 1867)—Western Rock Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Trachylepis suzanae, sp. nov.—Suzana’s Wedge-Snouted Skink . . . . . . . . . . . . . . . . . . . . . . . . . 112
Trachylepis vunongue, sp. nov.—Mwene Vunongue Skink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Trachylepis wahlbergii (Peters, “1869” 1870)—Wahlberg’s Striped Skink . . . . . . . . . . . . . . . . 126
Trachylepis wilsoni, sp. nov.—Wilson’s Wedge-Snouted Skink . . . . . . . . . . . . . . . . . . . . . . . . . 132
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . after page 147
Appendix SA1. List of Material Examined (online supplement: https://doi.org/10.5531/sd.sp.65)
Appendix SA2. Mean Pairwise Genetic Distances (online supplement: https://doi.org/10.5531/sd.sp.65)
2
ABSTRACT
The genus Trachylepis is currently represented by 21 species in Angola, most of them part of
nomenclaturally and taxonomically challenging species complexes. In this study we present an inte-
grative taxonomic revision of the genus in Angola and describe seven new species: Trachylepis atten-
boroughi, sp. nov., Trachylepis bouri, sp. nov., Trachylepis hilariae, sp. nov., Trachylepis ovahelelo, sp.
nov., Trachylepis suzanae, sp. nov., Trachylepis vunongue, sp. nov., and Trachylepis wilsoni, sp. nov.
As result of our taxonomic revisions, 25 valid Trachylepis species are now confirmed from Angola.
A phylogenetic analysis using a combination of mitochondrial (16S, ND2) and nuclear (RAG1)
markers, as well as morphological data, supports the recognition of the new species. In addition,
data support the revalidation of Trachylepis albopunctata (Bocage, 1867), Trachylepis notabilis (Peters,
1879), and Trachylepis ansorgii (Boulenger, 1907). We also provide a redefinition of Euprepes anchi-
etae Bocage, 1866, which we synonymize with Trachylepis maculilabris (Gray, 1845). Given that the
type material for Trachylepis albopunctata, T. angolensis, and T. anchietae has been lost or destroyed,
we designate neotypes for the purpose of nomenclatural stability. The description of the new species
and the revision and revalidation of previously described Angolan species contribute to a better
understanding of the taxonomy and biogeography of the genus, as well as to the general biogeo-
graphic patterns and evolution of the Angolan fauna.
INTRODUCTION
every researcher who has worked in Angolan her-
Trachylepis Fitzinger, 1843, is one of the most petofauna since the 19th century has examined spe-
diverse and species-rich genera in the family Scin- cies of the genus. Eight species have been described
cidae, currently comprising 87 recognized species based on Angolan material, including Trachylepis
(Uetz et al., 2024) distributed throughout the Afri- albopunctata (Bocage, 1867a), Trachylepis bayonii
can continent, Madagascar, Arabia, and one spe- (Bocage, 1872), Trachylepis binotata (Bocage,
cies occurring on Fernando de Noronha Island, 1867b), Trachylepis bocagii (Boulenger, 1887), Tra-
eastern Brazil. The genus belongs to the Mabuyinae chylepis chimbana (Boulenger, 1887), Trachylepis
group (Weinell et al., 2019) and its members are laevis (Boulenger, 1907), Trachylepis punctulata
the dominant and most conspicuous surface (Bocage, 1872), and Trachylepis raymondlaurenti.
active (e.g., terrestrial, arboreal, and rupiculous) Many of these are widespread species, while others
skinks throughout most of its range. In the last belong to problematic species complexes. Other
decade, numerous species have been described: taxa of dubious taxonomic validity, such as Euprepes
Trachylepis adamastor Ceríaco, 2015, Trachylepis notabilis Peters, 1879, Mabuya bayonii huilensis Lau-
principensis Ceríaco et al., 2016, Trachylepis rent, 1964, and Trachylepis monardi Marques et al.,
thomensis Ceríaco et al., 2016, Trachylepis gon- 2018 (nomen novum for Mabuia striata angolensis
wouoi Allen et al., 2017, and Trachylepis raymond- Monard, 1937), have also been described based on
laurenti Marques et al., 2019. Recently this genus Angolan material, but their taxonomic identity has
was the subject of a comprehensive species-level never been adequately assessed.
phylogeny by Weinell et al. (2019). Despite the considerable number of authors
Angola hosts an extraordinary diversity of Tra- who have dealt with Angolan Trachylepis, to date
chylepis species, with almost one fourth of all known there are no comprehensive reviews or identifica-
African species occurring in the country (Marques tion keys for the Angolan species as exist, for exam-
et al., 2018, 2019a). The most recent checklists ple, for southeast Africa (Broadley, 2000),
regarding the reptile fauna of the country note the Cameroon and the central African countries (Allen
existence of 21 species in Angola (Marques et al., et al., 2017), and Ethiopia (Spawls et al., 2023). As
2018, 2019a; Branch et al., 2019). As they are among part of ongoing research on the herpetofauna of
the most abundant animals in the territory, nearly Angola, we present a comprehensive review of the
3
4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 465
Trachylepis of Angola. We provide a phylogeny of Natural, Luanda, Angola; MNHN-RA, Muséum

the genus including all species purported to occur national d’Histoire naturelle, Paris; MTD, Museum
in Angola, present updated taxonomic accounts für Tierkunde, Senckenberg Natural History Col-
and new diagnoses for all the Angolan taxa, desig- lections Dresden, Dresden; MUHNAC, Museu
nate neotypes for T. albopunctata, T. angolensis, and Nacional de História Natural e da Ciência da Uni-
T. anchietae, and describe seven new species. versidade de Lisboa, Lisbon; NHMW, Naturhisto-
risches Museum, Vienna; NMZB, Natural History
Museum of Zimbabwe, Bulawayo, Zimbabwe;
MATERIALS AND METHODS PEM, the Port Elizabeth Museum, Bayworld, Gqe-
berha (Port Elizabeth), South Africa; SMF, Senck-
Specimen Collection and Sampling
enberg Forschungsinstitut und Naturmuseum,
Specimens collected for this study were fixed in Frankfurt am Main; TM, Ditsong National
the field with 10% buffered formalin and trans- Museum of Natural History, Pretoria; UF, Natural
ferred to 70% ethanol for long-term preservation. History Museum of the University of Florida,
Liver tissue was removed before formalin fixation Gainesville, FL; USNM, National Museum of Natu-
and preserved in 96%–100% ethanol or in RNAL- ral History, Smithsonian Institution, Washington,
ater and subsequently transferred to 95% ethanol DC; ZMB, Museum für Naturkunde, Berlin; ZMH,
for storage. A total of 774 specimens were con- Zoologisches Museum Hamburg, Hamburg; and
sulted for this study (appendix SA1 in the online ZSM, Zoologische Staatssammlung München,
appendix: https://doi.org/10.5531/sd.sp.65). For Munich (for a detailed list see respective taxonomic
mensural and meristic comparisons, we examined accounts and appendix SA1 in the online supple-
377 specimens of Trachylepys species (including ment). Specimen listed with the acronym AMB
type specimens of T. bayonii [ZMB 6477], T. bino- belong to Aaron M. Bauer field series will be acces-
tata [ZMB 7794], E. petersii [ZMB 6479], T. notabi- sioned in CAS. Additional specimens were also
lis [ZMB 9204], and T. punctulata [ZMB 6478]) examined but only to confirm specific identity.
deposited in the following collections: AMNH, Information on morphological characters of species
American Museum of Natural History, New York, and/or type material that could not be examined,
NY; ANSP, Academy of Natural Sciences of Drexel as well as supplemental data for all Trachylepis, was
University, Philadelphia, PA; BMNH, Natural His- obtained from the relevant literature (e.g., Bocage,
tory Museum, London; CAS, California Academy 1867a, 1867b, 1872, 1895; Boulenger, 1887, 1907;
of Sciences, San Francisco, CA; CM, Carnegie Schmidt, 1919; Laurent, 1947, 1954, 1964; Broadley,
Museum of Natural History, Pittsburgh, PA; 1974a, 1974b, 2000; Hoogmoed, 1974; Branch,
FMNH, Field Museum of Natural History, Chi- 1998; Ceríaco et al., 2016a; Allen et al., 2017; Wei-
cago, IL; IICT, Instituto de Investigação Científica nell and Bauer, 2018; Pietersen et al., 2021).
Tropical, Lisbon; INBAC, Instituto Nacional da
Biodiversidade e Áreas de Conservação, Kilamba,
DNA Sequences
Angola MCZ, Museum of Comparative Zoology,
Harvard University, Cambridge, MA; MD, Museu We generated novel DNA sequence data for
Regional do Dundo, Dundo, Angola; MHNC-UP, previously unsampled Angolan Trachylepis spe-
Museu de História Natural e da Ciência da Univer- cies, from tissues collected over multiple field
sidade do Porto, Porto, Portugal; MHNC, Musée expeditions to Angola since 2012. In addition to
d’Histoire Naturelle de La Chaux-de-Fonds, La new data, we included sequences from the Wei-
Chaux-de-Fonds, Switzerland; MHNG, Muséum nell et al. (2019) dataset for 76 Trachylepis and 13
d’Histoire Naturelle, Geneva; MHNN, Museum outgroup species. Outgroup taxa included spe-
d’Histoire Naturelle de Neuchâtel, Neuchâtel, Swit- cies from the mabuyine genera Chioninia, Copeo-
zerland; MNHNL, Museu Nacional de História glossum, Dasia, Eumecia, Eutropis, Heremites,
2024 CERÍACO ET AL.: TRACHYLEPIS (SQUAMATA: SCINCIDAE) OF ANGOLA 5
Lubuya, and Toenayar, and from the eugongyline taining ND2 and tRNAs. Best-fit substitution
genera Caledoniscincus and Cryptoblepharus. models were selected automatically for each par-
We extracted genomic DNA using a salt-extrac- tition by using the ‘-m TEST’ option in IQTREE,
tion protocol (Aljanabi and Martinez, 1997). Next, which implements the program ModelFinder
polymerase chain reactions (PCR) were performed (Kalyaanamoorthy et al., 2017). Clade support
to amplify portions of two mitochondrial loci and values were estimated from 10000 ultrafast boot-
one nuclear locus commonly used in systematic straps (UFBoot), and we considered UFBoot ≥95
studies. Mitochondrial loci included 16S ribosomal to be strong support for the monophyly of lin-
RNA (16S) and a locus containing the protein- eages (Minh et al., 2013; Hoang et al., 2018).
coding gene NADH dehydrogenase subunit 2
(ND2) and transfer RNA genes tRNA (Ala), tRNA
Morphological Methods
(Asx), tRNA (Cys), and tRNA (Tyr); the nuclear
locus was recombination activation protein 1 Specimens were measured with a digital caliper
(RAG1). Thermocycler profiles (PCR) and primers (0.1 mm), while lepidosis was observed with the
(PCR and sequencing) were the same as described help of a stereomicroscope. Scale nomenclature,
in Weinell et al. (2019). Products of PCR amplifica- scales counts, and measurements used in the
tion were confirmed using gel electrophoresis and descriptions follow Broadley (2000), Ceríaco
cleaned using a magnetic bead solution (Rohland (2015), Ceríaco et al. (2016a), and Marques et al.
and Reich, 2012) before cycle sequencing using (2019a). We measured the following 21 characters:
Big-Dye V3.1 chemistry. An additional magnetic SVL, snout–vent length, from snout tip to vent; TL,
bead clean up was performed before analyzing tail length, from cloaca to tip of tail, measured only
cycle sequencing on an ABI 3730xl sequencer at in specimens with complete, original tails; HH,
Villanova University. head height, from base of maxilla to top of head;
We used Geneious v11.0.2 to assemble and HL, head length, from tip of snout to anterior tym-
edit novel sequences, MAFFT v7.310 (Katoh and panum border; HW, head width, from lateral edge
Standley, 2013) to separately align sequences for of left parietal to lateral edge of right parietal, above
each locus, and SeqKit v2 (Shen et al., 2016) to eyes; EN, eye-nostril distance, from anterior edge
translate protein-coding regions to check for of eye to nostril; ES, eye-snout distance, from ante-
early stop codons and to trim and concatenate rior edge of eye to tip of snout; IN, internostril dis-
single-locus alignments. We calculated genetic tance, minimum distance between nostrils; MSR,
distances between each pair of individuals at number of scale rows at midbody; SAD, number of
each locus (ignoring sites with ambiguous or scales dorsally, from nuchal (excluded from count)
missing data) using function ‘stringDist’ in R to base of tail (above anal plate); SAV, number of
package Biostrings (Pagès et al., 2021), from scales ventrally, from mental (excluded from count)
which we calculated mean genetic distance at to anal plate (excluded from count); LUFF, number
each locus for each pair of species. We filtered of subdigital lamellae under Finger-IV; LUFT,
tRNA genes and other noncoding sites before number of subdigital lamellae under Toe-IV; SC,
calculating genetic distances for ND2. number of supraciliaries; SL, number of supralabi-
als, with those widened in subocular position indi-
cated between parentheses; CP, type of contact
Phylogenetic Relationships
between parietals; CFP, contact between frontopa-
To estimate phylogenetic relationships under rietals; CSN, contact between supranasals; CPF,
maximum likelihood (ML), we used the program contact between prefrontals; KDS, number of keels
IQTREE v1.6.10 (Nguyen et al., 2015), with sites on dorsal scales; and PS, type of plantar scales. Col-
partitioned by locus and separately for protein- oration pattern was reported and high-resolution
coding and noncoding regions of the locus con- photographs of preserved specimens were taken.
TABLE 1
Minimum Mean Pairwise Genetic Distance and Corresponding Species per Locus for
Trachylepis Species Described Herein
See appendix SA2 for all pairwise genetic distances (https://doi.org/10.5531/sd.sp.TK).
Taxon 16S ND2 RAG1

T. attenboroughi, sp. nov. T. spilogaster (2.14%) T. wahlbergii (12.03%) T. striata (0.78%)
T. bouri, sp. nov. T. bocagii (3.15%) T. bocagii (11.91%) T. bocagii (1.14%)
T. hilariae, sp. nov. T. cf. triebneri (0.69%) — T. variegata (0.74%)
T. ovahelelo, sp. nov. T. vunongue, sp. nov. (2.59%) T. vunongue, sp. nov. (8.15%) T. ansorgii (0.80%)
T. suzanae, sp. nov. T. wilsoni, sp. nov. (1.90%) T. wilsoni, sp. nov. (6.80%) T. wilsoni, sp. nov. (0.90%)
Trachylepis vunongue, sp. nov. Trachylepis ovahelelo, sp. nov. Trachylepis ovahelelo, sp. nov. T. variegata (0.38%)
(2.59%) (8.15%)
T. wilsoni, sp. nov. T. suzanae, sp. nov. (1.90%) T. suzanae, sp. nov. (6.80%) T. suzanae, sp. nov. (0.90%)
Specimens that were measured and scale counted lished genetic data, the number of individuals
are noted in Material Examined sections in the sampled was 221 at the 16S locus, 97 at ND2,
respective taxonomic accounts. Specimens that and 142 at RAG1 (see appendix SA2 in the
were physically examined (but not measured or online supplement: https://doi.org/10.5531/sd.
scale counted) either directly by the authors, sp.65). Our alignment with loci concatenated,
through photos, or by colleagues and whose taxo- which was used for phylogenetic analyses,
nomic identification was unambiguous are listed included 221 individuals (208 individuals from
under Additional Material sections in the same 69 species and 1 nonnominate subspecies of Tra-
taxonomic accounts. chylepis and 13 individuals from outgroup lin-
eages) and 3235 sites (550 sites from 16S; 1496
Additional Data sites from locus containing ND2 and tRNA
genes; and 1189 sites from RAG1). Percent of
Locality data are reported in the form of deci- missing or ambiguous data in our alignment was
mal degrees and use the WGS 84 map datum. 54.1% (in total), 12.82% (16S), 66.50% (ND2 and
Older (non-GPS) records are derived mostly tRNAs), and 57.58% (RAG1). Among Trachylepis
from Marques et al. (2018) and have been geore- species, mean pairwise genetic distances ranged
ferenced using the GEOLocate web application from 0%–11.18% for 16S, 6.67%–25.78% for
(https://www.geo-locate.org). Elevations are all ND2, and 0.24%–6.46% for RAG1 (table 1;
reported as meters above sea level. At the begin- appendix SA2 available online: https://doi.
ning of each account a list of name usages refer- org/10.5531/sd.sp.65).
ring specifically to Angolan records of the given
taxa, i.e., a “regional” chresonymy, is presented.
Phylogenetic Relationships
RESULTS The set of best-fit substitution models
included TIM3 (16S), TPM2 (ND2 protein-cod-
DNA Sequences
ing region), TPM3 (ND2 tRNA genes), and TN
We generated new DNA sequences for 51 (RAG1), with nucleotide frequencies estimated,
individuals from 20 species, including 51, 48, and with the invariable site plus discrete Gamma
and 45 sequences at 16S, ND2, and RAG1 loci, model (Gu et al., 1995) to account for rate het-
respectively. After inclusion of previously pub- erogeneity across sites.
Chioninia vaillantii
• Chioninia delalandii
100
Trachylepis tessellata TJR 38 Oman: Wadi Haql
Trachylepis tessellata TE 100363 Oman: Mughsayl
• 100
Trachylepis socotrana M74 Yemen: Socotra Isl.
Trachylepis socotrana M73 Yemen: Socotra Isl.
100 Trachylepis brevicollis 1 CAS 198871 Kenya: Kajiado
100 99
Trachylepis cristinae JR 15 Yemen: Socotra Archipelago: Abd al Kuri Isl.
Trachylepis brevicollis 2 be 100361 Oman: Dhofar
br 99
100 Trachylepis dichroma MAB 189 Tanzania: Dodoma
Trachylepis dichroma MAB 187 Tanzania: Dodoma
100
Trachylepis perrotetii MVZ 245351 Ghana: Greater Accra
99
Trachylepis perrotetii FMNH 262231 Ghana
Trachylepis perrotetii FMNH 26630 Ghana
• af1 100 100
Trachylepis gonwouoi CAS 261092 Cameroon: Fako
Trachylepis gonwouoi USNM 584340 Republic of Congo: Impongui
100
Trachylepis gonwouoi PEM R-20428 Republic of Congo
100 Trachylepis affinis BYU 62095 Cameroon: Mt. Cameroon
100
100 Trachylepis affinis NCSM 88359 Cameroon: Bioko Isl.
100 Trachylepis affinis PEM R-5425 Gabon: Loango NP
Trachylepis affinis PEM R-5340 Gabon: Rabi
88
99
Trachylepis paucisquamis UWBM 6041 Ghana
100
Trachylepis paucisquamis PEM R-4438 Ivory Coast
99
Trachylepis paucisquamis PEM R-4436 Ivory Coast
Trachylepis aureogularis MVZ 252612 Ghana: Eastern
93
100
Trachylepis albilabris PEM R-5422 Gabon: Loango NP
Trachylepis albilabris PEM R-5384 Gabon: Rabi
• 100
100 Trachylepis albilabris MUHNAC:MB 03-001374 Angola: Cabiri
Trachylepis albilabris MUHNAC:MB 03-001376 Angola: Cabiri
• 100
Trachylepis atlantica MRT 4427 Brazil: Fernando de Noronha
90 Trachylepis atlantica MRT 4429 Brazil: Fernando de Noronha
Trachylepis atlantica MRT 4428 Brazil: Fernando de Noronha
86 Trachylepis quinquetaeniata TJC 1372 Ethiopia: Awash NP: Lodge Falls
100
100 Trachylepis quinquetaeniata MVZ 249790 Ghana
98 Trachylepis quinquetaeniata FMNH 262232 Ghana
qt 100 Trachylepis margaritifera PEM R-9745 Malawi: Mt. Mulanje
100 Trachylepis margaritifera PEM R-5539 Mozambique: Niassa
100 Trachylepis margaritifera CAS 234173 South Africa: Limpopo
Trachylepis margaritifera CAS 234165 South Africa: Limpopo
Sch 100
Trachylepis wrightii MAWR 1 Seychelles
85
Trachylepis sechellensis TRASEY 1 Seychelles
79 Trachylepis polytropis PEM-R 5443 Gabon: Loango NP
af2 100
100
Trachylepis polytropis PEM-R 5337 Gabon: Rabi
100
Trachylepis notabilis MUHNAC:MB 03-001357 Angola: Malanje: Luaca Dam
Trachylepis notabilis MUHNAC:MB 03-001355 Angola: Malanje: Luaca Dam
100 Trachylepis maculilabris 1 PEM R-16325 Malawi: Mt. Mulanje
67 Trachylepis maculilabris 1 PEM R-9737 Malawi: Mt. Mulanje
84 100 Trachylepis thomensis CAS 218722 São Tomé Isl.
Trachylepis thomensis CAS 218821 São Tomé Isl.
95
87 Trachylepis thomensis MUHNAC:MB 03-000963 São Tomé Isl.
99 Trachylepis maculilabris 2 PEM R-19463 Angola: Lunda Norte
ma 78
Trachylepis maculilabris 2 PEM R-5844 Gabon: Nyanga
Trachylepis maculilabris 2 INBAC/AMB 9883 Angola: Cuanza Sul: Cada Amboim
100
100
Trachylepis boulengeri PEM R-5533 Mozambique: Nampula
Trachylepis boulengeri PEM R-16179 Mozambique: Niassa: Mt. Mekula
• 55
99 Trachylepis boulengeri PEM R-13442 Mozambique: Fogo Isl.
Trachylepis boulengeri PEM R-15451 Mozambique: Zambezi Delta: Malinga Pansi
85
100 Trachylepis adamastor CAS 238898 Principe Isl.
Trachylepis adamastor MUHNAC:MB 03-00979 Principe Isl.
85 Trachylepis adamastor MUHNAC:MB 03-00956 Principe Isl.
Trachylepis comorensis 1 ZFMK 62192 Madagascar: Nosy Tanikely
99
92 Trachylepis comorensis 2 MH 23 Comoros: Moheli: Ouhoni
Trachylepis casuarinae 71Mcaguri Mozambique: Fogo Isl.
to i
0.1 substitutions/site • = monophyly enforced during analysis
af1 = affinis group part 1 ma = maculilabris group
af2 = affinis group part 2 qt = quinquetaeniata group
br = brevicollis group Sch = Seychelles group
FIGURE 1. Bayesian inference phylogenetic tree (above and on next two pages; inset shows location on tree
of enlarged section). Tip labels highlighted in green correspond to species described herein. Color and width
of horizontal bars at internal nodes indicate clade support.
Trachylepis gravenhorstii 1 MAB 182 Madagascar: Tulear: Ifaty: Ranobe Village

i 100 Trachylepis elegans 1 PEM FNR 438 Madagascar: Basibass
94 Trachylepis madagascariensis UMMZ 209103 Madagascar: Antananarivo: Vakinankaratra
93
Trachylepis elegans 2 RAN 56248 Madagascar: Toliara: Mt. Vohidava
• Mal 100
Trachylepis gravenhorstii 2 RAX 00265 Madagascar: Namoroka Reserve
Trachylepis tandrefana AF153566 Madagascar: Kirindy
100
100
Trachylepis boettgeri ZSM 362-200 Madagascar
Trachylepis boettgeri UMMZ 208917 Madagascar: Antananarivo: Vakinankaratra
98
100
Trachylepis vato ZSM K-398-2000 Madagascar
Trachylepis vato RAN 56249 Madagascar: Toliara
100
100 Trachylepis aureopunctata RAN 56250 Madagascar: Toliara: Mt. Vohidava
98
Trachylepis aureopunctata RAN 54509 Madagascar: Ambalarano
Trachylepis aureopunctata PEM FNR 437 Madagascar: Toliara
100
Trachylepis depressa PEM-R 17745 South Africa: KwaZulu-Natal: Osabeni
Trachylepis depressa PEM-R 15573 Mozambique: Gaza: Chibuto
100
99
Trachylepis homalocephala homalocephala PEM-R 23918 South Africa: Western Cape
100
Trachylepis homalocephala homalocephala AMB 7072 South Africa: Western Cape
100 Trachylepis homalocephala peringueyi PEM R-17459 South Africa: Northern Cape
Trachylepis homalocephala peringueyi PEM R-17458 South Africa: Northern Cape
• 100
dp Trachylepis acutilabris AMB 6935 Namibia: Kunene
98
86
Trachylepis acutilabris MCZ A-28325 Namibia: Erongo: Farm Omandumba
Trachylepis acutilabris AMB 8847 Namibia: Khomas: vic. Solitaire
100 100 Trachylepis wilsoni, sp. nov. UF 187317 Angola: Namibe: Omahua Chitundolo
64
Trachylepis wilsoni, sp. nov. CAS 254930 Angola: Namibe
Trachylepis wilsoni, sp. nov. UF 187318 Angola: Namibe: Virei
55
100 Trachylepis suzanae, sp. nov. CAS 263357 Angola: Luanda: Kissama
54 Trachylepis suzanae, sp. nov. INBAC/AMB 36478 Angola: Luanda: Kissama
Trachylepis suzanae, sp. nov. PEM R-24047 Angola: Namibe
54 Trachylepis suzanae, sp. nov. CAS 223990 Namibia: Kunene: Khorixas
59 Trachylepis suzanae, sp. nov. CAS 214651 Namibia: Kunene: Khorixas
Trachylepis raymondlaurenti CAS 258401 Angola: Malanje: Cangandala NP
100
Trachylepis megalura PEM R-16775 Tanzania: Mara
98
vr 100
100 Trachylepis laevigata MCR R-184536 South Africa: Limpopo
Trachylepis laevigata M 158 South Africa: North West: Botsalano Nature Reserve
98
Trachylepis laevigata M 248 South Africa: North West: Enselsberg
100 100
Trachylepis damarana PEM R-20514 Angola: Cuando Cubango
Trachylepis damarana MCZ R-193241 Namibia: Otjozondjudpa
100
Trachylepis cf. varia G TJC 1409 Ethiopia: Somali: Filtu
83
100
Trachylepis cf. varia E PEM R-16789 Tanzania: Mara
Trachylepis cf. varia E PEM R-16768 Tanzania: Mara
100
Trachylepis varia PEM R-5542 Mozambique: Niassa
100 Trachylepis varia PEM R-5413 South Africa: Eastern Cape: Transkei: Mtentu Gorge
100
Trachylepis cf. varia C ELI 713 DRC: South Kivu: Lwemba
84
97
Trachylepis cf. varia C EBG 2256 DRC: Katanga: Kilwemapante
100 Trachylepis cf. varia D TJC 1336 Ethiopia: Oromia: Bale
99 96 Trachylepis cf. varia D TJC 1335 Ethiopia: Oromia: Bale
Trachylepis albopunctata CAS 263560 Angola: Namibe: Serra da Neve
100
Trachylepis albopunctata MUHNAC:MB03-001379 Angola: Malanje: Lauca Dam
100 Trachylepis albopunctata CAS 263406 Angola: Huila: Cascatad Huila
94 Trachylepis albopunctata PEM R-21739 Angola: Benguela
84 Trachylepis albopunctata GJ 2766 Angola: Malanje: Kalandula Falls
94 Trachylepis albopunctata PEM R-21738 Angola: Benguela
100 Trachylepis albopunctata INBAC/AMB 10297 Angola: Huambo: Kanjonde
Trachylepis occidentalis MCZ R-184266: Namibia: Kunene: Gai-As
ca 99
100
Trachylepis occidentalis CAS 214511: South Africa: Northern Cape: Farm Avondsehijn
96
Trachylepis capensis CAS 234152: South Africa: Limpopo: Farm Brenhilda
Trachylepis capensis LSUMZ 57253: South Africa: Northern Cape: Richtersveld NP
Trachylepis planifrons TJC 1478: Ethiopia: Oromia: Borena: Dollo Mena
68 Trachylepis binotata MCZ R-185902: Namibia: Kunene: Okangwati
100
100
Trachylepis binotata CAS 263496 Angola: Namibe: Virei Chipumpo
Trachylepis binotata INBAC/AMB 10351 Angola: Namibe: Camacuio
100 Trachylepis hoeschi PEM R-24143 Angola: Namibe: Farm Mucongo
87 Trachylepis hoeschi MCZ R-184257 Namibia: Kunene: Gai-As
100
0.1 substitutions/site ca = capensis group
dp = depressa group
97 • = monophyly enforced
during analysis Mal = Malagasy group
vr = varia group
to ii
Trachylepis spilogaster MB 20874 South Africa: Northern Cape: Farm Boseekoebaard

97 Trachylepis spilogaster MCZ R-183768 Namibia: Kunene
95
100 Trachylepis spilogaster MCZ A-38295 Namibia: Karas
99 Trachylepis spilogaster MCZ A-38261 Namibia: Karas: Helminghause
26
Trachylepis attenboroughi, sp. nov. PEM R-21842 Angola: Cuando Cubango
Trachylepis attenboroughi, sp. nov. CAS 263386 Angola: Huila: Bicuar
99
59 Trachylepis attenboroughi, sp. nov. UF 187322 Angola: Huila: Serra da Leba
Trachylepis attenboroughi, sp. nov. INBAC/AMB 10125 Angola: Huila: Lubango
Trachylepis punctatissima 1 S1031 South Africa: North West: Botsalano Nature Reserve
st 100 Trachylepis sparsa CAS 214458 South Africa: Northern Cape: Farm Avonschijn
93 99
100
Trachylepis sparsa MCZ R-184370 Namibia: Karas
ii Trachylepis sparsa MCZ A-38356 Namibia: Karas
90 Trachylepis punctatissima 3 S976 South Africa: North West: Mafikeng Game Reserve
96
99
Trachylepis punctatissima 3 AMB 8622 South Africa: Mpumalanga
91
Trachylepis punctatissima 3 NMBR 8610 South Africa: Free State: Phuthaditjhaba
99
Trachylepis cf. spilogaster 1 AMB 7614 Namibia: Kunene
35
Trachylepis punctatissima 2 MBUR 00138 South Africa: Mpumalanga: Schoemanskloof
88
Trachylepis cf. spilogaster 2 MCZ Z-37947 Namibia: Kunene: Marienflus
100
Trachylepis wahlbergii 1 MCZ R-193549 Zambia: Copperbelt
Trachylepis wahlbergii 1 MB 20380 Zambia: southwestern
100
Trachylepis wahlbergii 2 CAS 258400 Angola: Malanje: Cangandala NP
59 100 Trachylepis wahlbergii 2 PEM R-21819 Angola: Bie: Lunda Falls
Trachylepis wahlbergii 2 PEM R-21818 Angola: Bie: Lunda Falls
Trachylepis wahlbergii 3 MB 20393 Zambia: southwestern
57 63
Trachylepis punctatissima 4 PEM R-16655 South Africa: Mpumalanga: Lydenburg
59
Trachylepis striata 1 TJC 1373 Ethiopia: Afar: Awash NP
100
Trachylepis striata 2 MBUR 01733 South Africa: Limpopo
100
100 100 Trachylepis striata 2 MOZ 17-463 Mozambique: Nampula: Mount Inago
Trachylepis striata 2 MOZ 17-211 Mozambique: Nampula: Ribaue
99 Trachylepis striata 3 PEM R-16772 Tanzania: Mara
100 Trachylepis mlanjensis PEM R-17482 Malawi: Mt. Mulanje: Chilemba Peak
69
100 Trachylepis mlanjensis PEM R-17484 Malawi: Mt. Mulanje: Chilemba Peak
Trachylepis mlanjensis PEM R-17480 Malawi: Mt. Mulanje: Chilemba Peak
Trachylepis laevis CAS 263582: Namibe: Serra da Neve
100 Trachylepis laevis INBAC/AMB 10540 Angola: Namibe: Virei Calundolo
100 Trachylepis laevis MGR 301 Namibia: Kunene: Otjitambi Guest Farm
100
Trachylepis laevis MGR 300 Namibia: Kunene: Otjitambi Guest Farm
100 Trachylepis ansorgii MUHNAC:MB 03-001385 Angola: Malanje: Lauca Dam
Trachylepis ansorgii MUHNAC:MB 03-001384 Angola: Malanje: Lauca Dam
100
Trachylepis ansorgii UF 187314 Angola: Namibe: Serra da Neve
99 100 Trachylepis ansorgii CAS 263427 Angola: Huila: Caconda
su 100 97
Trachylepis ansorgii PEM R-21737 Angola: Huila
Trachylepis ansorgii MUHNAC:MB 03-001394 Angola: Huambo: Kanjonde
100
Trachylepis sulcata CAS 263485 Angola: Namibe: Virei Calundolo
97
Trachylepis sulcata CAS 263342 Angola: Huila: Tundavala
98
100
Trachylepis sulcata MCZ A-28869 Namibia: Hardap: Hardapdam
Trachylepis sulcata MCZ A-28324 Namibia: Erongo: Farm Omandumba
100
99
Trachylepis sulcata CAS 223979 Namibia: Kunene
Trachylepis sulcata AMB 8033 Namibia: Osmusati: Ruacana
Trachylepis ovahelelo, sp. nov. CAS 263493 Angola: Namibe
62 Trachylepis vunongue, sp. nov. MUHNAC:MB 03-001520 Angola: Cuatir
59 100
Trachylepis hilariae, sp. nov. CAS 254770 Angola: Namibe
90
54
Trachylepis hilariae, sp. nov. CAS 254769 Angola: Namibe
100 Trachylepis variegata MCZ R-184365 Namibia: Karas: Savanna Guest Farm
100 Trachylepis variegata CAS 200019 South Africa: Northern Cape: Richtersveld NP
100
Trachylepis punctulata PEM R-24137 Angola: Namibe: Farm Mucongo
Trachylepis punctulata CAS 263508 Angola: Namibe: Virei
100
100
Trachylepis punctulata PEM R-18001 Angola: Namibe
96 Trachylepis punctulata PEM R-17998 Angola: Namibe
99
Trachylepis cf. triebneri MCZ R-183759 Namibia: Erongo
Trachylepis cf. triebneri MCZ A-27711 Namibia: Kunene: Opuwo
59 58 Trachylepis cf. triebneri MCZ A-28045 Namibia: Erongo: vic. Brandberg
94 77
Trachylepis cf. triebneri MCZ Z-37900 Namibia: Kunene
Trachylepis cf. triebneri MCZ A-27750 Namibia: Kunene: Kamanjab
vg 100 Trachylepis bayonii MUHNAC:MB 03-001404 Angola: Huambo: Kanjonde
96
Trachylepis bayonii INBAC/JVV 9273 Angola: Malanje: Cangandala NP
99 Trachylepis bayonii PEM R-21822 Angola: Bie
100
Trachylepis huilensis PEM R-17963 Angola: Namibe
Trachylepis huilensis CAS 263565 Angola: Namibe: Serra da Neve
100
100
Trachylepis huilensis AG 201 Angola: Huila: Tundavala
90
Trachylepis huilensis CAS 263351 Angola: Huila: Tundavala
99
Trachylepis bouri, sp. nov. PEM R-24097 Angola: Namibe: Chicambi
100 Trachylepis bouri, sp. nov. CAS 354903 Angola: Namibe
96
Trachylepis bouri, sp. nov. PEM R-21727 Angola: Namibe st = striata group
Trachylepis bouri, sp. nov. CAS 263534 Angola: Namibe: Caraculo
100
Trachylepis brauni PEM R-20128 Tanzania: Iringa: Kitulo Plateau su = sulcata group
Trachylepis chimbana WRB WK 1 Angola: Sumbe
84
99
99
Trachylepis chimbana CAS 263543 Angola: Namibe: Dolondolo vg = variegata group
Trachylepis chimbana CAS 263563 Angola: Namibe: Serra da Neve
54
99 Trachylepis bocagii CAS 263436 Angola: Bengo: Luanda
100 Trachylepis bocagii MCZ A-36456 Angola: Bengo: Luanda
Trachylepis bocagii CAS 263434 Angola: Luanda: Kissama 0.1 substitutions/site
63 Trachylepis bocagii PEM R-21735 Angola: Bengo: Kwanza River
65
Trachylepis bocagii MUHNAC:MB 03-001341 Angola: Malanje: Lauca Dam
In our ML phylogeny (fig. 1), most lineages Species Concept

were strongly supported and most species were
Combining the morphological and molecu-
recovered as monophyletic. However, nine tra-
lar evidence presented above, we adopt the gen-
ditionally recognized species were recovered as
eral lineage species concept (de Queiroz, 1999)
paraphyletic, including Trachylepis brevicollis,
and recognize the 25 species of Trachylepis
T. comorensis, T. elegans, T. gravenhorstii, T.
occurring in Angola.
maculilabris, T. punctatissima, T. spilogaster,
T. striata, and T. wahlbergii. With respect to
new species that we describe in the Taxo- TAXONOMIC ACCOUNTS
nomic Accounts section, phylogenetic analyses
strongly supported Trachylepis attenboroughi, Trachylepis albilabris (Hallowell, 1857)—
sp. nov., as sister to T. spilogaster (in part.). Guinea Skink
Trachylepis wilsoni, sp. nov., was recovered sis- Figures 2, 3, plate 1
ter to Trachylepis suzanae, sp. nov., with strong
support, and these two species were together Euprepes albilabris Hallowell, 1857: 51. Holo-
recovered as the sister lineage to T. acutilabris. type: ANSP (specimen number unknown,
Trachylepis bouri, sp. nov., was strongly sup- lost fide Uetz et al., 2022) collected by Ford.
ported and recovered as the sister lineage to Type locality: “Gaboon.”
the lineage comprising T. bocagii, T. brauni,
Euprepes Blandingii: Bocage (1866a: 44); Peters
and T. chimbana. Trachylepis ovahelelo, sp.
(1877: 614).
nov., Trachylepis vunongue, sp. nov., and Tra-
Mabuia Raddonii: Bocage (1895: 40).
chylepis hilariae, sp. nov., are monophyletic
Mabuia raddonii: Boulenger (1887: 165).
groups, closely related to T. variegata, and the
Mabuya radoni: Ferreira (1903: 14); Parker
group composed by these four taxa is sister to
(1936: 138); Hellmich (1957b: 55).
the group composed by T. punctulata and T. cf.
triebneri. Some relationships depicted in figure Trachylepis affinis: Marques et al. (2018: 253);
1 should be treated with skepticism. We did Branch et al. (2019a: 318).
not examine the specimens associated with T. Trachylepis cf. affinis: Ernst et al. (2020: 242).
wrightii and T. seychellensis or the Madagascan
Previous records of this species in Angola have
members of the genus, so the identity of the
been attributed to “Euprepes blandingii,” “Mabuya
sequenced individuals is in question. Finally,
raddonii,” or Trachylepis affinis (see chresonymy
relationships within the T. striata group should
above). This is illustrative of the difficulties sur-
not be considered definitive, and a separate
rounding the taxonomy of the Trachylepis affinis
revision of this difficult group is in prepara-
species complex (which includes the synonyms T.
tion (Portik et al., in prep.).
raddonii and T. blandingii) and its resembling
regional congeners, which have already been high-
Morphology lighted by several authors, including Hoogmoed
(1974) and Allen et al. (2017, 2019). The confusion
Mensural and meristic data for the studied
between T. albilabris and T. affinis dates back to the
species are presented in table 2. The 25 Trachy
19th century, but the debate about the species iden-
lepis species occurring in Angola differ from
tity continued into the mid-20th century, with
each other and from other congeners in several
authors such as Boulenger (1887), Schmidt (1919),
consistent morphological characters. Detailed
Chabanaud (1921), Loveridge (1936), and Manaças
diagnoses for each species are provided in the
(1951) considering T. albilabris as a synonym of T.
taxonomic accounts below. Diagnoses are based
affinis (or of its current synonyms). Hoogmoed
only on the Angolan populations.
TABLE 2
2024
Morphological and Meristic Comparisons between Angolan Trachylepis
Abbreviations are listed in the Materials and Methods. Measurements are presented in millimeters and ratios as percentages (data presented as [Min.–Max.]).
Type of contact between head shields coded as follows:
in contact (C), in contact at a single point (SPC), separated (S), fused (F)
Trachylepis albilabris Trachylepis Trachylepis ansorgii Trachylepis Trachylepis bayonii Trachylepis binotata Trachylepis bocagii
(N = 13) albopunctata (N = 26) attenboroughi, sp. (N = 10) (N = 11) (N = 12)
(N = 41) nov. (N = 24)
SVL 56.2–75.8 31.2–62.5 41.7–98.9 51.4–79.4 50.2–83.0 56.5–134.3 35.7–76.3
TL 78.0–131.8 61.4–111.4 70.2–159.1 64.2–111.9 79.1–131.3 65.0–123.0 52.7–92.1
HW 7.5–10.6 5.3–8.3 6.4–15.0 8.0–12.1 7.4–11.3 9.0–25.3 5.6–9.9
HH 6.2–8.2 3.2–6.2 3.9–10.6 4.8–8.6 5.8–8.3 6.7–17.9 3.7–7.4
HL 12.0–15.4 8.0–13.2 10.6–20.3 13.1–19.4 10.4–15.6 13.5–31.3 9.9–14.9
IN 1.8–3.0 0.8–2.4 1.4–4.3 1.8–2.6 1.5–2.5 2.0–4.5 1.3–2.3
EN 3.0–5.0 1.8–4.3 3.3–7.6 2.8–5.8 2.6–4.6 5.0–10.7 2.2–4.8
ES 4.2–5.8 3.3–5.6 4.0–9.6 4.5–7.3 4.0–6.5 5.8–14.2 3.2–6.1
TL/SVL 105–215 106–204 102–178 94–162 123–193 87–138 109–154
HL/SVL 19–22 17–26 18–25 18–27 14–25 21–26 17–28
ES/HL 30–41 49–70 35–52 30–46 32–54 35–48 32–45
HW/HL 63–72 57–80 48–85 58–81 60–96 59–82 57–77
LUFT 13–15 17–26 18–24 17–22 15–16 17–22 21–27
LUFF 9–13 13–18 13–18 13–18 12–13 14–17 13–20
MSR 31–36 31–38 38–45 34–40 28–35 34–42 36–41
SAD 46–48 41–49 50–55 52–57 50–56 51–60 55–61
SAV 47–54 50–59 54–64 53–61 58–61 59–72 58–67
KDS 3–4 3–4 3–6 3–4 3–7 3–5 3–5
CERÍACO ET AL.: TRACHYLEPIS (SQUAMATA: SCINCIDAE) OF ANGOLA
SC 6–7 5–6 5 5 4–5 5–6 4–6

SL 7 7–8 7–8 8 (rarely 7) 8 (rarely 9) 8 (rarely 7) 8 (rarely 9)
PS Smooth Spinose Spinose Spinose Smooth Smooth Spinose
11
12
TABLE 2 continued
Trachylepis albila- Trachylepis Trachylepis ansorgii Trachylepis Trachylepis bayonii Trachylepis binotata Trachylepis bocagii
bris (N = 13) albopunctata (N = 26) attenboroughi, sp. (N = 10) (N = 11) (N = 12)
(N = 41) nov. (N = 24)
CP Usually S (N = 9) or Usually C (N = 29) Usually S (N = 9) or Usually C (N = 18) Usually S (N = 8), Usually S (N = 8), Usually C (N = 11),
SPC (N = 3), rarely or SPC (N = 2), SPC (N = 16), rarely or SPC (N = 3), rarely C (N = 1) or rarely SPC (N = 2) rarely SPC (N = 1)
C (N = 1) sometimes S (N = 8) C (N = 1) rarely S (N = 3) SPC (N = 1)
CFP Always C Always C Always C Always C, rarely Always F Always C Always C
SPC (N = 1)
CSN Usually S (N = 12), Usually C (N = 24) Usually C (N = 25) Always C, rarely Always C Always C Always C
rarely SPC (N = 1) or SPC (N = 4), or SPC (N = 1) SPC (N = 1)
sometimes S (N =
10)
CPF Always C Usually S (N = 35) Usually S (N = 20) Usually S (N = 17) Usually S (N = 8), Usually C (N = 10), Usually S (N = 9),
rarely C (N = 1) or or SPC (N = 3), or SPC (N = 4), rarely C (N = 2) rarely S (N = 1) SPC (N = 3)
SPC (N = 2) sometimes C (N = sometimes C (N =
3) 3)
TABLE 2
(continued)
Trachylepis bouri, sp. Trachylepis chimbana Treachylepis Trachylepis hilariae, Trachylepis hoeschi Trachylepis huilensis Trachylepis laevis
nov. (N = 6) (N = 16) damarana (N = 12) sp. nov. (N = 4) (N = 3) (N = 10) (N = 7)
SVL 40.5–56.3 34.5–56.0 45.6–56.2 32.8–43.1 59.1–76.5 42.0–60.4 40.6–63.0
TL 40.5–54.0 44.1–80.1 72.1–93.0 47.0–55.3 105.9–114.0 65.0–89.0 65.6–100.0
HW 5.6–8.5 5.2–8.1 6.3–8.2 4.7–5.9 10.6–13.5 6.2–8.6 5.7–9.7
HH 4.1–6.4 3.5–6.1 4.2–5.7 3.3–4.1 7.0–8.6 4.6–6.3 2.9–4.2
HL 9.2–13.7 8.3–13.3 9.3–11.5 7.9–8.9 14.4–18.4 9.4–12.9 9.9–15.2
IN 1.2–1.9 1.2–1.8 1.5–1.8 1.0–1.2 1.8–2.3 1.4–1.8 0.8–2.2
BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
EN 3.5–4.6 2.4–3.7 3.4–6.8 2.4–2.7 4.9–5.6 2.5–3.9 2.4–3.9

ES 4.1–6.8 3.2–5.6 4.8–5.7 3.3–3.6 6.0–7.4 4.1–5.2 4.1–5.8
TL/SVL 132–176 102–173 150–184 109–162 145–179 112–172 119–202
HL/SVL 22–26 18–25 20–22 21–25 24–25 20–25 21–25
ES/HL 40–50 33–50 47–55 40–43 37–42 35–45 35–45
HW/HL 59–66 61–76 62–71 58–66 69–74 60–73 58–66
NO. 465
TABLE 2 continued
Trachylepis bouri, sp. Trachylepis chimbana Treachylepis Trachylepis hilariae, Trachylepis hoeschi Trachylepis huilensis Trachylepis laevis 2024
nov. (N = 6) (N = 16) damarana (N = 12) sp. nov. (N = 4) (N = 3) (N = 10) (N = 7)
LUFT 21–23 18–24 14–17 21–22 18–19 16–18 19–22
LUFF 15–17 15–17 19–24 15–16 13–15 12–14 15–16
MSR 36–38 33–39 31–34 29–30 33 34–39 30–33
SAD 50–54 49–56 40–43 46–48 43–48 51–56 49–53
SAV 54–60 55–63 50–55 58 55–62 55–66 56–57
KDS 5 4–5 3–4 5 3 3–5 Smooth
SCL 5 5 4–5 5 5–6 3–4 4–5
SL 8–9 8 (rarely 9) 7 8 10 8 9 (rarely 8)
PS Spinose Spinose Spinose Spinose Smooth Smooth Smooth
CP Always C Usually C (N = 12) or Always SPC Always C Always S Usually C (N = 7), Always C
SPC (N = 3), rarely S sometimes SPC
(N = 1) (N = 3)
CFP Always C Always C Always C Always C Always C Always C Always C
CSN Always C Always C Usually C (N = 10) Always C C (N = 2) or SPC Always C Always S
or SPC (N = 2), (N = 1)
CPF Usually S (N = 5), Always S Usually SPC (N = 9) Always S Always S Always S Always S
rarely C (N = 1) or C (N = 1), rarely
S (N = 2)
TABLE 2
(continued)
Trachylepis maculilabris Trachylepis notabilis Trachylepis Trachylepis ovahelelo, Trachylepis punctulata Trachylepis Trachylepis sulcata
(N = 20) (N = 6) occidentalis (N = 5) sp. nov. (N = 1) (N = 20) raymondlaurenti (N = 17)a (N = 41)
SVL 44.2–91.9 45.1–97.9 44.4–110.7 37.3 35.7–48.5 46.9–80.0 39.5–87.9
TL 89.9–201.3 91.0–189.2 57.2–185.0 70.9 50.0–78.0 126.1–215.8 51.2–150.0
HW 6.1–15.0 7.6–16.3 7.7–17.7 5.8 4.5–7.0 5.9–8.6 6.6–14.7

HH 4.4–10.6 5.3–11.4 4.9–12.0 3.3 3.2–5.1 4.8–7.0 3.9–10.1
HL 9.0–21.3 11.2–22.4 11.7–25.9 8.0 8.8–12.2 9.2–13.1 9.7–20.2
IN 1.6–3.0 2.3–3.7 1.6–3.4 0.9 1.0–2.0 1.8–2.6 1.4–3.5
13
EN 2.3–5.2 3.0–6.5 3.6–6.2 2.5 2.5–3.4 2.5–3.7 2.6–6.1

14
TABLE 2 continued
Trachylepis maculilabris Trachylepis notabilis Trachylepis Trachylepis ovahelelo, Trachylepis punctulata Trachylepis Trachylepis sulcata
(N = 20) (N = 6) occidentalis (N = 5) sp. nov. (N = 1) (N = 20) raymondlaurenti (N = 17)a (N = 41)
ES 3.4–7.6 4.4–10.1 5.0–9.6 3.2 3.2–4.8 4.1–5.7 3.6–8.9
TL/ 135–249 109–195 77–183 190 128–184 192–331 99–193
SVL
HL/ 16–25 20–25 22–26 21 21–27 16–20 19–28
SVL
ES/HL 28–47 35–50 33–44 40 34–47 19–32 25–39
HW/ 56–82 64–81 66–74 73 51–75 62–69 55–92
HL
LUFT 12–19 13–18 18–24 18 21–24 15–17 20–27
LUFF 10–15 11–15 13–16 14 12–18 10–13 16–21
MSR 32–40 31–38 30–34 34 28–35 24–28 33–42
SAD 47–62 49–59 45–50 43 42–50 48–53 48–58
SAV 50–66 50–60 56–58 53 46–60 50–58 57–68
KDS 3–6 4–5 3 5 3–5 Smooth 3–7 (usually 5)
SCL 4–7 5–6 5 5 4–6 5 4–6
SL 7 7 (rarely 8) 7 (rarely 10) 7 8 (rarely 7) 7–8 6–9
PS Smooth Smooth Smooth Spinose Spinose Smooth Spinose
CP Usually C (N = 16) or Usually S (N = 4), S (N = 3) or SPC C Usually C (N = 17) or Usually S (N = 15), Usually C (N =
SPC (N = 3), rarely S rarely C (N = 2) (N = 2) SPC (N = 3) rarely C (N = 1) or SPC 15) or SPC (N =
(N = 1) (N = 1) 16), sometimes S
(N = 9)
CFP Always C Always C Always C C Always C Always C Always C
BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
CSN Usually C (N = 7) or Usually C (N = 4) Always C C Always C Usually S (N = 16), Usually C, rarely S

SPC (N = 9), rarely S or SPC (N = 2) rarely C (N = 1) (N = 4)
(N = 4)
CPF Usually C (N = 13) or Always C Usually S (N = 4), S Always S Usually S (N = 12b), Usually S, rarely C
SPC (N = 3), rarely S rarely C (N = 1) sometimes C (N = 5) (N = 6)
(N = 4)
aData from Marques et al. (2019).
bSeparated by additional median scale in two specimens.
NO. 465
TABLE 2 continued
TABLE 2 2024
(continued)
Trachylepis suzanae, sp. nov. (N = 16) Trachylepis vunongue, sp. nov. (N = 10) c Trachylepis wahlbergii (N = 20) Trachylepis wilsoni, sp. nov. (N = 26)
SVL 41.6–62.25 max. 46.0 31.6–85.8 38.6–57.3
TL 60.8–110.3 max 63.0 47.5–113.7 60.3–91.1
HW 6.2–9.8 5.4–6.2 5.6–13.0 6.0–8.7
HH 4.5–7.4 3.9–5.0 3.7–9.5 4.3–6.1
HL 10.6–15.1 8.1–9.4 8.3–19.0 8.2–12.6
IN 1.5–2.4 0.8–1.5 1.3–3.0 1.2–2.5
EN 3.5–4.8 2.0–3.0 2.4–5.7 2.2–4.8
ES 4.6–6.7 3.4–3.9 3.2–7.6 3.5–5.3
TL/SVL 146–167 111–152 117–155 109–197
HL/SVL 25–27 20–24 19–27 21–25
ES/HL 39–43 37–43 34–49 38–49
HW/HL 58–70 63–70 56–78 58–76
LUFT 19–26 15–22 15–18 22–26
LUFF 12–15 9–14 12–15 15–17
MSR 28–31 30–35 35–40 29–33
SAD 43–49 40–52 46–55 48–52
SAV 48–57 45–55 53–63 52–57
KDS 3–4 5 3–4 2–4
SCL 4–7 4–6 4–7 5
SL 7 (rarely 6) 7–8 (rarely 9) 8 (rarely 7 or 9) 8
PS Spinose Spinose Spinose Spinose
CP Usually C C or SPC Usually S (N = 14), rarely C (N = 1) Usually C (N = 21), rarely S (N = 1)
or SPC (N = 1)
CFP C C Always C Always C
CSN Usually C C Always C Always C

CPF Usually S S Usually S (N = 11) or SPC (N = 4), Always S
rarely C (N = 1)
cData from Conradie et al. (2022) and the type series, including non-Angolan material.
15
FIGURE 2. Specimen of Trachylepis albilabris from Cabiri, Bengo Province (MUHNAC/MB03 1374). Photos
by L.M.P.C.
(1974) was the first to consider T. albilabris as a (this paper) and Uíge provinces (Ernst et al., 2020;
valid species, different from the T. affinis species this paper), where the habitat presents a southern
complex. This was recently confirmed by Allen et continuation of the Congolese habitats and biomes.
al. (2017) and Weinell et al. (2019) through molec- Diagnosis: A medium-sized skink (max. SVL
ular data. Trachylepis albilabris was originally 75.8 mm, MUHNAC/MB03-001371), with fully
described from “Gaboon” (Hallowell, 1857), and its developed, pentadactyl limbs (figs. 2, 3); dorsal
distribution ranges from Guinea eastward to scales tricarinate to quadricarinate; ventral scales
Uganda and southward to Angola (Hoogmooed, smooth; 47–54 SAV; 46–48 SAD [43–49 in extra-
1974; Kingdon and Spawls, 2010; Allen et al., 2017, limital populations; see Hoogmoed, 1974]; 31–36
2019; this paper). Our molecular and morphologi- MSR [27–32 in extralimital populations; see Allen
cal data allow us to correct the historical misiden- et al., 2019]; lamellae beneath fingers and toes
tification of this species in Angola. Trachylepis smooth; plantar scales smooth; 13–15 LUFT [up to
albilabris is a representative of the central/western 18 in extralimital populations; see Hoogmoed,
African clade of the genus (Weinell et al., 2019), 1974], 9–13 LUFF [up to 15 in extralimital popula-
historically recorded in the northwestern areas of tions; see Hoogmoed, 1974]; supranasals usually
the country by several authors (Peters, 1877; separated; parietals usually separated; prefrontals
Bocage, 1895; Ferreira, 1903; Parker, 1936; always in contact; frontoparietals always in contact;
Hellmich, 1957b) and recently collected in Bengo one pair of enlarged nuchal scales present; ear open-
FIGURE 3. Life photo of Trachylepis albilabris from Serra Pingano, Uíge Province (MUHNAC/MB03-001369).
Photo by L.M.P.C.
ing vertically ovoid and smaller than the eye; lacking farms [-7.6729°, 14.9339°, 631 m] (MUHNAC/
subtriangular auricular scales on the anterior mar- MB03-001365–1370, 001521); Camp site near
gin of the ear opening. Seven SL, the fifth subocular; water pump, E of Ponte village [-7.6829°,
six to seven supracilliaries, the third usually longest; 14.9340°, 774 m] (MUHNAC/MB03-001371,
nostril oriented laterally. Dorsum olive-brown with 001372).
dark flecks; a dark brown lateral band running from Additional Material: Kwanza Sul Province:
the snout to the hind limb insertion, bordered Roça Canzele, 30 km W of Camabatela [-8.1830°,
below by a white stripe, followed by brownish col- 15.0940°, 718 m] (FMNH 74303, 74304, 74306, 74307,
oration on the lower flank. Top of head uniform 74309, 74311; MCZ R-110306); 30 km S of Gabela
[-11.0982°, 14.3342°, 1022 m] (FMNH 74310); Con-
brown; labials white to grayish, sometimes brownish
gulu [-10.8667°, 14.2833°, 639 m] (MCZ R-112221;
on the upper margin of supralabials. Scales of the BMNH 1936.8.1.601–611). Malanje Province: 16 km
venter yellowish or white, without markings. Some W of Lucala [-8.9116°, 15.8044°, 1167 m] (TM 45495).
specimens present dark speckling on the chin and Uíge Province: University Campus, Kimpa Vita
gular area. There is often a yellowish to orange patch [-7.6176°, 15.0654°, 835 m] (MTD 48611, 48928,
near the forelimb insertion. 48931, 49784).
Material Examined: Bengo Province: Historical Localities (no extant
dirt road to Cabiri, in a small, forested area on specimens): Cabinda Province: “Chinchoxo”
the left side of the road [-8.8710°, 13.6057°, (côte d’Loango) [-5.1000°, 12.1000°, 45 m]
13.5 m] (MUNHAC/MB03-001373–001376). (Peters, 1877; Bocage, 1895). Huíla Province:
Uíge Province: Ponte village, roads through Caconda [-13.3333°, 15.0667°, 1644 m]
FIGURE 4. Distribution of Trachylepis albilabris in Angola.
(Bocage, 1895). Kwanza Norte Province: Habitat and Natural History Notes: The
“Piri-Dembos” [-8.5299°, 14.4377°, 712 m] species is found in humid habitats, with mosaics
(Hellmich, 1957b). Malanje Province: Duque of forest and savannah (Grandvaux-Barbosa,
de Bragança [-9.0740°, 15.9999°, 1065 m] 1970; fig. 5). Ernst et al. (2020) reported that the
(Ferreira, 1903). species was frequently associated with degraded
Distribution in Angola: The species is dis- forest or forest-edge habitats in Serra Pingano,
tributed in the northwestern regions of Angola, Uíge Province. This observation is in line with our
from the Cabinda enclave southward to Huíla own recent observations of the species in Cabiri
Province, along the more forested areas and and Serra Pingano. The species is usually sympat-
escarpment regions (fig. 4). ric with T. maculilabris. It is quite active during
Global Distribution: A central African the day, sometimes seen basking on tree trunks
species, occurring from Gabon southward to and shrubby vegetation. In the localities where it
Angola. exists, it is very common and abundant.
FIGURE 5. Typical habitat of Trachylepis albilabris in Serra Pingano, Uíge Province. Photo by L.M.P.C.
Trachylepis albopunctata (Bocage, 1867)— National Park, Malanje Province, central

Angolan Variable Skink Angola [-9.8546°, 16.7100°, 1104 m].
Mabuia varia: Boulenger (1887: 202; 1905: 111);
Figures 6, 7, plate 1
Bocage (1895: 43, 1896: 111).
Euprepes Oliveiri var. albo-punctatus Bocage Mabuya acutilabris [part]: Schmidt (1919: 551).
1867a: 223. Neotype [here designated]: Mabuya varia [part]: Schmidt (1933: 12);
An unsexed adult (CAS 258379, field number Monard (1937: 87); Mertens (1938: 437);
JVV 9180), collected by Luis M.P. Ceríaco, Parker (1936: 128); Laurent (1964: 72);
Mariana P. Marques, Suzana A. Bandeira, Branch (1998: 157).
Jens V. Vindum, and Edward L. Stanley on 14 Mabuya varia varia: Hellmich (1957a: 66, 1957b:
September 2016. Type locality: Cangan- 57); Loveridge (1957: 212).
dala National Park, Malanje Province, central Trachylepis varia: Ceríaco et al. (2016b: 31;
Angola [-9.8546°, 16.7100°, 1104 m]. 2016c: 67); Conradie et al. (2016: 26).
Euprepes angolensis Bocage 1872: 78. Neotype Trachylepis varia Clade B: Weinell and Bauer
[here designated]: The same specimen as (2018: 107).
the neotype for Euprepes Oliveiri var. albo- Trachylepis cf. albopunctata: Marques et al.
punctatus. Type locality: Cangandala (2018: 254); Branch et al. (2019a: 319).
Trachylepis albopunctata: Butler et al. (2019: identifying the species as varia. Schmidt (1919),
233); Baptista et al. (2019: 109); Santos et al. however, confounded some specimens of this
(2021: 23); Conradie et al. (2022: 201). species from northern Angola with Mabuya acu-
Trachylepis varia complex: Ceríaco et al. (2020a: tilabris (now T. suzanae; see account below).
403). Only recently, based on a phylogenetic revision
of the T. varia species complex, Weinell and
Trachylepis albopunctata is probably one of Bauer (2018) showed evidence that the Angolan
the most widely distributed species of the genus populations likely constitute a distinct taxon,
in the country (Marques et al., 2018) but, at the suggesting that the names albopunctata and/or
same time, one of the species for which taxo- angolensis would possibly be available for the
nomic and nomenclatural history is most com- Angolan lineage. This led subsequent authors to
plex. Long considered part of the Trachylepis adopt the oldest nomen, albopunctata, as the
varia species complex, with which it was synony- applicable name for the Angolan representatives
mized by the majority of authors who historically of the T. varia complex (Marques et al., 2018;
dealt with it in Angola, the species was only Baptista et al., 2019; Branch et al., 2019a; Butler
recently reinstated to full species level by Weinell et al., 2019). However, as noted by Weinell and
and Bauer (2018). The species was originally Bauer (2018) and Marques et al. (2018), the situ-
described by Bocage (1867a) as a variety of ation is nomenclaturally unstable primarily
Euprepes olivieri (currently Heremites vittatus because of the loss of the type material of both
(Olivier, 1804)), based on “several specimens Bocage’s albopunctata and angolensis, and the
from Benguella and Catumbella” (currently Ben- secondary homonymy of Mabuia angolensis,
guela and Catumbela towns, respectively), Ben- erected by Monard (1937) to describe a putative
guela Province. The author provided a detailed new species within the T. striata/wahlbergii spe-
(by mid-19th century standards) description of cies complex (see account of T. monardi, comb.
the species, focusing especially on coloration nov., in Marques et al., 2018, and that of T. wahl-
pattern (Bocage, 1867a). A few years later, bergii in this paper for further details). The only
Bocage (1872) described Euprepes angolensis sensible solution to stabilize the nomenclature of
based on two specimens from “Biballa” and three the group is to designate neotypes for both
specimens from “Dondo,” currently Bibala and nomina—albopunctata and angolensis. Following
Dondo in Namibe and Kwanza Norte provinces, the original opinion of Bocage (1895) and con-
respectively. This description was considerably sidering that the distribution of our samples
more detailed than the former, but, in general its across Angola largely corresponds to the type
characters agreed entirely with his own E. o. var. localities of both nomina, we opt to designate the
albo-punctatus and, in Bocage’s own words, was same neotype for both albopunctata and angolen-
similar to “E. damaranus” (currently Trachylepis sis, rendering the latter an objective junior syn-
damarana). In his major revision of Angolan onym of the former.
herpetofauna, Bocage (1895) opted to synony- Diagnosis: A medium-sized skink (max. SVL
mize both albo-punctatus and angolensis with 62.5 mm, CAS 258389; max. SVL of extralimital
Mabuia (currently Trachylepis) varia. For more populations 65 mm, see Weinell and Bauer, 2018),
than 120 years, all subsequent authors that have with fully developed, pentadactyl limbs (figs. 6, 7);
dealt with this species in Angola (Bocage, 1896; dorsal scales tricarinate or quadricarinate; ventral
Schmidt, 1933; Parker, 1936; Monard, 1937; scales smooth; 50–59 SAV; 41–49 SAD; 31–38
Mertens, 1938; Hellmich, 1957a, 1957b; Lov- MSR; lamellae beneath fingers and toes keeled
eridge, 1957; Laurent, 1964; Branch, 1998; and spinose; plantar scales spinose; 17–26 LUFT,
Ceríaco et al., 2016a, 2016b; Conradie et al., 13–18 LUFF; supranasals usually in contact, but
2016) followed the decision of Bocage (1895) in sometimes separated; parietals usually in contact,
FIGURE 6. Neotype of Trachylepis albopunctata from Cangandala National Park, Malanje Province (CAS
258379). Photos by L.M.P.C.
but sometimes separated; prefrontals usually sepa- nostril and usually becomes more reticulated pos-
rated, rarely in contact; frontoparietals always in teriorly, often forming irregular blotches or verti-
contact; one pair of enlarged nuchal scales pres- cal bars on the upper flanks. Limbs variegated
ent; ear opening vertically ovoid and smaller than above, with irregular dark and pale speckles. Top
the eye; 2–4 subtriangular auricular scales (shorter of head uniform brown or with diffuse dark stip-
than the diameter of ear) extend posteriorly and pling; labials white, sometimes with dark vertical
usually slightly upward from the anterior margin bars on the posterior margin. Venter white to blu-
of the ear opening. Supralabials usually eight ish without markings.
(rarely seven), the sixth being subocular; usually Neotype [for both Euprepes Oliveiri var.
five supracilliaries, the second notably longer; albo-punctatus Bocage, 1867, and Euprepes
nostril oriented laterally. Dorsum olive brown, angolensis Bocage, 1872]: An unsexed adult
with a pair of faint pale dorsolateral stripes; (CAS 258379, field number JVV 9180; fig. 6) col-
between the stripes there is usually heavy white lected at Cangandala National Park, Malanje
and black speckling, the latter with tendency to Province, central Angola [-9.8546°, 16.7100°,
form transverse bars starting behind the neck and 1104 m], by Luis M.P. Ceríaco, Mariana P.
extending through the tail; there may also be an Marques, Suzana A. Bandeira, Jens V. Vindum,
indistinct pale vertebral stripe. A bright white lat- and Edward L. Stanley on 14 September 2016.
eral stripe, usually bordered with black, starts at Description of the Neotype: Well-pre-
the subocular and extends through the flank to served, unsexed adult. Body cylindrical and
the hind limb insertion; a black stripe starts at the robust with a poorly defined neck and well-
FIGURE 7. Life photo of Trachylepis albopunctata from Bicuar National Park, Huíla Province (CAS 263387).
Photo by L.M.P.C.
developed pentadactyl limbs; tail long and parietal and interparietal. Frontoparietal plus
robust, its length slightly larger than SVL, interparietal length greater than frontal length.
smoothly tapered. Fore- and hind limbs over- Interparietal twice as long as broad, with a
lap when adpressed against the body. SVL 60.5 visible parietal foramen. Parietals larger than
mm, TL 65.7 mm, HL 12.7 mm. Additional frontoparietals. Parietals in contact with each
measurements are presented in table 3. Ear other. Five supraciliaries, second largest. Eight
opening small, with three anterior subtriangular supralabials, sixth subocular. Eight infralabials.
auricular scales. Rostral visible from above. Nos- Postmental bordering seven scales (mental, two
trils set posteriorly, so that postnasal effectively infralabials on each side and two primary chin
borders nostril. Supranasals in contact. Fronto- shields). Transparent scale present in lower eye-
nasal broader than long, in contact with loreal lid, as is usual for Trachylepis. Tympanum vis-
scale. Prefrontals pentagonal, separated from ible, at same level as mouth. Dorsal scales each
one another, each in contact with the following with three smooth keels. Ventral scales smooth.
head shields: frontonasal, loreal, first and second MSR 35, SAD 49, SAV 54. Limbs with five digits;
supraocular, first supraciliary and frontal. Two scales on palms soles spinose. Relative length of
loreals. Frontal longer than the distance between fingers IV > III > V > II > I, relative length of
anterior tip of frontal and tip of snout. Frontal toes IV > III > V > II > I. Finger-IV lamellae 17,
in contact with three supraoculars on each side. Toe-IV lamellae 20.
Two frontoparietals, in contact with each other Coloration in Ethanol: Background color of
and the frontal, third and fourth supraoculars, flanks and upper side of head, neck, dorsum, and
TABLE 3
Mensural and Meristic Data for the Examined Trachylepis Neotypes and Syntypes
Abbreviations are listed in the Materials and Methods. Measurements are presented in millimeters
and ratios as percentages
Trachylepis Trachylepis Trachylepis Trachylepis Trachylepis

albopunctata bayonii bocagii maculilabris [Euprepes notabilis
[Euprepes angolensis] anchietae]
CAS 258379 ZMB 6477 ZMB 6479 MUHNAC/MB03- ZMB 9204
Neotype Syntype Syntype 1390 Neotype Lectotype
Sex unsexed unsexed unsexed unsexed unsexed

SVL 60.5 65.6 68.8 79.9 84.4
TL 65.7 88.7 – 185 165
TL/SVL 110 140 – 230 190
HL 12.7 12.7 13.5 17.8 18.2
HL/SVL 20 20 20 20 20
SVL/HL 480 520 510 450 460
HW 8.2 8.8 9.4 11.2 12.2
HW/HL 60 70 70 60 70
HH 5.5 6.9 7.4 8.9 10.9
IN 1.8 1.6 1.5 3 2.6
EN 3.7 4.4 4.4 4.5 5.8
ES 5.6 5.2 6.1 6.3 8
MSR 35 35 36 32 32
SAD 49 55 57 57 59
SAV 55 61 62 57 60
LUFF 17 12 17 14 15
LUFT 20 15 24 16 18
SC 5 4 4 5 6
SL (SO) 8 (6) 8 7 7 (3) 7 (5)
CP C S C C S
CFP C F C C C
CSN C C C S C
CPF S S S C C
KDS 3 3–4 3–4 5 5
Plantar spinose smooth keeled smooth smooth
scales
tail olive brown, with scattered black and white crosses the eye and becomes fainter and reticulated
speckles forming series of transverse bars starting posteriorly; labials cream white; a bright white
behind the nape and extending to the tip of the tail; stripe starts at the subocular and extends to the
limbs variegated above. There is a pair of faint pale hind limb insertion. Venter white with some gray-
dorsolateral stripes; a black stripe on the snout ish speckling, especially near the flanks.
Material Examined: Bié Province: Chitau 13.2569°, 1585 m] (MUNHAC/MB03-001468,

[-11.4333°, 17.1500°, 1510 m] (CM 5750); Cas- 001469, 001486–001498, 001500–001502,
sumbi, camp established near Cassumbi village, 001504–001508); Serra da Neve base, 2km N of
14 km dirt road, rocky area [-11.0764°, 16.6628°, Maylowe [-13.8280°, 13.2625°, 818 m] (MUH-
1244 m] (CAS 266020; MUNHAC/MB03- NAC/MB03-001509); Serra da Neve, rocky area
001380); 14 km ENE from Mumbue village, dis- near base camp [-13.7653°, 13.2571°, 1645 m]
turbed miombo, walk margin of the river bed (MUNHAC/MB03-001499, 001503). Zaire
[-13.8653°, 17.4255°, 1543 m] (CAS 266038; Province: Noqui [-5.8750°, 13.4322°, 38 m]
MUHNAC/MB03-001466). Huambo Province: (AMNH R11095–96).
Kajonde village, Mt. Moco [-12.4294°, 15.1545°, Additional Material: Bengo Province: Ambriz
1890 m] (MUNHAC/MB03-001381, 001382). [-7.8612°, 13.1182°, 26 m] (BMNH 18511.1.19.3–6). Bié
Huíla Province: Bicuar National Park, on a Province: Bihe (= Bié) [-12.3833°, 16.9500°, 1660 m]
dam wall [-15.1271°, 14.7715°, 1246 m] (CAS (FMNH 18511–18517); Chitau [-11.4333°, 17.1500°, 1510
263398, 263399); Bicuar National Park, camp m] (AMNH 49012; CM 5671, 5883; NMB 13228, 13229);
headquarters [-15.1016°, 14.8399°, 1224 m] 10 km west of Cuemba village [-12.0348°, 18.0487°, 1437
m] (PEM R23344–5); Stop 2: road to Cuito River source
(CAS 263387, 263388, 263389, 263390, 263391;
[-12.2823°, 18.6291°, 1487 m] (PEM R23355); EN140
UF 187324); Bicuar National Park, E of camp North of Menongue [-13.8470°, 17.2531°, 1503 m]
[-15.1028°, 14.8477°, 1243 m] (UF 187325); (PEM R23543). Cuando Cubango Province: south of
Bicuar National Park, on the rd to Nongalafa, ca Menongue en route to Cuebe River [-14.9628°, 17.6909°,
8 km W of camp headquarters [-15.1274°, 1300 m] (PEM R23256–8); Cuchi River gorge [-14.5900°,
14.7714°, 1225 m] (CAS 263392); Lubango, 16.9076°, 1350 m] (PEM R23265, INBAC no number
Nossa Senhora do Monte [-14.9434°, 13.4644°, fide Conradie et al., 2022). Cunene Province: Kuve-
1983 m] (CAS 263408, 263409; UF 187326, lai (= Cuvelai) [-15.6500°, 15.8000°, 1217 m] (MHNG
1545.013, 1545.014). Huambo Province: Mt. Moco
187327); Lubango, Cristo Rei [-14.9401°,
[-12.4167°, 15.1833°, 2393 m] (BMNH 1936.8.1.592);
13.5117°, 2200 m] (CAS 263410); Huíla water-
Cubango River campsite 2 near mission [-13.3289°,
fall [-15.0549°, 13.5349°, 1658 m] (CAS 263406). 16.4117°, 1520 m] (PEM R23389; INBAC/WC-5207);
Malanje Province: Pungo Andongo near foot- Cubango River, campsite 1 below rapids, west of Fundo
prints of the Queen [-9.6755°, 15.5836°, 1107 village [-13.0448°, 16.3750°, 1557 m] (PEM R23390).
m] (UF 185744); Kalandula Falls, road down to Huíla Province: 15 km N Quilengues [-13.9526°,
the corn field from the hotel [-9.0755°, 16.0112°, 14.0470°, 939 m] (MD 1970.1); Capelonga (=Capelongo)
1063 m] (UF 185747); Kalandula Falls, road [-14.9167°, 15.0833°, 1220 m] (AMNH 49013–49018);
outside the hotel [-9.0825°, 16.0121°, 1049 m] Kuwanga (currently Kuvango) [-14.4667°, 16.3000°, 1453
m] (MHNG 856.006); Caconda [-13.7333°, 15.0667°,
(UF 185746); Pungo Andongo near town
1674 ] (BMNH 1906.8.24.61; MNHN 1907.201; NHMW
grounds [-9.6673°, 15.5909°, 1155 m] (UF 16208); Humpata [-15.0251°, 13.3761°, 1872 m] (MD
185745); Cangandala National Park [-9.8276°, 1840.2) Stone bridge over Albul stream, 12 km North of
16.6699°, 1097 m] (CAS 258379, 258389); Laúca Humpata South West Angola [-14.9271°, 13.3359°, 2057
Dam, flooded area [-9.7627°, 15.1438°, 750 m] m] (PEM R17940); Christo Rei, Sada Bandeira (= Cristo
(MUNHAC/MB03-001242–001243; 001346– Rei, Sá da Bandeira (currently Lubango)) [-14.6667°,
001348); Gauca [-11.1833°, 17.4500°, 1163 ] 13.5137° 1365 m] (TM 40833); Hungueria [-15.3167°,
(CM 5950, 5952, 5953). Namibe Province: 13.5333°, 1297 m] (TM 40940); Sá da Bandeira (cur-
Serra da Neve [-13.7864°, 13.2575°, 1596 m] rently Luabango), Toco 16 km NE [-14.9167°, 13.5000°,
1751 m] (TM 45184); Leba Pass, between river and
(CAS 263560); dirt road to Quilenges [-13.8159°,
highway [-15.0703°, 13.2438°, 1670 m] (CAS 254874,
13.3264°, 587 m] (MUNHAC/MB03-001383); 254884); Entre Rios [-13.0167°, 14.6333°, 1267 m] (ZSM
Serra da Neve, 2016 base camp [-13.7770°, a102/1953); Cubal [-13.0333°, 14.2500°, 921 m] (SMF
13.2591°, 1488 m] (MUNHAC/MB03-001384); 25404/15); Kuvango River hydro plant site [-14.3877°,
Serra da Neve, Catchi surroundings [-13.7620°, 16.2937°, 1429 m] (PEM R23379). Kwanza Sul Prov-
FIGURE 8. Distribution of Trachylepis albopunctata in Angola. Black star denotes the newly designated neo-
type locality.
ince: Namba [-11.9167°, 14.8500°, 1783 m] (TM 46643). Pungo-Andongo [-9.6667°, 15.5833°, 1220 m] (BMNH
Lunda Norte Province: Capaia (Capaceira) [-8.3333°, 1904.5.2.38–40); Gauca, 20 mi E of Dando (on Quanza
20.2000°, 1010 m] (NHMW 766). Lunda Sul Prov- River) [-11.1833°, 17.4500°, 1163 m] (CM 5730–5743,
ince: Alto Chicapa, Lunda [-10.9333°, 19.1500°, 1373 5746–5749–5764, 5941, 5942, 5944–5949, 5951, 5954–
m] (MCZ R-74118; MD 5466); Main track at turning 5960, 5962, 5963). Moxico Province: Sandando, 85
to Luanda, Mussuco diversion, Cuango river [-8.9833°, km east from Luso [-11.6167°, 20.6333°, 1177 m] (MD
18.0167°, 786 m] (PEM R13480); Ridge 18 km North of 5718); Quembo River source trap 4 [-13.1359°, 19.0471°,
Tarjua Falls, Cuango River [-9.1750°, 18.1333°, 996 m] 1369 m] (PEM R23479). Namibe Province: Serra da
(PEM R13486). Malanje Province: Pavalange [-11.3667°, Neve camp [-13.7770°, 13.2591°, 1488 m] (MUHNAC/
17.6167°, 1176 m] (ANSP 32191, 32193–3219); Duque MB03-001516; INBAC/LMPC 1265); dirt road to Qui-
de Bragança Falls (currently Kalandula Falls) [-9.1333°, lenges [-13.8159°, 13.3264°, 587 m] (CAS 266148);
16.0667°, 1010 m] (TM 45464, 45467, 45480, 45481); 8.5 km North of Rito [-16.6232°, 19.0535°, 1128 m]
Cangandala National Park [-9.7588°, 16.8023°, 1134 m] (PEM R20508, 20509). Undetermined locality: Angola
(CAS 258376–258378, 258380–258387, 258390–258393); (MHNC 91.0449).
FIGURE 9. Typical habitat of Trachylepis albopunctata in Cangandala National Park, Malanje Province. Photo
by L.M.P.C.
Historical Localities (no extant speci- Kwanza Norte Province: Ambaca [-9.2667°,
mens): Benguela Province: Cahata [-12.3500°, 15.1833°, 738 m] (Bocage, 1895); Dondo [-9.6833°,
14.8167°, 1202 m] (Bocage, 1895); Catumbela 14.4333°, 33 m] (Bocage, 1872, 1895). Namibe
[-12.4333°, 13.5000°, 7 m] (Bocage, 1867a; Lov- Province: Biballa [-14.7667°, 13.3667°, 955 m]
eridge 1957); Quissange [-12.4333°, 14.0500°, 870 (Bocage, 1872, Ceríaco et al., 2016b). Undeter-
m] (Bocage, 1895); Benguela [-12.5833°, 13.4167°, mined locality: Between Benguela and Bihé
15 m] (Bocage, 1867a; Loveridge 1957); Hanha (Boulenger, 1905).
[-13.4941°, 14.5276°, 1669 m] (Bocage, 1896); Distribution in Angola: The species is
Sangevé [-13.8833°, 15.8333°, 1634 m] (Monard, widely distributed in the country, ranging from
1937); Ebanga [-12.7333°, 14.7333°, 1334 m] the southwestern Namibe Province to Lunda
(Monard, 1937); Lobito [-12.3709°, 13.5542°, 3 m Norte Province (fig. 8). The species appears to be
a.s.l] (Parker, 1936). Cunene Province: Kuvelai (= absent from the southeastern region of Cuando
Cuvelai) [-15.6500°, 15.8000°, 1217 m] (Monard, Cubango, where it is replaced by Trachylepis
1937); “Fleuve Mbalé” [-15.1667°, 16.7500°, 1245 damarana (see respective account).
m] (Monard, 1937); Chimporo [-16.0333°, Global Distribution: The species occurs
17.1500°, 1206 m] (Monard, 1937). Huíla Prov- from Angola to western Zambia (Weinell and
ince: Caconda [-13.7333°, 15.0667°, 1674 m] Bauer, 2018; Pietersen et al., 2021).
(Bocage, 1895); Kuvangu (= Kuvango) [-14.4667°, Habitat and Natural History Notes: The
16.3000°, 1453 m] (Monard, 1937); “Kapelongo” species is usually found in miombo habitats (fig.
[-14.8833°, 15.0833°, 1192 m] (Monard, 1937). 9), but can also be located in more disturbed
areas, like plantations and human settlements specimens from “Sangevé (Galangue),” Huíla
(Grandvaux-Barbosa, 1970). Butler et al. (2019) Province, ansorgii was closely related to T. sul-
noted that the species is a habitat generalist and cata (instead of T. bocagii as originally noted by
can be found on rock walls and manmade struc- Boulenger, 1907). Subsequent records of ansorgii
tures in addition to natural rocky outcrops and are to be treated with caution. According to the
downed logs. molecular results of Butler et al. (in review), T.
ansorgii occurs only in the highlands of Angola,
from northern Namibe and Huíla provinces
Trachylepis ansorgii (Boulenger, 1907)— northward to Malanje Province, with scattered
Ansorge’s Skink records following the escarpment and with an
Figures 10–12, plate 1 isolated record at Epupa Falls on the Namib-
ian side of the Cunene River, while T. sulcata is
Mabuia ansorgii Boulenger, 1907: 213. Holo- more widely distributed in the southern regions
type: BMNH 1946.8.3.33 (formerly BMNH of Namibe and Huíla provinces (and widely dis-
1906.8.24.62), collected by W.J. Ansorge. tributed in Namibia and western South Africa).
Type locality: “Caconda, Benguella” [= As there are no clearcut diagnostic features that
Caconda, Huíla Province], Angola. one could discern from most published accounts,
Euprepes olivaceus [part] [?]: Bocage (1870: 68). the allocation of historical specimens to either
Mabuia sulcata [part]: Bocage (1895: 41; 1896: taxon is based mostly on the distribution range
111). and should be considered provisory until molec-
Mabuia ansorgii: Monard (1937: 90). ular sampling from the same area offers insight
Mabuia bocagei ansorgei: (Laurent (1947: 8). into the given population’s taxonomic identity.
Mabuya sulcata ansorgii [part]: Hellmich (1957a: All Angolan T. ansorgii records subsequent to
64); Branch (1998: 157). those of Boulenger (1907) and Monard (1937)
Trachylepis sulcata [part]: Marques et al. (2018: most likely refer to nominotypical sulcata based
269). on distribution ranges. These include the records
Trachylepis sulcata ansorgii [part]: Portik et al. from Otschinjau, Cunene Province by Hellmich
(2010: 147, 2011: 1745); Masterson et al. (1957b), from “Munhino, 50 km à l’ouest de Sá
(2014b: 267); Baptista et al. (2019: 109); da Bandeira, district de Mocâmedes, 1000 m”
Branch et al. (2019a: 319); Butler et al. [= “Munhino,” 50 km west of Lubango, Namibe
(2019: 235); Conradie et al. (2022: 204). Provice; georeferenced to -14.96667, 12.96667
by Marques et al., 2018] by Laurent (1964), and
Boulenger (1907) described Mabuia Ansorgii from “11 km south of Chibemba,” Huíla Province
based on a “single, somewhat damaged specimen by Haacke in Baptista et al. (2019). Besides the
from Caconda, Benguella” [currently Caconda, most recently collected specimens (Butler et al.,
Huíla Province]. According to the author, the 2019, this paper, and another in review), the only
new taxon was “closely allied to M. Bocagii” historical specimens (excluding the type material
[= Trachyelpis bocagii Boulenger, 1887], but no and those recorded by Monard, 1937) that can be
comparisons were provided against other species safely allocated to T. ansorgii are those recorded
of the genus. The description noted a few diag- from Entre Rios, Benguela Province, and Nova
nostic characters of the newly described species Lisboa [currently Huambo], Huambo Province,
(Boulenger, 1907), but it was sufficiently vague to by Hellmich (1957a).
cause further confusions. Monard (1937) was the Diagnosis: A large-sized skink (max. SVL
first author to directly deal with the form after its 98.9 mm, MUHNAC/MB03-001340) with fully
original description, considering it as a valid full developed, pentadactyl limbs (figs. 10–12); dor-
species. According to Monard and based on three sal scales usually pentacarinate, but sometimes
FIGURE 10. Male specimen of Trachylepis ansorgii from Serra da Neve, Namibe Province (UF 187314). Photos
by L.M.P.C.
tricarinate, quadricarinate, or hexacarinate; ven- brown above, with six dark longitudinal stripes
tral scales smooth; 54–64 SAV; 50–55 SAD; (a pair of paravertebral stripes, a pair of dorso-
38–45 MSR; lamellae beneath fingers and toes lateral stripes, and another pair on the flanks)
spinose; plantar scales spinose; 18–24 LUFT; that are usually prolonged on the tail as irregular
13–18 LUFF; supranasals usually in contact; lines or series of spots; lower flanks whitish,
parietals usually separated or touching at a single sometimes with an irregular dark line or series of
point, rarely in broad contact; prefrontals usually spots. In adult females, the paravertebral and
separated, rarely touching at a single point; fron- dorsolateral stripes usually start at the frontopa-
toparietals always in contact; one pair of enlarged rietals and supraoculars, respectively, and there
nuchal scales present; ear opening vertically is often a dark spot on the frontal. Dorsum uni-
ovoid and smaller than the eye, lacking subtrian- form golden brown to dark brown in adult males;
gular auricular scales on the anterior margin of flanks and limbs light grayish; tail often with
the ear opening. Supralabials usually eight remnants of dark stripes. Ventral parts bluish
(sometimes seven), the sixth being subocular; white; labials, throat and chin sometimes yellow-
five supraciliaries, the second longest; nostril ori- ish to orange in live specimens, usually with
ented laterally. Color pattern exhibits pro- heavy black speckling, in some cases restricted to
nounced sexual and ontogenetic dimorphism. the labials and in others extending to the sides
Adult females and juveniles are pale to golden and top of the head.
FIGURE 11. Life photo of a gravid female Trachylepis ansorgii from Caconda, Huíla Province (CAS 263427).
Photo by L.M.P.C.
Material Examined: Benguela Province: 001472, 001475, 001477, 001478, 001482); Serra
Dembi, on the road from Cubal to Caimbambo da Neve, Catchi surroundings [-13.7620°,
[-13.0592°, 14.1279°, 906 m] (MUHNAC/MB03- 13.2569°, 1585 m] (MUNHAC/MB03-001473,
001391). Huambo Province: rocky outcrops 001474, 001479–001481, 001485); Serra da Neve
near Kanjonde village, base of Mt Moco base, 2km N of Maylowe [-13.8280°, 13.2625°,
[-12.4294°, 15.1545°, 1890 m] (MUHNAC/ 818 m] (MUHNAC/MB03-001476); Serra da
MB03-001392–001395). Huíla Province: Neve, Lutala crater surroundings [-13.7630°,
Caconda, rock outcrops [-13.7548°, 15.0422°, 13.2514°, 1598 m] (MUHNAC/MB03-1484).
1620 m] (CAS 263425, 263426, 263427, 263428, Additional Material (* denotes type material):
263429). Malanje Province: Laúca Dam, flooded Benguela Province: Entre Rios [-13.0167°, 14.6333°, 1267
area [-9.7627°, 15.1438°, 750 m] (MUHNAC/ m] (ZSM 103/1953). Cuando Cubango Province: en
route to Cuito, east of Huambo [-12.7362°, 15.9744°, 1777
MB03-001340, 001342–001345, 001363).
m] (PEM R23368). Huambo Province: Huambo
Namibe Province: base of Serra da Neve, May- [-12.7667°, 15.7333°, 1671 m] (ZSM 104/1953). Huíla
lowe village [-13.8357°, 13.2763°, 800 m] (MUH- Province: Sangevé [-13.8833°, 15.8333°, 1634 m] (MHNC
NAC/MB03-001396, 001397); Serra da Neve 91.0494, 91.0495); Caconda [-13.3333°, 15.0667°, 1644 m]
base camp [-13.7770°, 13.2591°, 1488 m] (CAS (BMNH 1946.8.3.33*); Caconda, rock outcrops [-13.7548°,
263567; UF 187313); Serra da Neve [-13.7865°, 15.0422°, 1620 m] (INBAC/AMB 10925, 10826).
13.2572°, 1594 m] (UF 187314); vic Dolondolo Historical Localities (no extant speci-
[-13.8104°, 13.1361°, 713 m] (CAS 263545); mens): Benguela Province: Hanha [-13.4941°,
Serra da Neve, rocky area near base camp 14.5276°, 1669 m] (Bocage, 1896). Huíla Prov-
[-13.7653, 13.2571, 1645 m] (MUNHAC/MB03- ince: Rio Cuce, près de Caconda (Bocage, 1895).
FIGURE 12. Life photo of a male Trachylepis ansorgii from Caconda, Huíla Province (CAS 263427). Photo by
L.M.P.C.
Distribution in Angola: Endemic to cen- Mabuya spilogaster [part]: Branch (1998: 156);
tral/western areas of the country, from northern Broadley (2000: 105).
Namibe and Huíla provinces to southern Malanje Trachylepis cf. spilogaster: Conradie et al. (2016:
Province (fig. 13). 25).
Global Distribution: Endemic to Angola. Trachylepis spilogaster: Marques et al. (2018:
Habitat and Natural History Notes: 268); Baptista et al. (2019: 116); Branch et
This is a rock-dwelling species, commonly found al. (2019a: 319); Butler et al. (2019: 315);
in rocky outcrops among miombo woodlands in Ceríaco et al. (2020a: 402); Conradie et al.
high elevation (above 700 m) areas (Grandvaux- (2022: 204).
Barbosa, 1970; fig. 14). It tends to be absent from
human-dominated landscapes and was never The first record attributable to this new species
found in human-made habitats such as houses or is from the Pulitzer Angola Expedition in Humbe,
on walls. Cunene Province, and reported by Schmidt (1933)
as “Mabuya chimbana” (accession number CM
Trachylepis attenboroughi, sp. nov.— 5655). A second specimen was reported by Lau-
Attenborough’s Skink rent (1964) from Mount Moco, in Huambo Prov-
Figures 15, 16, plate 2 ince (specimen still extant at the collections of the
Museu Regional do Dundo, Angola—accession
Mabuya chimbana: Schmidt (1933: 12). number MD 1832—see Ceríaco et al., 2020a). A
Mabuya striata spilogaster: Laurent (1964: 71). recent survey of the Mount Moco region has
FIGURE 13. Distribution of Trachylepis ansorgii in Angola. Black star denotes the type locality.
failed to collect more specimens in the region (2019) provided additional records for the same
(L.M.P.C., personal obs.). Conradie et al. (2016) province. According to our molecular data (fig. 1)
collected a considerable number of specimens in the central and southern Angolan populations
both Bié and Cuando Cubango provinces and constitute a distinct taxon, sister to T. spilogaster
attributed them to Trachylepis cf. spilogaster, not- sensu stricto, and morphological data also allow
ing however some ecological differences between us to separate these two taxa.
the Angolan populations and those from Namibia Holotype: An adult male (CAS 263386, field
and the rest of the species distribution. Marques number AMB 10669; fig. 15) collected at Bicuar
et al. (2018) also noted that it was “highly unlikely National Park, camp headquarters, Huíla Prov-
that these [Angolan] records refer to the same lin- ince [-15.1016°, 14.8399°, 1224 m], collected by
eage as true T. spilogaster.” Butler et al. (2019) pro- Mariana P. Marques, Luis M.P. Ceríaco, Suzana
vided the first records of the species for Huíla A. Bandeira, Brett O. Butler, Matthew P. Heinicke
Province, and some months later Baptista et al. and Timóteo Júlio, on 28 July 2017.
FIGURE 14. Typical habitat of Trachylepis ansorgii in Caconda, Huíla Province. Photo by L.M.P.C.
Paratypes: All 15 specimens from Angola: an and Timóteo Júlio, on 4 August 2017; an adult
adult male (CM 5655), collected at Humbe, female (CAS 263424, field number 10791), col-
Cunene Province [-16.6872°, 14.9065°, 1106 m] lected at Instituto Superior de Ciências da Edu-
by Rudyerd W. Boulton and Laura T. Boulton on cação (ISCED), Lubango, Huíla Province
13 November 1930; four adult females (CAS [-14.9182°, 13.4849°, 1867 m] by Mariana P.
263384, 263385, field numbers AMB 10168, Marques, Luis M.P. Ceríaco, Suzana A. Bandeira,
10666; UF 187320, 187321, field numbers AMB Brett O. Butler, and Matthew P. Heinicke, on 6
10153, 10166), same data as holotype with excep- August 2017; an unsexed adult (CAS 263397,
tion of the collecting dates, 27 and 28 July 2017 field number AMB 10811), collected at Chibia,
(respectively); two adult females (CAS 263415, near children’s park, Huíla Province [-15.1894°,
field number AMB 10130; UF 187319, field num- 13.6905°, 1486 m] by Mariana P. Marques, Luis
ber 10139) and one adult male (CAS 263416, M.P. Ceríaco, Suzana A. Bandeira, Brett O. But-
field number AMB 10138), collected at Kimbo ler, and Matthew P. Heinicke, on 7 August 2017;
do Soba, Lubango, Huíla Province [-14.9342°, three unsexed adults (MUNHAC/MB03-
13.4693°, 1866 m] by Mariana P. Marques, Luis 001398–001400, field numbers AMB 10991,
M.P. Ceríaco, Suzana A. Bandeira, Brett O. But- 10992, 11021), collected at Bicuar National Park,
ler, Matthew P. Heinicke and Timóteo Júlio, on Matunto Ranger Station, Huíla Province
21 and 22 July 2017; an adult female (CAS [-15.3694°, 15.2751°, 1159 m] by Mariana P.
263403, field number AMB 10737) and an Marques, Luis M.P. Ceríaco, and Hilária Valério,
unsexed adult (UF 187322, field number AMB on 10 March 2018.
10739) collected at Serra da Leba overlook, Huíla Material Examined: Huambo Prov-
Province [-15.0770°, 13.2329°, 1680 m] by Mari- ince: Serra do Moco, Luimbale [-12.5333°,
ana P. Marques, Luis M.P. Ceríaco, Suzana A. 15.1833°, 1748 m] (MD 1832). Huíla Prov-
Bandeira, Brett O. Butler, Matthew P. Heinicke ince: Bicuar National Park, Matunto Ranger
FIGURE 15. Holotype of Trachylepis attenboroughi, sp. nov., from Bicuar National Park, Huíla Province (CAS
Station [-15.3694°, 15.2751°, 1159 m] (MUN- National Park (6) [-15.1305°, 14.6837°, 1257 m]
HAC/MB03-001377–001379, 001423–001427, (NB532 fide Baptista et al., 2019); Bicuar National
001456–001462). Park (9) [-15.1267°, 14.6377°, 1281 m] (NB528 fide
Baptista et al., 2019); Bicuar National Park (10)
Additional Material: Bié Province: Cubango
[-15.1260°, 14.6012°, 1298 m] (NB527 fide Baptista et
basin (12b) [-13.5964°, 16.8772°, 1517 m] (PEM
al., 2019); Bicuar National Park (13) [-15.1005°,
R20019). Cuando Cubango Province: Cubango basin
14.8451°, 1250 m] (NB529 fide Baptista et al., 2019);
(6a) [-14.6716°, 17.7353°, 1391 m] (PEM R21842);
Bicuar National Park, Main Camp (31) [-15.1007°,
Cubango basin (20) [-14.6718°, 17.1533°, 1355 m]
14.8396°, 1244 m] (NB519 fide Baptista et al., 2019).
(PEM R20007); Cuito basin (30a) [-16.9098°, 19.3077°,
Moxico Province: Cuito River source lake [-12.6894°,
1106] (PEM R20517); Cuito basin (35) [-16.6232°,
18.3601°, 1435 m] (PEM R23334–5); Quembo River
19.0535°], 1128 m (PEM R20518); Cuito basin (32)
source [-13.1069° 19.0179°, 1545 m] (PEM R23528);
[-17.0488°, 19.5333°, 1105 m] (PEM R20519);
DOR en route to village [-13.0597°, 18.8324°, 1567 m]
Cubango basin (47) [-14.7021°, 17.3781°, 1392 m]
(PEM R23358–60); DOR en route between Cuanavale
(PEM R21828‒21840); Cubango basin (49) [-14.6630°,
River source and Tempué [-13.3395°, 18.8512°, 1386
17.6655°, 1384 m] (PEM R21483); Cuito basin (57)
m] (PEM R27441); Quembo River, walk back from
[-15.4597°, 18.7633°, 1242 m] (PEM R21486‒7); Cuito
small waterfall [-13.5298°, 19.2834°, 1242 m] (INBAC/
basin (62) [-17.5088°, 20.0661°, 1079 m] (PEM
WC-6813).
R21498‒9); Hotel Mulombe, Menongue [-14.6641°,
17.6926°, 1363 m] (CM 172593-4); Cuatir main camp Diagnosis: A medium-sized skink (max. SVL
[-16.4852°, 18.2030°, 1148 m] (CM 172595-8). 79.4 mm, CAS 263424), with fully developed,
Cunene Province: Otchinjau [-16.5025°, 13.9240°, pentadactyl limbs (figs. 15, 16); dorsal scales tri-
363 m] (ZSM 68-1954). Huíla Province: Bicuar carinate or quadricarinate; ventral scales smooth;
FIGURE 16. Life photo of the paratype of Trachylepis attenboroughi from Bicuar National Park, Huíla Province
(UF 187321). Photo by L.M.P.C.
53–61 SAV; 52–57 SAD; 34–40 MSR; lamellae brown above, speckled with white or light gray.
beneath fingers and toes keeled and spinose; There may be scattered black and/or white speck-
plantar scales spinose; 17–22 LUFT; 13–18 LUFF; les on the flanks and dorsal surfaces. Labials
supranasals always in contact; parietals usually in cream white, often darker at the sutures and
contact, but sometimes separated or touching at sometimes with irregular dark speckling. Lower
a single point; prefrontals usually separated, flanks grayish; venter surfaces white with grayish
sometimes touching at a single point; frontopa- speckling, especially laterally and on the chin
rietals always in contact, rarely touching at a and throat area.
single point; one pair of enlarged nuchal scales Description of the Holotype: A well-pre-
present; ear opening vertically ovoid and smaller served adult male. Body cylindrical and robust
than the eye, lacking subtriangular auricular with a poorly defined neck and well-developed
scales on the anterior margin of the ear opening. pentadactyl limbs; tail long and robust, its
Supralabials eight (rarely seven), the sixth suboc- length greater than the SVL, smoothly tapering.
ular; five supraciliaries, the second longest; nos- Fore- and hind limbs overlap when adpressed
tril oriented laterally. Dorsum grayish to reddish against the body. SVL 71.4 mm, TL 98.3 mm.
brown, with a pair of pale dorsolateral stripes HL 17.8 mm, with relatively long and promi-
with tendency to become broader and fainter nent snout. Additional measurements are pre-
posteriorly; upper flanks with a black band that sented in table 4. Ear opening medium, lacking
starts at the snout and extends to the tail base, anterior subtriangular auricular scales. Rostral
becoming fainter posteriorly; limbs grayish visible from above. Nostrils oriented laterally
TABLE 4
Mensural and Meristic Data for the Type Series of Trachylepis attenboroughi, sp. nov. 2024
Abbreviations are listed in the Materials and Methods. Measurements are presented in millimeters and ratios as percentages
CAS CM CAS CAS CAS CAS CAS CAS CAS UF UF UF UF MUHNAC/ MUHNAC/ MUHNAC/
263386 5655 263384 263385 263415 263416 263403 263424 263397 187320 187321 187319 187322 MB03-1398 MB03-1399 MB03-1400
Holo- Para- Para- Para- Para- Para- Para- Para- Paratype Para- Para- Para- Paratype Paratype Paratype Paratype
type type type type type type type type type type type
Sex ♂ ♂ ♀ ♀ ♀ ♂ ♀ ♀ unsexed ♀ ♀ ♀ unsexed unsexed unsexed unsexed
SVL 71.4 62 77 66 69 69.7 77.5 79.4 51.4 56.3 75 67.7 59.7 68.4 69.8 62.4
TL 98.3 – – 74 – 90 – 74.3 77.7 87.1 101.6 – 74.9 110.6 111.9 –
TL/ 280 – – 110 – 130 – 90 150 150 140 – 130 160 160 –
SVL
HL 17.8 14.4 16.8 15.5 17 16.9 18.2 19.4 13.7 13.1 17.6 16.2 14.9 13.3 15.7 12.4
HL/ 20 20 20 20 20 20 20 20 30 20 20 20 20 20 20 20
SVL
SVL/ 400 430 460 420 410 410 430 410 370 430 430 420 400 510 440 500
HL
HW 11 9.4 11.2 9.4 10.1 10.4 11.9 12.1 8 8.4 10.7 9.6 8.8 10.8 10.8 8
HW/ 60 70 60 60 60 60 70 60 60 60 60 60 60 80 70 60
HL
HH 2.5 6.6 7.6 6.9 7.5 7.6 4.8 8.6 6.1 5.7 7.7 7.2 6.6 6.9 7.3 6.6
IN 2.4 2.5 2.1 2.1 2.3 2.4 2.2 2 2 2 2 2.1 1.8 2.4 2.5 2.3
EN 5.3 3.6 4.5 4.4 4.6 4.6 5.8 4.8 3.7 3.9 4.6 4.8 4.2 3.8 3.7 3.7
ES 7 6.3 6.5 6.4 6.9 7 7.3 7.1 5.5 5.6 6.5 6.8 6 5.2 5.8 5
MSR 35 35 36 36 35 37 34 35 38 38 37 37 37 39 37 39
SAD 57 54 56 54 57 53 54 56 54 54 54 57 54 54 54 53
SAV 61 60 59 58 59 59 58 59 61 58 58 61 60 55 58 56
LUFF 19 17 16 16 17 18 16 17 16 16 16 17 17 15 15 14
LUFT 16 22 21 20 22 22 20 20 22 20 19 22 20 19 20 20
SC 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5
SL (SO) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6)
CP C C C SPC C SPC C C C S SPC C S C C C
CFP C C C C C C C C C C C C C SPC C C
CSN C C C C C C C C C C C C C C C SPC
CPF SPC S S SPC SPC S C S SPC C S S S S S S
KDS 3 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4
Plantar
spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose
scales
35
and set posteriorly, so that postnasal effectively attenboroughi differs from all the other spe-
borders nostril. Supranasals in contact. Fronto- cies of Trachylepis known to occur in Angola,
nasal broader than long, in contact with loreal with the exception of T. wahlbergii, by having
scale. Prefrontals irregularly pentagonal, sepa- a combination of keeled plantar scales and
rated from one another, each in contact with the absence of auricular scales on the anterior
following head shields: frontonasal, loreal, first margin of the ear opening. It differs from T.
and second supraocular and frontal. Two lore- wahlbergii (and from T. striata) in having a
als. Frontal longer than the distance between dorsum brownish, with conspicuous black or
anterior tip of frontal and tip of snout. Frontal white speckles (vs. dorsum homogeneous
in contact with three supraoculars on each side. brownish, without black or white speckles in
Two frontoparietals, in contact with each other T. wahlbergii and T. striata). Regarding the
and the frontal, third and fourth supraoculars, other southwest African congeners of the stri-
parietal, and interparietal. Frontoparietal plus ata subgroup sensu Weinell et al. (2019), T.
interparietal length equal to frontal length. attenboroughi differs from T. punctatissima by
Interparietal twice as long as broad, with a vis- having three to four rows of scales above win-
ible parietal foramen. Parietals of greater length dow in eyelid (vs. two rows in T. punctatis-
than frontoparietals. Parietals in contact with sima), from T. sparsa by having a clear pattern
each other. Five supraciliaries, second largest. of lateral stripes and a nonmelanistic dorsum
Eight supralabials, sixth being subocular. Eight (vs. a black dorsum with small scattered light
infralabials. Postmental bordering seven scales speckling in T. sparsa) and before the entire
(mental, two infralabials on each side, and two ventrum heavily speckled with black (vs. ven-
primary chin shields). Transparent scale pres- trum uniform in T. sparsa), and finally from
ent in lower eyelid, as is usual for Trachylepis. its sister taxon, T. spilogaster, by having 52–57
Tympanum visible, at same level as mouth. SAD (vs. 48–52 in T. spilogaster).
Dorsal scales each with three smooth keels. Distribution in Angola: The species occurs
Ventral scales smooth. MSR 35, SAD 57, SAV in the southern Angolan Plateau (Huíla, Cunene,
61. Limbs with five digits; scales on palms and Bié, and Cuando Cubango provinces), being
soles keeled and spinose. Relative length of fin- absent from the coastal and lower elevation areas
gers IV > III > II > V > I, relative length of toes of Namibe Province (fig. 17).
IV > III > V > II > I. Finger-IV lamellae 16, Global Distribution: The bulk of the dis-
Toe-IV lamellae 19. tribution of this species is in Angola, but it
Coloration in Ethanol: Dorsum reddish extends into neighboring northern Namibia both
brown with a pair of pale dorsolateral stripes that in the Kunene Region and the Kavango Region
progressively start fading around midbody; top of and probably in intervening areas (Bauer et al.,
head, limbs, and tail grayish brown; upper flanks unpubl. data).
with a black stripe that starts at the loreals and Habitat and Natural History Notes: The
extends to the tail, gradually becoming fainter species is mostly associated with sparse miombo
posteriorly from midbody on; labials cream white woodlands at high elevation areas (Grandvaux-
with dark speckling, especially near the posterior Barbosa, 1970; fig. 18). Most of the specimens
edge. Venter white, with extensive gray speckling collected were found in either sparsely vegetated
near the flanks and on the chin and throat area. areas, or sometimes near rocky outcrops. Con-
Variation: Variation in scalation and mea- radie et al. (2016) note that the species is more
surements among the type series is reported commonly found on the ground, in rock piles,
in table 4. and on village houses. Baptista et al. (2019) note
Comparison with Other Angolan and that it “was the most frequently observed rep-
Southwestern African Trachylepis: T. tile species” in the Bicuar National Park, and
FIGURE 17. Distribution of Trachylepis attenboroughi in Angola. Black star denotes the type locality.
“together with Pachydactylus punctatus, was one Trachylepis bayonii (Bocage, 1872)—
of only two species to be found in the park’s Bayão’s Skink
woodlands after recent fires.”
Etymology: The specific epithet “attenbor- Figures 19–22, plate 2
oughi” is formed in the genitive singular and is
masculine. It is given in honor of David F. Atten- Euprepes Bayonii var. A Bocage, 1872: 75. Syn-
borough (1926–), British naturalist and science types: MBL (specimen numbers unknown,
communicator in recognition of his inspiring destroyed by fire on March 18, 1978),
works promoting the study of natural history BMNH 1946.8.19.13 (formerly MBL,
and biodiversity conservation. We suggest BMNH 66.6.11.8), ZMB 6477, collected by
“Attenborough’s Skink” and “Lagartixa do Atten- F.A.P. Bayão. Type locality: “Duque de
borough” as the English and Portuguese com- Bragança, dans l’intérieur d’Angola” [=
mon names, respectively, for this species. Calandula, Malanje Province], Angola.
FIGURE 18. Typical habitat of Trachylepis attenboroughi in Bicuar National Park, Huíla Province. Photo by
L.M.P.C.
Euprepes Gravenhorstii: Bocage (1866a: 44). (1872) based on specimens from Duque de Bran-
Euprepes Bayonii: Bocage (1870: 68; 1872: 75; gança (currently Calandula), Malanje Province,
1879b: 95; 1887: 179); Bauer et al. (2003: and Huíla, Huíla Province. According to Bocage
270). (1872), the species contained two varieties: “var.
Mabuia bayonii: Boulenger (1887: 201; 1905: A,” originating from Duque de Bragança, and
111); Bocage (1895: 38; 1897a: 195); Monard “var. B,” originating from Huíla (subsequently
(1937: 87). recognized as T. huilensis, see account below).
Mabuya bayonii: Schmidt (1933: 11); Hellmich The main differences between these two varieties
(1957b: 54); Manaças (1963: 234); Bauer et were in their coloration patterns. Bocage’s (1870)
al. (2003: 270). reference to “Eupr. Bayonii. nov. sp.” represents a
Mabuya bayoni bayoni: Laurent (1964: 67). nomen nudum, as no description accompanied
Trachylepis bayonii [part]: Ceríaco et al. (2016b: the new nomen. Bocage (1895) emphasized the
57; 2018: 423; 2020a: 402); Marques et al. difference between the two varieties and pro-
(2018: 256); Ceríaco et al. (2021: 115); San-
vided an illustration of the species (fig. 19).
tos et al. (2021: 24). Bauer et al. (2003) noted that two of the syn-
Trachylepis bayonii bayonii: Branch et al. (2019: types, both from Duque de Bragança, were still
extant in the collections of the ZMB and BMNH
318); Conradie et al. (2022: 202).
(accession numbers ZMB 6477 and BMNH
Trachylepis bayonii is an Angolan endemic, 1946.8.19.13, respectively), while the remaining
easily distinguished from all of its congeners by specimens that were housed in the Lisbon
fused frontoparietal scales, a diagnostic charac- Museum were lost during the fire that destroyed
ter that is absent in all other Angolan species the museum in 1978. The species has been
and particularly rare amongst other Trachylepis. noted in the country by several authors since
The species was originally described by Bocage the original description (Boulenger, 1887, 1905;
FIGURE 19. Fragment of plate III of Bocage’s (1895) “Herpétologie d’Angola et du Congo” (color edition),
depicting Trachylepis bayonii.
Schmidt, 1933; Monard, 1937; Hellmich, 1957b; seven keels; ventral scales smooth; 58–61 SAV;
Manaças, 1963; Laurent, 1964; Ceríaco et al., 50–56 SAD; 28–35 MSR; lamellae beneath fin-
2018a). Marques et al. (2018) noted that the gers and toes smooth; plantar scales smooth;
record from “S. Salvador do Congo” (currently 15–16 LUFT; 12–13 LUFF; supranasals always in
M’Banza Kongo), Zaire Province, from Bocage contact; parietals usually separated; prefrontals
(1895), was out of the expected species range usually separated; frontoparietals fused; enlarged
and probably represents a misidentification, but nuchal scales absent; ear opening vertically ovoid
a recent sighting of a road-kill specimen (not and smaller than the eye; 3–4 subtriangular
collected; fig. 20) near Zulumongo [-7.207669°, auricular scales (shorter than the diameter of
15.094092°, 1184 m], Uíge Province, proves that ear) extend posteriorly and usually slightly
the species distribution reaches these northern upward from the anterior margin of ear opening.
regions of the country. However, as we have no Supralabials usually eight (rarely nine), the sixth
molecular data from these northern specimens, subocular; four supraciliaries, the first notably
we cannot reject the hypothesis that they repre- longer; nostrils oriented dorsally. Dorsum olive
sent a different taxon. to reddish brown, with a pair of faint pale dorso-
Loveridge (1956) described a subspecies of T. lateral stripes; between these there are usually
bayonii, T. b. keniensis based on specimens from two series of longitudinally aligned black spots.
central Kenya and southern Tanzania. Recent Top of head uniform brownish; labials cream
authors (e.g., Spawls et al., 2018) still considered white; a narrow white stripe extends from the
it as a subspecies of bayonii, but, based on mor- labials to the hind limb insertion, sometimes
phological differentiation between nominotypic bordered with black; flanks and lateral side of
bayonii and keniensis (pentacarinate dorsal scales head and neck usually with extensive black
in bayonii versus tricarinate in keniensis) and speckling. Venter bluish white, with some gray-
their geographic separation, we consider kenien- ish speckling near the flanks; subdigital lamellae
sis as a full species. and plantar scales brown.
Diagnosis: A medium-sized skink (max. SVL Material Examined (* denotes type mate-
83.0 mm, CAS 258387), with fully developed, rial): Bié Province: Chitau [-11.4333°, 17.1500°,
pentadactyl limbs (figs. 19–22); dorsal scales 1510 m] (CM 5866, 5881). Huambo Province:
usually pentacarinate, but may have three to Kajonde village, Mt. Moco [-12.4294°, 15.1545°,
FIGURE 20. Specimen of Trachylepis cf. bayonii found dead on the road near Zulumongo, Uíge Province. Note
the fused fronto-parietal scales. Photo by L.M.P.C.
1890 m] (MUNHAC/MB03-001385). Malanje (=N’dalatando) [-9.3000°, 14.9167°, 782 m] (BMHN

Province: Duque de Bragança (currently Kalan- 1909.10.29.100); Duque de Bragança (currently Kalan-
dula) [-9.0740°, 15.9999°, 1065 m] (ZMB 6477*); dula) [-9.0740°, 15.9999°, 1065 m] (BMHN 1904.5.2.41,
1946.8.19.13*); Pavalange [-11.3667°, 17.6167°, 1176 m]
Cangandala National Park [-9.8269°, 16.6500°,
(ANSP 32199); Cangandala National Park [-9.82689°,
1189 m] (CAS 258387); right bank of the
16.65003°, 1086 m] (CAS 258375); Gauca, 20 mi E of
Kwanza River [-9.8853°, 16.2872°, 1005 m] (CAS Dando (on Quanza river) [-11.1833°, 14.4500°, 1163 m]
258388); Gauca, 20 mi E of Dando (on Quanza (CM 5738, 5935, 5937, 5940, 5943). Moxico Province:
river) [-11.1833°, 14.4500°, 1163 m] (CM 5736, environs du lac Calundo [-11.8000°, 20.8667°, 1119 m]
5739). Undetermined locality: Angola (MHNC- (MD 5602); Calombe [-11.8333°, 19.9333°, 1355 m]
UP/REP 276, 277). (IICT 243/1959; 246/1959; 254/1959; 255/1959;
Additional Material: Bié Province: Bihe (= Bié) 258/1959; 268–278/1959; 288/1959; 324/1959,
[-12.3833°, 16.9500°, 1660 m] (FMNH 18508, 18509, 325/1959; 327–338/1959; 364–373/1959; 376–
18510); Chitau [-11.4333°, 17.1500°, 1510 m] (AMNH 385A/1959; 421–427/1959); Cuito River source lake
48649; CM 5869, 5890, 5891, 5893, 5902; NMB 13227); [-12.6894°, 18.3601°, 1435 m] (PEM R23336–8); Kulua
Kwanza River Bridge [-11.9935°, 17.6696°, 1273 m] River source lake, 6 km SE of Cuito River source
(PEM R23378). Huambo Province: Huambo [-12.7368°, 18.3931°, 1446 m] (PEM R23354); Quembo
[-12.7667°, 15.7333°, 1671 m] (AMNH 48650, 48651). River trap 2 [-13.1354°, 19.0440°, 1369 m] (PEM
Kwanza Norte Province: Ambaca [-9.2667°, 15.1833°, R23477); Quembo River source lake [-13.1410°,
738 m] (BMNH 1904.5.2.42); Piri-Dembos [-8.5299°, 19.0543°, 1399 m] (PEM R23501); Cuito River source
14.4377°, 712 m] (ZMH 1086). Lunda Norte Province: lake [-12.6886°, 18.3603°, 1426 m] (PEM R23514);
Luzamba compound [-9.1167°, 18.0500°, 861 m] (PEM Kulua River source [-12.7372°, 18.3934°, 1444 m] (PEM
R13474, 13488). Lunda Sul Province: Alto Cuílo, R23516); Quembo River source camp [-13.1410°,
Lunda [-10.0500°, 19.5170°, 1260 m] (MCZ R-74109, 19.0543°, 1371 m] (PEM R23553–5; INBAC/WC-4674);
74110; MD 5306). Malanje Province: Ndalla Tando Rio Comba [-12.6244°, 18.6516°, 1299 m] (PEM
FIGURE 21. Specimen of Trachylepis bayonii from Cangandala National Park, Malanje Province (CAS 258388).
Photos by L.M.P.C.
R23971); Lungwebungu River trap 1 [-12.5801°, (Bocage, 1872, 1895); Kalukembé [-13.7833°,
18.66740°, 1298 m] (PEM R23987); Lungwebungu 14.683°, 1699 m] (Monard, 1937); Kuvangu (=
River camp, at bridge [-12.5839°, 18.6655°, 1295 m] Kuvango) [-14.4667°, 16.3000°, 1453 m]
(PEM R27420); Lungwebungu River camp [-12.5844°, (Monard, 1937). Lunda Norte Province: Cas-
18.6675°, 1297 m] (PEM R27421); Quembo River
sange [-9.5833°, 17.8667°, 955 m] (Bocage,
bridge camp, trap 1 [-13.5280°, 19.2815°, 1236 m]
(PEM R27422); Quembo River right side tributary
1879b, 1895, 1897a). Malanje Province: Duque
(Micongo River) past village [-13.5187°, 19.2849°, 1248 de Bragança [-9.1000°, 15.9500°, 1010 m]
m] (PEM R27423–4). Undetermined locality: (Bocage, 1866a, 1872, 1895, 1897a). Zaire Prov-
Unknown location (AMNH 48652–48661; FMNH ince: S. Salvador do Congo [-6.2667°, 14.2333°,
74298, 74299; MNHN 1866.72); Angola (MNCN 5851, 481 m] (Bocage, 1887).
5880; MHNC 91.0538–91.541; USNM 26389); between Other Localities (specimen not vouch-
Benguela and Bihe (BMHN 1905.5.29.19). ered): Uíge Province: near Zulumongo [-7.2077°,
Historical Localities (no extant speci- 15.0941°, 1184 m] (LMPC, personal obs.).
mens): Benguela Province: Cahata [-12.3500°, Distribution in Angola: The species occurs
14.8167°, 1202 m] (Bocage, 1895). Huambo on the Angolan Plateau, especially in the central
Province: Santo Amaro [-12.7000°, 15.8500°, and northern provinces of Huíla, Huambo, Bié,
1707 m] (Monard, 1937). Huíla Province: Benguela, Moxico, Kwanza Norte, Kwanza Sul,
Caconda [-13.7333°, 15.0667°, 1674 m] (Bocage, Bengo, Malanje, Lunda Norte, Lunda Sul, Uíge,
1895); Huilla [-15.0500°, 13.5500°, 1733 m] and Zaire, and absent from the coastal and lower-
FIGURE 22. Life photo of Trachylepis bayonii from Mount Moco, Huambo Province (MUNHAC/MB03-
001385). Photo by L.M.P.C.
elevation areas, as well as the southern provinces Orthoptera, arachnids and larvae of other small
of Cuando Cubango and Cunene (fig. 23). invertebrates in their stomach contents.
Global Distribution: Confirmed records of
the species are restricted to Angola, although it
Trachylepis binotata (Bocage, 1867)—
possibly extends to adjoining areas of the Demo-
Ovambo Tree Skink
cratic Republic of Congo, as some northern
records occur in habitats that are continuous Figures 25–27, plate 2
between the two countries. In Huila province,
where the species is sympatric with T. huilensis Euprepes binotatus Bocage, 1867b: 230. Syn-
(see account below), T. bayonii is limited to the types: MBL (specimen numbers unknown,
northern parts of the province. destroyed by fire on March 18, 1978),
Habitat and Natural History Notes: The MNHN 1462 fide Brygoo (1985), BMNH
species occurs in a wide range of habitats, from 1946.8.15.32 (formerly BMNH 67.7.23.26)
miombo dominated forests to grasslands (Grand- fide Boulenger (1887), ZMB 5830 fide Bauer
vaux-Barbosa, 1970; fig. 24). Manaças (1963) et al. (2003), collected by J.A. Anchieta.
notes that some females collected between Sep- Type locality: “Benguella,” Benguela
tember and November had well developed Province, Angola.
embryos. The same author noted that the Cal- Euprepes binotatus: Bocage (1867a: 223; 1879a:
ombe, Luso (Moxico Province) specimens con- 88); Brygoo (1985: 13).
tained a large number of termites, Coleoptera, Mabuya quinquetaeniata: Boulenger (1887: 198).
FIGURE 23. Distribution of Trachylepis bayonii in Angola. Black star denotes the type locality.
Mabuia binotata: Bocage (1895: 46, 1897a: 196); tos et al. (2021: 24); Lobón-Rovira et al.
Monard (1937: 91). (2022: 309).
Mabuya binotata: Hellmich (1957a: 59, 1957b: Trachylepis binotata is one of the most conspicu-
54); Laurent (1964: 68); Branch (1998: 151); ous species in the arid regions of southern Angola
Bauer et al. (2003: 270). and Namibia given its considerable size and arbo-
Euprepes binotata: Bauer et al. (2003: 270). real habits. The species was described by Bocage
Mabuya quinquetaeniata binotata: Mertens based on three specimens from Benguela Province
(1926: 152, 1937: 10, 1938: 437). (Bocage, 1867b), and the description was accompa-
Trachylepis binotata: Ceríaco et al. (2016b: 57, nied by an illustration of the species (fig. 25). How-
2021: 402); Marques et al. (2018: 257); ever, in his previous paper (Bocage, 1867a)
Branch et al. (2019a: 318); Butler et al. published in the same issue as the description
(2019: 234); Baptista et al. (2019: 109); San- (Bocage, 1867b), the author presented “Euprepes
FIGURE 24. Typical habitat of Trachylepis bayonii in Kajonde, Huambo Province. Photo by L.M.P.C.
binotatus. Nov. sp.,” based on several specimens smooth; 59–72 SAV; 51–60 SAD; 34–42 MSR;
from Benguella, Dombe, and Catumbella collected lamellae beneath fingers and toes smooth; plantar
by José d’Anchieta. This first mention, however, scales smooth; 17–22 LUFT, 14–17 LUFF; suprana-
lacked any formal description and diagnosis, and sals always in contact; parietals usually separated;
thus is considered here a nomen nudum. It is likely prefrontals usually in contact; frontoparietals
that these specimens are the same as those used in always in contact; one pair of enlarged nuchal
the description (Bocage, 1867b), of which one syn- scales present; ear opening vertically ovoid and
type is still extant in the collections of ZMB (ZMB smaller than the eye; lacking subtriangular auricu-
5830 fide Bauer et al., 2003), while another is in the lar scales on the anterior margin of the ear opening.
collection of the BMNH (BMNH 1946.8.15.32), Supralabials usually eight, the sixth subocular;
whence they were sent in the 19th century as dupli- supraciliaries usually five, the second the longest;
cates. The latter syntype was erroneously listed by nostrils oriented laterally. Dorsal surfaces grayish
Boulenger (1887) as Mabuya quinquetaeniata. Due brown, uniform or with scattered black spots that
to its large size and conspicuous eye mask, the spe- tend to form transverse bars on the back; there may
cies has never posed any taxonomic problems to also be scattered pale spots, especially on the flanks
authors who have worked with it, and its identity and limbs. Labials cream white, sometimes with
has not been disputed. light grayish stippling; there is a characteristic
Diagnosis: A large and robust skink (max. SVL broad, black band that starts at the eye and ends
134.3 mm, MUNHAC/MB03-1386), with fully above the forelimb insertion, often bordered above
developed, pentadactyl limbs (figs. 25–27); dorsal by a faint pale stripe. Lower flanks and venter
scales tricarinate or pentacarinate; ventral scales white, often with light grayish speckling on the chin
FIGURE 25. Section of the plate of Bocage (1867b) depicting Trachylepis binotata.
and throat; subdigital lamellae and plantar scales Additional Material: Benguela Province: Ben-
usually brownish. guela [-12.58333°, 13.41667°, 15 m] (MNHN 0.1462;
Material Examined (* denotes type mate- BMNH 1946.8.15.32*, 1906.8.24.69, 1906.8.24.70;
ZSM 85/1953); Catengue [-13.0333°, 13.7333°, 553 m]
rial): Benguela Province: Catumbela [-12.4333°,
(MCZ R-22420, 22421, 28636; NHMW 9677; SMF
13.5500°, 15 m] (ZMB 5830*). Huíla Province: 21579–21588); Cubal [-13.0333°, 14.2500°, 921 m]
Bicuar National Park, Matunto Ranger Station (SMF 25393–25403). Cunene Province: Mupa
[-15.3694°, 15.2751°, 1159 m] (MHNC-UP/REP [-16.1833°, 15.7500°, 1166 m] (MHNC 91.0497,
441). Namibe Province: Capangombe 91.0498, 91.0499); Kuvelai (= Cuvelai) [-15.6500°,
[-15.0974°, 13.1391°, 542 m] (MUNHAC/ 15.8000°, 1217 m] (NMB 12993); Ruacana Falls
MB03-001386, 001452); Maconjo [-15.0167°, [-17.3876°, 14.2118°, 897 m] (TM 38642–38645).
13.2000°, 856 m] (ZMB 7794); Virei-Chipumpo Namibe Province: Maylowe [-13.8357°, 13.2763°, 800
m] (INBAC/LMPC 1150); Capangombe [-15.0974°,
[-16.2442°, 12.9179°, 617 m] (CAS 263496);
13.1391°, 542 m] (CAS 266055, MB03-001452,
Virei camp [-16.1196°, 12.8346°, 522 m] (UF INBAC/AMB 9835); Maungo farm [-14.3934°,
187303); Camucuio [-14.1137°, 13.2432°, 670 12.8289°, 367 m] (INBAC/AMB 11461); Camacuio
m] (CAS 263535); Maungo houses [-14.5380, [-14.1137°, 13.2432°, 670 m] (AMB 10351); Virei
12.7448, 365 m] (CAS 264721); base camp, camp [-16.1196°, 12.8346°, 522 m] (UF 187303); 50
located in a dry river line at Maungo farm km Moçâmedes road to Sá da Bandeira [-15.0136°,
[-14.3934°, 12.8289°, 367 m] (CAS 264733); 12.5186°, 531 m] (MD 1967); Vila Arriaga [-14.7667°,
base of Serra da Neve, Maylowe village 13.3667°, 955 m] (TM 22605); 11 km south of
Chibemba [-15.8400°, 14.1134°, 1280 m] (TM 40103);
[-13.8357°, 13.2763°, 800 m] (CAS 266149);
Iona National Park [-16.9000°, 12.5833°, 1000 m] (TM
Serra da Neve base, 2km N of Maylowe 40738); Humbe [-16.6833°, 14.9000°, 1105 m] (TM
[-13.8280°, 13.2625°, 818 m] (MUHNAC/ 45125); Oncocua-Otchinjau 37 km NE of [-16.7213°,
MB03-001453, 001454). Undetermined local- 13.1569°, 882 m] (TM 40795); 12 km from Oncocua
ity: Unknown locality (MHNC-UP/REP 268). [no exact location] (TM 40765); 38 km from Oncocua
FIGURE 26. Specimen of Trachylepis binotata from Maungo Farm, Namibe Province (CAS 264721). Photos
by L.M.P.C.
[no exact location] (TM 40770); Otchifengo Global Distribution: Endemic to south-
[-16.6849°, 12.8413°, 584 m] (AMB 13076); Otchi- western Angola and northwestern Namibia
fengo to Montenegro road [-16.8712°, 13.1356°, 764 (Bauer et al., 1993).
m] (AMB 13174). Undetermined locality: Bombome,
Habitat and Natural History Notes: The
Moçamedes (BMNH 1907.6.29.33–34); Unknown
locality (AMNH 48763–48769).
species is associated with dry habitats, like
Historical Localities (no extant speci- mopane and miombo forests and savannahs
mens): Benguela Province: Catumbella (Grandvaux-Barbosa, 1970; fig. 29). It is usually
[-12.3500°, 14.8167°, 1202 m] (Bocage, 1867a, seen in trees, hence the English common name
1895, 1897a); Benguella [-12.5833°, 13.4167°, 15 “Ovambo Tree Skink,” although it also frequents
m] (Bocage, 1867a, 1867b, 1895, 1897a); Dombe boulders and koppies (Bauer et al. 1993). Bocage
[-12.9500°, 13.1000°, 50 m] (Bocage, 1867a, (1867a) notes that the species “lives close to
1895, 1897a). Huíla Province: Caconda human populations and dwells in ruins, wall
[-12.4333°, 13.5500°, 15 m] (Bocage, 1879a). holes or in retreats of small flying mammals.”
Namibe Province: Capangombe [-15.1000°,
13.1500°, 553 m] (Bocage, 1895). Trachylepis bocagii (Boulenger, 1887)—
Distribution in Angola: The species is Bocage’s Skink
widely distributed in the southwestern parts of Figures 30, 31, plate 2
the country, from Benguela to southern Bié
Province, southward through Huíla, Namibe, Euprepes Petersi Bocage, 1872: 74 [preoccupied
and Cunene provinces (fig. 28). by Euprepes petersi Steindachner, 1867].
FIGURE 27. Life photo of Trachylepis binotata from Maylowe, Namibe Province (CAS 266149). Photo by
L.M.P.C.
Holotype: MBL (specimen catalog number Mabuya bocagei: Frade (1963: 252).
unknown, destroyed by fire on March 18, Euprepes quinquetaeniata: Brygoo (1985: 90).
1978), collected by F.A.P. Bayão. Type Trachylepis bocagii: Marques et al. (2018: 258);
locality: “Duque de Bragança dans Branch et al. (2019: 318).
l’intérieur d’Angola” [= Calandula, Malanje
Trachylepis bocagii has a somewhat convo-
Province], Angola.
luted taxonomic and nomenclatural history that
Mabuia bocagii Boulenger, 1887: 203 [replace-
can be understood only in the context of the
ment name for Euprepes petersi Bocage,
authors at the time of its description. Shortly
1872]. after Bocage was entrusted with the position
Euprepes quinquetaeniatus [nec]: Bocage (1866a: of the director of the Zoological Section of the
44). National Museum of Lisbon, he started to receive
Mabuya Petersi: Bocage (1895: 42, 1897a: 197); specimens from the Portuguese overseas territo-
Ferreira (1900: 49, 1903: 15, 1906:170); Tie- ries in Africa (Ceríaco, 2021). As a competent
demann and Häupl (1980: 43); Tiedemann zoologist but still unfamiliar with the herpeto-
et al. (1994: 53). fauna of Africa, Bocage trusted in his colleagues
Mabuya quinquetaeniata: Boulenger (1905: 111). from other European museums in order to con-
Mabuya bocagii: Parker (1936: 136); Mertens firm his identifications and to provide helpful
(1938: 437); Bauer et al. (2003: 275). comments, in a kind of peer-review system. In
Mabuya quinquetaeniata quinquetaeniata: 1865 Bocage received several specimens from
Hellmich (1957b: 54). Duque de Bragança that were sent by the Por-
FIGURE 28. Distribution of Trachylepis binotata in Angola. Black stars denote the syntype localities.
tuguese colonial officer and amateur naturalist Museu Bocage (AHMB) in MUHNAC, Lisbon.
Francisco António Pinheiro Bayão (1833–1883) In a letter from Peters to Bocage dated July 7,
(Ceríaco, 2021; Ceríaco et al., in prep). Among 1869, the German zoologist noted to Bocage
these specimens, there were a considerable that the E. quinquetaeniatus specimens were not
number of specimens of what Bocage (1866a) conspecific with the type of the species (Peters,
considered Euprepes quinquetaeniatus (cur- July 7, 1869, in litt.). As an acknowledgment to
rently Trachylepis quinquetaeniata (Lichten- Peters, Bocage (1872) decided to describe it as a
stein, 1823)). He sent some of these specimens new species, naming it Euprepes Petersi. For this
to Wilhelm Peters (1815–1883) in Berlin, Franz description, Bocage provided the measurements
Steindachner (1834–1919) in Vienna, Auguste of a single specimen from the original series of
Duméril (1828–1870) in Paris, and Albert Gün- “Euprepes quinquetaeniata” sent to him by Pin-
ther (1830–1914) in London. These shipments heiro Bayão (Bocage, 1872). However, this name
are still recorded in the historical archives of was already preoccupied by E. petersii Stein-
FIGURE 29. Typical habitat of Trachylepis binotata in Capangombe, Namibe Province. Photo by L.M.P.C.
dachner, 1867 (currently a synonym of Eutropis author of the nomenclatural act at the time of
dissimilis (Hallowell, 1857)). description, but also any other specimens col-
Having recognized this situation, Boulenger lected on the same occasion and at the same
(1887) proposed Mabuia bocagii as a replace- locality.
ment name. He provided a completely new diag- In the type catalog of the Lisbon Museum,
nosis and listed three syntypes—one from Duque Bocage (1897a) referenced syntypes in the col-
de Bragança (the same specimen originally sent lections for Euprepes Petersi from Duque de Bra-
by Pinheiro Bayão to Bocage, and subsequently gança and from Dondo, although he had never
by the latter to Günther) and two others col- mentioned specimens from Dondo in his origi-
lected by the botanist Friedrich Welwitsch nal description. Broadley was able to locate only
(1806–1872) in “Pungo Andongo” and an two “syntypes” from Dondo (MBL 847a, 847b)
unspecified location in Angola. Since then, there during his visit to the Museu Bocage in the late
have been incongruities about the interpretation 1960s (Broadley, 1968, unpubl. data). Other
of the number of type specimens for this taxon. specimens wrongly labelled as syntypes are listed
This can be explained by the different interpreta- in the collections of NHMW (16677, from
tions of what a type series is. We follow a more Dondo), BMNH (1946.8.15.27, from Duque de
strict and exclusive interpretation, wherein only Bragança), MNHN (1286 and 1286a from Duque
such specimens or group of specimens that were de Bragança), and ZMB (6479, from Duque de
explicitly referenced in the text of the original Bragança) and represent those that Bocage
description are regarded as belonging to the type offered to the aforementioned naturalists.
series. On the other hand, Bocage and other Bocage’s (1872) description of Euprepes petersii
authors had a more open and inclusive interpre- contained measurements of only one specimen,
tation of what constituted a type series, which which would likely have been deposited in the
included not only the specimens analyzed by the Lisbon Museum and should be considered the
FIGURE 30. Specimen of Trachylepis bocagii from Kissama National Park, Luanda Province (CAS 263436).
Photos by L.M.P.C
holotype. As Broadley was unable to locate any therefore, should not be considered as part of the
specimen from Duque de Bragança in his visit to type series.
MBL in 1968 (Broadley, 1968, unpubl. data), it is Diagnosis: A medium-size skink (max. SVL
likely that the holotype could have been lost at 76.3 mm, MUNHAC/MB03-001341), with fully
that stage or was overlooked. In any case, all developed, pentadactyl limbs (fig. 30, 31); dorsal
specimens in MBL were destroyed by the fire of scales tricarinate to pentacarinate; ventral scales
1978. Several putative syntypes exist, especially smooth; 67–68 SAV; 55–61 SAD; 36–41 MSR;
considering Bocage’s interpretation of type series, lamellae beneath fingers and toes keeled and spi-
in which the holotype was part of a series of nose; plantar scales spinose; 21–27 LUFT; 13–20
“several specimens” collected by Bayão in Duque LUFF; supranasals always in contact; parietals
de Bragança and identified as Euprepes quinque- usually in contact; prefrontals usually separated;
taeniatus years before (Bocage, 1866a). This is frontoparietals always in contact; one pair of
the case for BMNH 1946.8.15.27 (formerly MBL, enlarged nuchal scales present; ear opening verti-
BMNH 66.6.11.7, collected by F.A.P. Bayão), sent cally ovoid and smaller than the eye; 3–4 subtri-
by Bocage to London in 1866 as E. quinquetae- angular auricular scales (shorter than the diameter
niatus. Given the fact that this specimen was sent of ear) extend posteriorly and usually slightly
to London six years before the description and upward from the anterior margin of the ear open-
identified as E. quinquetaeniatus, it is safe to ing. Supralabials usually eight, the sixth subocular;
assume that this was not in the hands of Bocage supraciliaries usually 5, the second the longest;
when he wrote the description of petersii and, nostrils oriented dorsally. Dorsum olive to dark
FIGURE 31. Life photo of Trachylepis bocagii from Kissama National Park, Luanda Province (CAS 263439).
Photo by John Cavagnaro.
brown with three white to yellowish stripes bor- m] (ZMB 6479*); Laúca Dam, flooded area
dered with black; vertebral stripe extends from the [-9.7627°, 15.1438°, 750 m] (MUNHAC/MB03-
nape to the base of the tail; dorsolateral stripes 001341); Top area of Kalandula falls [-9.0720°,
start behind the eye and continue along the tail; 16.0016°, 1073 m] (UF 187306); Pungo Andongo
there are usually thick black speckles between near footprints of the Queen [-9.6755°, 15.5836°,
dorsal stripes. Flanks usually darker, with a white 1107 m] (CAS 263592).
to yellowish stripe bordered with black, starting Additional Material: Benguela Province: Huxe
below the eye and extending to the hind limb [-12.7167°, 13.2000°, 65 m] (BMNH no number);
insertion; limbs uniform brownish above, some- Cubal [-13.0333°, 14.2500°, 921 m] (SMF 25716/8).
Kwanza Norte Province: Dondo [-9.6833°, 14.4333°,
times with pale spots on the thighs. Top of head
33 m] (NHMW 16677); Nova Oeiras [-9.4507°,
uniform brownish or with light grayish stippling; 14.4160°, 43 m] (ZMB 48887); Piri-Dembos [-8.5299,
labials cream-white, usually with brown to black 14.4377, 712 m] (ZMH no number); Piri-Dembos,
mottling. Venter whitish, sometimes with dark Roça Nova Douro [-8.5500°, 14.4333°, 630 m] (ZSM
spots or streaks under the tail. 69/1954). Kwanza Sul Province: Congulu, Amboim
Material Examined (* denotes type mate- [-10.8667°, 14.2833°, 640 m] (BMNH 1936.8.1.543–
546). Luanda Province: Kissama National Park, Bravo
rial): Luanda Province: Kissama National Park,
1 [-9.3255°, 13.2183°, 9 m] (AMB 12380); Kissama
KAWA camp headquarters [-9.1839°, 13.3722°, National Park, KAWA camp headquarters [-9.1839°,
151 m] (CAS 263434–263439; UF 187304, 13.3722°, 144 m] (AMB 12367, 12390, 12415, 12450–
187305). Malanje Province: Duque de Bragança 54, 12474); Kissama National Park, Romeo 1 [-9.1831°,
(currently Kalandula) [-9.0740°, 15.9999°, 1065 12.3700°, 62 m] (AMB 12458). Malanje Province:
FIGURE 32. Distribution map of Trachylepis bocagii in Angola. Black star denotes the type locality.
Duque de Bragança (currently Kalandula) [-9.1000°, [-9.6833°, 14.4333°, 33 m] (Bocage, 1872,

15.9500°, 1010 m] (BMNH 1946.8.15.27; TM 45460, 1895); Cambondo [-9.1596°, 16.6577°, 1145 m]
45461, 45463, 45479, 45492); N’Dalla Tando (= (Ferreira, 1906). Malanje Provice: Duque de
N’dalatando) [-9.3000°, 14.9167°, 782 m] (BMNH
Bragança [9.1000°, 15.9500°, 1010 m] (Bocage,
1909.10.29.101); Pungo Andongo [-9.6667°, 15.5833°,
1220 m] (BMNH 1864.7.13.35, 1904.5.2.37; ZMB
1866a, 1872, 1895); Pungo Andongo [-9.6667°,
9205). Undetermined locality: Unknown locality 15.5833°, 1220 m] (Bocage, 1895). Undeter-
(AMNH 48744–48745; MNHN 1866.71); Angola mined locality: Unknown locality (Ferreira,
(BMNH 1946.8.15.30*; MNCN 5788; MNHN 0.1286; 1900).
ZMH no number). Distribution in Angola: The species is
Historical Localities (no extant widely distributed in the western parts of the
specimens): Benguela Province: Quibula country, from Malanje to Luanda, Kwanza Norte,
[-12.2833°, 14.6833°, 1142 m] (Bocage, 1895, Kwanza Sul, and Benguela provinces (fig. 32).
1897a). Kwanza Norte Province: Dondo Global Distribution: Endemic to Angola.
FIGURE 33. Typical habitat of Trachylepis bocagii in Kalandula, Malanje Province. Photo by L.M.P.C.
Habitat and Natural History Notes: Holotype: An unsexed adult (CAS 263534,
This species is usually found in a mosaic of forest field number AMB 10366; fig. 34) collected at
and savannah, as well as miombo woodlands Caraculo, Namibe Province [-15.0165°,
(Grandvaux-Barbosa, 1970; fig. 33). 12.6425°, 489 m], by Luis M.P. Ceríaco, Suzana
A. Bandeira, and Ishan Agarwal on 23 Novem-
Trachylepis bouri, sp. nov.— ber 2016.
Bour’s Skink Paratypes: All from Angola. Two specimens:
an unsexed adult (CAS 85961, field number 603),
Figure 34, 35, plate 3 collected at 41 mi NE Mossamedes (= Caraculo)
[-14.7705°, 12.5942°, 530 m] by R. Leech and E.
Mabiua chimbana [part]: Bocage (1895: 45;
Ross on 22 May 1958; an unsexed adult (CAS
1897a: 195).
254903, field number JVV 8594), collected at
Trachylepis punctulata [part]: Ceríaco et al.
Namibe-Lubango road, marker 59, 1.8 km W of
(2016b: 31).
Caraculo, N side of road, Namibe Province
This newly described species was previously [-15.0153°, 12.6424°, 497 m] by Luis M.P.
misidentified by Ceríaco et al. (2016b) as T. punct- Ceríaco, Edward L. Stanley, Arianna Kuhn, Jens
ulata. According to our molecular results, this V. Vindum, Sango de Sá, Suzana A. Bandeira,
species is sister to a clade containing both topo- and Hilária Valério on 6 December 2013.
typical T. chimbana and T. bocagii (fig. 1). The Material Examined: Namibe Province:
species is known only from Namibe Province. Shortcut Road to Bibala from Namibe (9.24)
Historical records of Mabuia chimbana from [-15.0339°, 12.9858°, 380 m] (MUNHAC/MB03-
“Maconjo” and “Capangombe” (Bocage, 1895, 001510); Serra da Neve, Catchi surroundings
1897a) are provisionally assigned to this newly [-13.7620°, 13.2569°, 1585 m] (MUNHAC/
described species based on overall similarity. MB03-001511); Serra da Neve, rocky area near
FIGURE 34. Holotype of Trachylepis bouri, sp. nov., from Caraculo, Namibe Province (CAS 263534). Photos
by L.M.P.C.
base camp [-13.7653°, 13.2571°, 1645 m] (MUN- developed, pentadactyl limbs (figs. 34, 35); dor-
HAC/MB03-001512). sal scales pentacarinate; ventral scales smooth;
Additional Material: Namibe Province: 50 km 54–60 SAV; 50–54 SAD; 36–38 MSR; lamellae
West of Humpata, by roadside in boulders [-15.0507°, beneath fingers and toes keeled and spinose;
13.0279°, 464 m] (PEM R17961); along fence leading to
Baptista farm (Omauha Lodge) [-15.9968°, 12.4068°,
plantar scales spinose; 21–23 LUFT, 15–17 LUFF;
301 m] (PEM R17962); Assunção [-14.8667°, 13.1000°, supranasals always in contact; parietals always in
505 m] (TM 40164, 40165); Cainde [-15.4833°, contact; prefrontals usually separated; frontopa-
13.3667°, 631 m] (TM 40956, 40957); Caraculo rietals always in contact; one pair of enlarged
[-15.0167°, 12.6667°, 463 m] (TM 40191, 40233, 46729, nuchal scales present; ear opening vertically
46730), [-15.0165°, 12.6425°, 464 m] (AMB 13024,
ovoid and smaller than the eye; three subtrian-
13035); Saiona River 25 km N of Cainde [-15.4000°,
13.2000°, 534 m] (TM 40982); Munhino [-14.9788°, gular auricular scales (shorter than the diameter
12.9781°, 415 m] (AMB 13048, 13055, 13056). of ear) extend posteriorly and usually slightly
Historical Localities (no extant speci- upward from the anterior margin of the ear
mens): Namibe Province: Maconjo [-15.0167°, opening. Supralabials usually eight, the sixth
13.2000°, 865 m] (Bocage, 1895, 1897a); Capan- being subocular; five supraciliaries, the second
gombe [-15.1000°, 13.1500°, 553 m] (Bocage, notably longer; nostrils oriented dorsally. Dor-
1895, 1897a). sum bronze to golden-brown, uniform or with
Diagnosis: A small-sized skink (max. SVL scattered black speckles; flanks mottled with
56.3 mm, MUNHAC/MB03-001511), with fully black and white, forming a band that extends
FIGURE 35. Life photo of Trachylepis bouri from Namibe Province. Note the flanks mottled with black and
white, forming an irregular stripe that extends from the eye to the hind limb insertion, and compare to its
sister species, T. chimbana, in fig. 39. Photo by L.M.P.C.
from the eye to the hind limb insertion; a pair of tact with loreal scale. Prefrontals pentagonal,
weakly defined pale dorsolateral stripes may be contacting in a single point, each in contact with
present. Labials white, sometimes with black the following head shields: frontonasal, loreal,
spots or vertical bars; head uniform or with scat- first and second supraocular, and frontal. Two
tered black spots. Venter uniformly white. loreals. Frontal length similar to the distance
Description of the Holotype: A well-pre- between anterior tip of frontal and tip of snout.
served, unsexed adult. Body cylindrical and Frontal in contact with two supraoculars on each
robust with a poorly defined neck and well- side. Two frontoparietals, in contact with each
developed pentadactyl limbs; tail truncated. other and the frontal, third and fourth supraocu-
Fore- and hind limbs overlap when adpressed lars, parietal, and interparietal. Frontoparietal
against the body. SVL 42.3 mm. HL 11.1 mm, plus interparietal length similar to frontal length.
with relatively long and prominent snout. Addi- Interparietal twice as long as broad, with a visible
tional measurements are presented in table 5. parietal foramen. Parietals greater than frontopa-
Three subtriangular auricular scales extend pos- rietals. Parietals in contact with each other. Five
teriorly from the anterior margin of the ear supraciliaries, second largest. Eight supralabials,
opening. Rostral visible from above. Nostrils ori- sixth being subocular. Eight infralabials. Post-
ented dorsally and set posteriorly so that postna- mental bordering seven scales (mental, two
sal effectively borders nostril. Supranasals in infralabials on each side and two primary chin
contact. Frontonasal broader than long, in con- shields). Transparent scale present in lower eye-
TABLE 5
Mensural and Meristic Data for the Type Series of Trachylepis bouri, sp. nov.
CAS 263534 CAS 254903 CAS 85961

Holotype Paratype Paratype
Sex Unsexed unsexed unsexed
SVL 42.3 53.5 41
TL – – 54
TL/SVL – – 130
HL 11.1 12.8 9.2
HL/SVL 30 20 20
SVL/HL 380 420 450
HW 6.5 7.7 5.6
HW/HL 60 60 60
HH 4.5 5.6 4.3
IN 1.3 1.7 1.2
EN 3.6 3.6 3.5
ES 4.4 5.1 4.1
MSR 36 36 36
SAD 52 50 50
SAV 54 58 54
LUFF 17 17 17
LUFT 22 23 23
SC 5 5 5
SL (SO) 8 (6) 8 (6) 8 (6)
CP C C C
CFP C C C
CSN C C C
CPF S C S
KDS 5 5 5
Plantar scales Smooth smooth smooth
lid, as is usual for Trachylepis. Tympanum visible, Coloration in Ethanol: Background

at same level as mouth. Dorsal scales each with color of flanks and upper side of head, neck,
three smooth keels. Ventral scales smooth. MSR dorsum, legs and tail homogenously brown,
36, SAD 52, SAV 54. Limbs with five digits; with some white speckles running from the
scales on palms and soles keeled. Relative length labials along the body to the anterior half of
of fingers IV > III > V > II > I, relative length of the tail. Above the labials the head is uni-
toes IV > III > V > II > I. Finger-IV lamellae 17, formly brown. Eyelids dark brown. Ventral
Toe-IV lamellae 22. surfaces uniformly whitish with some scat-
FIGURE 36. Distribution of Trachylepis bouri in Angola. Black star denotes the type locality.
tered dark speckles near the flanks and under sulcata, T. ansorgii, T. albopunctata, and T. dama-
the tail. rana by having spinose plantar scales. It differs
Variation: Variation in scalation and mea- from T. wahlbergi and T. attenboroughi, by the
surements among the type series is reported in presence of subtriangular auricular scales on the
table 5. Both paratypes generally agree with the anterior margin of the ear opening (vs. absent in
holotype in terms of coloration. the latter). T. bouri is readily distinguished from T.
Comparison with Other Angolan and suzanae and T. wilsoni, which have wedged-
Southwest African Trachylepis: Trachylepis shaped snouts. It differs from T. hilariae by having
bouri, sp. nov., differs from all the other species of a higher number of MSR (36–38 in T. bouri vs.
Trachylepis known to occur in Angola, with the 29–30 in T. hilariae), from T. punctulata by having
exception of T. wahlbergii, T. attenboroughi, T. 50–54 SAD (vs. 42–50 in the latter) and by having
chimbana, T. punctulata, T. hilariae, T. bocagii, T. a speckled pattern (vs. mostly uniform dorsum in
ovahelelo, T. suzanae, T. vunongue, T. wilsoni, T. topotypical T. punctulata), and from T. vunongue
FIGURE 37. Typical habitat of Trachylepis bouri in Caraculo, Namibe Province. Photo by Arianna Kuhn.
by having 36–38 MSR and 15–17 LUFF (vs. 30–35 Distribution in Angola: The species is
MSR and 9–14 LUFF in the latter). It differs from known only from central and northern Namibe
T. bocagii by absence of dorsal bands and from T. Province (fig. 36).
ovahelelo by presence of a dark lateral patch. T. Global Distribution: So far only known
bouri differs from T. ansorgii, T. sulcata, T. albo- from Angola. Haacke’s (1997) records from
punctata and T. damarana by having its nostrils northern Namibia may belong to T. bouri, but
situated more dorsally, directed upward (vs. nos- these specimens need to be reviewed.
trils situated more laterally, directed sideward in Habitat and Natural History Notes:
the latter). In comparison to its sister taxon T. This species is associated with the dry habitats of
chimbana, T. bouri has an extensive lateral patch Namibe Province, inhabiting dry mopane wood-
of black and white speckles extending almost to lands and savannah-steppe mosaic (Grandvaux-
the hind limb insertion area (vs. a more limited Barbosa, 1970; fig. 37).
and subtle speckling, not extending past midbody Etymology: The specific epithet “bouri”
in T. chimbana). Regarding the remainder of the is formed in the genitive singular and is mas-
southwest African congeners of the variegata sub- culine. It is given in honor of French herpe-
group sensu Weinell et al. (2019), T. bouri is read- tologist Roger Henri Bour (1947–2020). This
ily differentiated from T. variegata by having five recognition is due to the inspiration, friend-
keels on dorsal scales (vs. three in T. variegata). ship, and support for the early herpetologi-
FIGURE 38. Specimen of Trachylepis chimbana from N’Dolondolo, Namibe Province (CAS 263542). Photos
by L.M.P.C.
cal career of the two first authors. We suggest Mabuia chimbana [part]: Bocage (1895: 45,
“Bour’s Skink” and “Lagartixa de Bour” as 1897a: 195); Frade (1963: 252).
the English and Portuguese common names, Mabuya bocagii bocagii: Hellmich (1957a: 60).
respectively, for this species. Mabuya striata angolensis: Hellmich (1957b: 56).
Mabuya chimbana: Broadley (1974b: 13); Branch
Trachylepis chimbana (Boulenger, 1887)— (1998: 153).
Chimba Skink Trachylepis chimbana [part]: Portik and Bauer
(2012: 128), Ceríaco et al. (2016b: 57, 2020a:
Figures 38, 39, plate 3 402); Marques et al. (2018: 259); Branch et
Euprepes affinis [nec] Bocage, 1872: 77 [preoc- al. (2019a: 318).
cupied by Tiliqua affinis Gray, 1838]. Syn- This small-sized Angolan endemic has been
types: MBL 822 (2 specimens), 824, both among the most taxonomically problematic spe-
destroyed by fire on March 18, 1978. Type cies. It was originally described by Bocage (1872)
locality: “Rio Chimba, dans l’intérieur de as Euprepes affinis, based on an unspecified num-
Mossamedes” [= Rio Chimba (currently ber of specimens (although Bocage provides the
Bentiaba), Namibe Province], Angola. maximum size of a single specimen) collected by
Mabuia chimbana Boulenger, 1887: 204 [replace- José d’Anchieta in “Rio Chimba” (currently Ben-
ment name for Euprepes affinis Bocage, tiaba), in northern part of Namibe province. The
1872]. name affinis was preoccupied by Tiliqua affinis
FIGURE 39. Life photo of Trachylepis chimbana from N’Dolondolo (CAS 263543). Photo by Ishan Agarwal.
Gray, 1838 (currently Trachylepis affinis (Gray, of chimbana but included among his analysis
1838)), which led Boulenger (1887) to provide a specimens from southern Namibe province,
replacement name—Mabuia chimbana—that has namely those from Assunção, 10 km E of Caem-
remained in use since then (Bocage, 1895, 1897a; bombo, Cainde, Caraculo, 14 km NE of Cara-
Broadley, 1974b; Branch, 1998; Portik and Bauer, culo, Coporolo, 12 km W of Humbia, Huke,
2012; Ceríaco et al., 2016a, 2021; Marques et al., Maconjo, and Saiona River, NW of Cainde,
2018; Branch et al., 2019a). Other authors who which belong to its recently recognized sister
had worked with Angolan skinks, such as species T. bouri (see account above).
Hellmich (1957a), Schmidt (1933) and Laurent Our morphological and molecular results
(1964), have confounded the taxon with others, restrict the distribution of the species to
such as Trachylepis bocagii, T. wahlbergi, or T. northern Namibe and southern Benguela
attenboroughi (see respective accounts). provinces. Records from the northeastern
Broadley (1968, personal commun.) exam- provinces of Moxico and Lunda Sul provided
ined the then extant syntypes in the collections by Laurent (1964) represent T. wahlbergii.
of Museu Bocage, Lisbon, before fire destroyed The record from Humbe (Schmidt, 1933) rep-
the collections in 1978. According to his obser- resents T. attenboroughi and the specimens
vations, the type series had 34–38 scale rows from Humbia cited by Baptista et al. (2019)
around the midbody, 5–7 keels on the dorsal likely also represent T. bouri. Records from
scales, 22–24 lamellae under the fourth toe, 3–4 Namibia (Haacke, 1997; Branch, 1998) likely
ear lobules, with a maximum size (SVL + TL) of represent a misidentification with T. bouri or
57 mm. Broadley (1974b) confirmed the validity another small and putatively undescribed
FIGURE 40. Distribution of Trachylepis chimbana in Angola. Black star denotes the type locality.
species of Trachylepis. These specimens need vertically ovoid and smaller than the eye; three
to be reviewed to establish their identity. subtriangular auricular scales (shorter than the
Diagnosis: A small skink (max. SVL 56.0 diameter of ear) extend posteriorly and usu-
mm, AMNH 40657), with fully developed ally slightly upward from the anterior margin
pentadactyl limbs (fig. 38, 39); dorsal scales of the ear opening. Supralabials usually eight,
usually pentacarinate; ventral scales smooth; the sixth being subocular; five supraciliaries,
55–63 SAV; 49–56 SAD; 33–39 MSR; lamellae the second longest; nostrils oriented dorsally.
beneath fingers and toes keeled and spinose; Dorsum bronze to grayish brown, uniform or
plantar scales spinose; 18–24 LUFT; 15–17 with scattered black speckles; anteriorly flanks
LUFF; supranasals always in contact; parietals usually with mottling of black and white,
usually in contact; prefrontals always separated; forming a band that extends from the eye to
frontoparietals always in contact; one pair of the forelimb insertion or midbody; a pair of
enlarged nuchal scales present; ear opening weakly defined pale dorsolateral stripes may be
FIGURE 41. Typical habitat of Trachylepis chimbana, south of Bentiaba, Namibe Province. Photo by L.M.P.C.
present. Labials white, sometimes with black 12.5269°, 6 m] (TM 41187); dirt road to Chingo [-14.3934°,
spots or vertical bars; head uniformly colored 12.8289°, 620 m] (CAS 264272). Undetermined locality:
or with scattered black spots. Venter uniformly Unknown locality (AMNH 48663–48674).
white. Historical Localities (no extant speci-
Material Examined: Benguela Province: mens): Benguela Province: Quindumbo
Hanha [-12.2506°, 13.7116°, 83 m] (AMNH [-12.4667°, 14.9333°, 1462 m] (Bocage, 1895,
40656–40662). Namibe Province: vic. 1897a). Namibe Province: Rio Chimba [-14.3000°,
N’Dolondolo [-13.8105°, 13.1361°, 707 m] 12.4000°, 204 m] (Bocage, 1872, 1895, 1897a).
(CAS 263542, 263543, 263544); Serra da Neve Distribution in Angola: The species is
[-13.8105°, 13.2581°, 1502 m] (CAS 263562, known to occur only n Huambo, southern
263563); base of Serra da Neve, Maylowe village Benguela, and northern Namibe provinces
[-13.8355°, 13.2755°, 798 m] (MUNHAC/ (fig. 40). Extralimital records likely represent
MB03-001387); Serra da Neve base, 2km N of misidentifications. The species is replaced in
Maylowe [-13.8280º, 13.2625º, 818 m] (MUH- the central and southern parts of Namibe
NAC/MB03-001518); road from Camucuio to Province by its sister species, T. bouri, reflect-
Serra da Neve [-14.0227°, 13.1941°, 665 m] ing a distributional turnover observed in other
(MUNHAC/MB03-001513); Serra da Neve, reptile taxa in the region (e.g., Parrinha, et al.
Catchi surroundings [-13.7619°, 13.2569°, 1585 2021; Marques et al. 2022).
m] (MUNHAC/MB03-001514); Serra da Neve, Global Distribution: Endemic to Angola.
rocky area near base camp [-13.7653°, 13.2571°, Habitat and Natural History Notes:
1645 m] (MUNHAC/MB03-001515, 001519). This rupiculous species is usually found on gran-
Additional Material: Benguela Province: Entre- ite outcrops in a variety of habitats, from mopane
Rios [-13.0167°, 14.6333°, 1267 m] (ZSM 101/1953). and miombo woodlands and shrubland to savan-
Huambo Province: Sanguengue [-12.5667°, 16.2333°, 1806 nah-steppe mosaic (Grandvaux-Barbosa, 1970;
m] (ZSM 171/1954). Namibe Province: Lucira [-13.8654°, Broadley, 1974b; fig. 41).
FIGURE 42. Specimen of Trachylepis damarana from Munguenga, Cuando Cubango Province (PEM R20507).
Photos by Werner Conradie.
Trachylepis damarana (Peters, 1870)— Cuando Cubango province as T. varia, but

Damaraland Skink Weinell and Bauer (2018) later confirmed that
these represented true T. damarana. So far,
these are the only confirmed records of the
Euprepes damaranus Peters, 1870: 20. Lecto- species in the country.
type (designated by Weinell and Bauer, Diagnosis: A medium-sized skink (max.
2018): ZMB 6153 (collected by Johan SVL on Angolan specimens 56.2 mm, PEM
August Wahlberg). Type locality: R20514; max. SVL of extralimital populations
“Damaraland,” Namibia. 67.6 mm, see Weinell and Bauer, 2018), with
Trachylepis varia: Conradie et al. (2016: 26). fully developed pentadactyl limbs (figs. 42, 43);
Trachylepis damarana: Weinell and Bauer (2018: dorsal scales tricarinate or quadricarinate; ven-
215); Marques et al. (2018: 260); Branch et tral scales smooth; 50–55 SAV [40–56 in extra-
al. (2019: 318); Conradie et al. (2022: 203). limital populations, see Weinell and Bauer,
2018]; 40–43 SAD; 31–34 MSR [30–36 in extra-
This species has only recently been con- limital populations, see Pietersen et al., 2021];
firmed in Angola. Conradie et al. (2016) pre- lamellae beneath fingers and toes keeled and
liminarily identified some specimens from spinose; plantar scales spinose; 14–17 LUFT;
FIGURE 43. Life photo of Trachylepis damarana from North of Rito, Cuando Cubango Province. Photo by
Werner Conradie.
19–24 LUFF [up to 26 in extralimital popula- stripe starts at the nostril and usually becomes
tions, see Weinell and Bauer, 2018]; supranasals more reticulated posteriorly, often forming
always in contact; parietals always in contact at irregular blotches or vertical bars on the upper
a single point; prefrontals usually touching at a flanks. Limbs variegated above, with irregular
single point; frontoparietals always in contact; dark and pale speckles. Top of head uniformly
one pair of enlarged nuchal scales present; ear brown or with light dark stippling; labials white,
opening vertically ovoid and smaller than the sometimes with dark vertical bars on the poste-
eye; 2–4 subtriangular auricular scales (shorter rior margin. Venter white without markings.
than the diameter of ear) extend posteriorly and Material Examined: Cuando Cubango
usually slightly upward from the anterior mar- Province: 1st dry pan 13 km from Chetto Angola
gin of the ear opening. Supralabials usually Border [-17.6783°, 22.6147°, 1021 m] (PEM
seven, the fifth being subocular; five supracili- 20511, 20512); 2nd dry pan 25 km from Chetto
aries, the second longest; nostril oriented later- Angola Border [-17.5883°, 22.6569°, 1004 m]
ally. Dorsum olive brown, with a pair of faint (PEM R20513); camp site 15 km North of
pale dorsolateral stripes; between the stripes Mavengue [-17.0488°, 19.5344°, 1086 m] (PEM
there is usually heavy black speckling with ten- R20510); Camp site 6 km west of Sasha E, Jamba
dency to form transverse bars, starting behind [-17.5733°, 23.2267°, 991 m] (PEM R21489);
the neck and extending extending along the tail. Camp site 6 km west of Sasha E, Jamba
A bright white stripe starts at the subocular and [-17.5733°, 23.2600°, 987 m] (PEM R20516); pan
extends through the flanks to the hind-limb 18 km East of Jamba [-17.4633°, 22.8664°, 995
insertion, usually bordered with black; a black m] (PEM R20514); Lagoon 6 km SE of Bonavae
FIGURE 44. Distribution map of Trachylepis damarana in Angola.
(Jamba) [-17.4961°, 23.1344°, 980 m] (PEM 19.2806°, 1241 m] (PEM R27428–9, 27432–3; INBAC/
R20515); Trap array near flooded rocky drossy to WC-6769); Quembo River bridge camp, trap 3
Mpupa falls (Munguenga) [-17.5122°, 20.0602°, [-13.5278°, 19.2745°, 1256 m] (PEM R27434, 27436);
left side tributary (Condinde River) at Cuando River
1079 m] (PEM R20507); 8.5 km North of Rito
bridge [-13.6007°, 19.5267°, 1219 m] (PEM R27435);
(Rito) [-16.6232°, 19.0535°, 1155 m] (PEM Luvu River camp [-13.7120°, 21.8353°, 1082 m] (PEM
R20508, 20509). Moxico Province: en route to R27437).
Cuanavale River Source [-12.7237°, 18.6228°, Distribution in Angola: The species is only
1355 m] (PEM R23266). known from the province of Cuando Cubango,
Additional Material: Moxico Province: camp at
southeastern Angola (fig. 44).
side tributary (Luandai River) of the Luanguinga River
[-13.7089°, 21.2623°, 1116 m] (PEMR27425–6, 27430);
Global Distribution: Trachylepis dama-
Lake Hundo [-14.9743°, 21.6297°, 1100 m] (PEM rana is known to occur in southeastern Angola
R27427, 27431); Quembo River bridge camp [-13.5275°, and northeastern Namibia, westward through
FIGURE 45. Typical habitat of Trachylepis damarana in Cuando Cubango Province. Photo by Werner
Conradie.
western Zambia, northern Botswana, Zimbabwe, Holotype: An unsexed adult (CAS 254775,
to northeastern South Africa and western field number JVV 8721; fig. 46) collected at
Mozambique (Weinell and Bauer, 2018). Tombwa dunes, Namibe Province [-16.2776°,
Habitat and Natural History Notes: 11.8224°, 11 m], by Luis M.P. Ceríaco, Edward L.
This species is often found near dry pans in a Stanley, Arianna Kuhn, Jens V. Vindum, Sango
mosaic of savannah and miombo woodlands de Sá, Suzana A. Bandeira, and Hilária Valério,
(Grandvaux-Barbosa, 1970; Conradie et al., on 8 December 2013.
2016; fig. 45). Paratypes: All specimens from Angola.
Three unsexed adults: (CAS 254769, 254770,
Trachylepis hilariae, sp. nov.—Hilária’s Skink 254771, field numbers JVV 8713, 8715, 8716,
Figure 46, plate 3 respectively), same data as holotype.
Diagnosis: A small-sized skink (max. SVL
Trachylepis punctulata [part]: Ceríaco et al. 43.1 mm, CAS 254770), with fully developed,
(2016b: 31). pentadactyl limbs (fig. 46); dorsal scales penta
carinate; ventral scales smooth; 58 SAV; 46–48
This presently described species was first SAD; 29–30 MSR; lamellae beneath fingers and
reported by Ceríaco et al. (2016b) as T. punctu- toes keeled and spinose; plantar scales spinose;
lata. Our molecular results indicate that these 21–22 LUFT, 15–16 LUFF; supranasals in con-
specimens belong to a taxon that is sister to a tact; parietals in contact; prefrontals separated;
clade comprising true T. punctulata and T. cf. frontoparietals in contact; one pair of enlarged
triebneri from Namibia, and closely related to T. nuchal scales present; ear opening vertically
variegata from Namibia, T. ovahelelo, and T. ovoid and smaller than the eye; three subtrian-
vunongue (fig. 1). It is known only from a small gular auricular scales (shorter than the diameter
area south of Tombwa in Namibe Province. of ear) extend posteriorly and usually slightly
FIGURE 46. Holotype of Trachylepis hilariae, sp. nov., from Tombwa, Namibe Province (CAS 254775). Photos
by L.M.P.C.
upward from the anterior margin of the ear developed pentadactyl limbs; tail long, its
opening. Supralabials eight, the fifth being length greater than the SVL, smoothly tapering.
subocular; five supraciliaries, the second longest; Fore- and hind limbs overlap when adpressed
nostril oriented dorsally. Dorsum and upper against the body. SVL 37.2 mm, TL 55.3 mm.
flanks grayish brown, with series of dark spots or HL 8.8 mm, with relatively long and thin snout.
dashes with tendency to form longitudinal lines, Additional measurements are presented in table
usually prolonged through the tail; limbs with a 6. Ear opening medium sized. Three subtrian-
variegated aspect above, with black edged scales gular auricular scales extend posteriorly from
and scattered pale spots. Top of head brown, the anterior margin of the ear opening. Rostral
with light grayish speckling; anterior labials pale visible from above. Nostrils oriented dorsally
brown to cream-white; there may be a more or and set posteriorly, so that postnasal effectively
less distinct pale stripe from the subocular to the borders nostril. Supranasals in contact. Fronto-
forelimb insertion, bordered with black. Venter, nasal broader than long, in contact with loreal
gular region and lower flanks grayish to bluish, scale. Prefrontals pentagonal, contacting at a
often with grayish speckling near the flanks. single point, each in contact with the following
Description of the Holotype: A well- head shields: frontonasal, loreal, first and sec-
preserved, unsexed adult. Body cylindrical and ond supraocular, first and second supraciliary
robust with a poorly defined neck and well- and frontal. Two loreals. Frontal length similar
TABLE 6
Mensural and Meristic Data for the Type Series of Trachylepis hilariae, sp. nov.
CAS 254775 CAS 254769 CAS 254770 CAS 254771

Holotype Paratype Paratype Paratype
Sex unsexed unsexed unsexed unsexed
SVL 37.2 32.8 43.1 33.4
TL 55.3 53 47 50
TL/SVL 150 160 110 150
HL 8.8 7.9 8.9 8.3
HL/SVL 20 20 20 20
SVL/HL 420 420 480 400
HW 5.4 4.7 5.9 4.8
HW/HL 60 60 60 60
HH 3.8 3.3 4.1 3.4
IN 1.1 1 1.2 1
EN 2.5 2.4 2.7 2.5
ES 3.6 3.3 3.6 3.6
MSR 30 29 29 30
SAD 46 47 48 46
SAV 58 58 58 58
LUFF 16 16 15 16
LUFT 22 21 22 21
SC 5 5 5 5
SL (SO) 8 (5) 8 (5) 8 (5) 8 (5)
CP C C C C
CFP C C C C
CSN C C C C
CPF S S S S
KDS 5 5 5 5
Plantar scales spinose spinose spinose spinose
to the distance between anterior tip of frontal Parietals greater than frontoparietals. Parietals
and tip of snout. Frontal in contact with two in contact with each other. Five supraciliaries,
supraoculars on each side. Two frontoparietals, second largest. Eight supralabials, fifth being
in contact with each other and the frontal, third subocular. Six infralabials. Postmental border-
and fourth supraoculars, parietal, and interpa- ing seven scales (mental, two infralabials on
rietal. Frontoparietal plus interparietal length each side and two primary chin shields). Trans-
similar to frontal length. Interparietal twice as parent scale present in lower eyelid, as is usual
long as broad, with a visible parietal foramen. for Trachylepis. Tympanum visible, at same level
FIGURE 47. Distribution of Trachylepis hilariae in Angola. Black star denotes the type locality.
as mouth. Dorsal scales each with three smooth tered pale and dark speckles. Top of head bluish
keels. Ventral scales smooth. MSR 30, SAD 46, posteriorly, with light grayish speckling; anterior
SAV 58. Limbs with five digits; scales on palms labials brownish, subocular bluish; a dark stripe
and soles spinose. Relative length of fingers IV starts behind the eye and extends to the forelimb
> III > V > II > I, relative length of toes IV > III insertion, becoming reduced to scattered spots
> V > II > I. Finger-IV lamellae 16, Toe-IV posteriorly. Lower flanks, venter, and gular
lamellae 22. region blueish, with light grayish speckling near
Coloration in Ethanol: Background color the flanks; underside of limbs and tail whitish,
of flanks, upper side of head, neck, dorsum, legs, with light grayish speckling under the tail.
and tail grayish brown, with series of dark spots Variation: Variation in scalation and mea-
that tend to form longitudinal lines from the surements among the type series is reported in
nape to the tail; limbs variegated above with scat- table 6.
FIGURE 48. Typical habitat of Trachylepis hilariae in Tombwa Dunes, Namibe Province. Photo by L.M.P.C.
Comparison with Other Angolan and situated more laterally, directed sideward in the
Southwest African Trachylepis: Trachylepis latter). It can be distinguished from T. chimbana,
hilariae, sp. nov., differs from all other species of T. bouri, T. ovahelelo, and T. bocagii by having
Trachylepis known to occur in Angola, with the <31 MSR (vs. >32 in the latter). In comparison
exception of T. albopunctata, T. damarana, T. with its sister taxon T. punctulata, T. hilariae
chimbana, T. bouri, T. bocagii, T. attenboroughi, presents a grayish-bluish venter (vs. uniformly
T. wahlbergii, T. sulcata, and T. ansorgi, by having white in T. punctulata). It differs from T.
spinose plantar scales. It differs from T. attenbor- vunongue by having 29–30 MSR (vs. 30–35 in the
oughi, and T. wahlbergii by the presence of sub- latter), 58 SAV (vs. 45–55) and 15–16 LUFF (vs.
triangular auricular scales on the anterior margin 9–14). Regarding the other southwestern African
of the ear opening (vs. absent in the latter). Tra- congeners of the variegata subgroup sensu Wei-
chylepis hilariae is readily distinguished from T. nell et al. (2019), T. hilariae is readily differenti-
suzanae and T. wilsoni by not having a wedge- ated from T. variegata by having five keels on
shaped snout (vs. wedge-shaped in the latter dorsal scales (vs. three in T. variegata).
two). The new species can be readily distin- Distribution in Angola: This species is
guished from T. albopunctata, T. damarana, T. known only from a small area south of Tombwa,
ansorgii, and T. sulcata by having its nostrils situ- Namibe Province (fig. 47).
ated more dorsally, directed upward (vs. nostrils Global Distribution: Endemic to Angola.
FIGURE 49. Specimen of Trachylepis hoeschi from Virei-Calundolo, Namibe Province (CAS 263483). Photos
by L.M.P.C.
Habitat and Natural History Notes: Trachylepis hoeschi (Mertens, 1954)—

This species is found in the mobile sand dunes of Hoesch’s Skink
the Namib Desert in southwestern Angola
(Grandvaux-Barbosa, 1970; fig. 48).
Etymology: The specific epithet “hilariae” Mabuya hoeschi Mertens, 1954: 178. Holotype:
is formed in the genitive singular and is femi- SMF 45681 (collected by W. Hoesch). Type
nine. It is given in honor of Hilária Valério locality: “Roessing-Berge, östlich von
(1991–), Angolan herpetologist and professor Swakopmund, SW-Afrika” [= Rössing Mts.,
at Agostinho Neto University’s department of east of Swakopmund, Erongo Region],
biology in Luanda. Hilária has been a funda- Namibia.
mental part of our team since the beginning of Mabuya hoeschi: Laurent (1964: 68); Branch
our joint herpetological studies in the country. (1998: 153).
Hilária initiated her scientific career collecting Trachylepis hoeschi: Ceríaco et al. (2016b: 32,
Trachylepis and other reptiles in the Namibe 57); Marques et al. (2018: 261); Baptista et
desert in 2013 and is now an established and al. (2018: 402); Branch et al. (2019a: 318);
respected member of the Angolan scientific Ceríaco et al. (2020a: 402); Lobón-Rovira et
community. We suggest “Hilária’s Skink” and
al. (2022: 309).
“Lagartixa de Hilária Valério” as the English
and Portuguese common names, respectively, Laurent (1964) provided the first record of
for this species. this species for Angola based on a single speci-
FIGURE 50. Life photo of Trachylepis hoeschi from Munhino, Namibe Province (AMB 13048). Photo by
L.M.P.C.
men collected by António de Barros Machado in smooth; plantar scales smooth; 18–19 LUFT,
“Plage das Conchas, Moçâmedes.” This new 13–15 LUFF; supranasals usually in contact;
record represented a considerable extension to parietals always separated; prefrontals always
the species’ known distribution range, which at separated; frontoparietals always in contact; one
the time was only known from Namibia. Addi- pair of enlarged nuchal scales present; ear open-
tional material was collected from Iona National ing vertically ovoid, smaller than the eye; subtri-
Park by Wulf Haacke in the 1970s and deposited angular auricular scales on the anterior margin
in the Ditsong National Museum of Natural His- of the ear opening absent. Ten supralabials;
tory. New records of this species were recently subocular not reaching the lip; supraciliaries
published by Ceríaco et al. (2016b) from Pediva usually five, the second longest. Dorsum grayish
Hot Springs and Namibe Reserve. brown, with series of dark brown blotches form-
Diagnosis: A medium-sized skink (max. ing transverse bars that may extend to the flanks,
SVL 76.5 mm, CAS 254851, max. SVL of extra- interrupted by two to three pale longitudinal
limital populations 100 mm, see Branch, 1998), stripes. Belly, throat, and lower flanks white.
with fully developed, pentadactyl limbs (figs. 49, Scales on palms and soles with dark brown
50); dorsal scales tricarinate; ventral scales tubercules along the digits, except for the distal
smooth; 55–62 SAV; 43–48 SAD; 33 scales rows subdigital lamellae (usually three to five). Juve-
around the midbody [starting at 32 in extralim- niles have a more contrasting pattern, with scat-
ital populations, see Branch, 1998]; lamellae tered white speckles on the back and irregular
beneath fingers and toes both keeled and dark mottling on the hind limbs, while adults
FIGURE 51. Distribution of Trachylepis hoeschi in Angola.
are usually duller, with the pale vertebral stripe [-16.6849°, 12.8413°, 584 m] (AMB 13085, 13086,
often indistinct. Labials white. 13097, 13132, 13136, 13137, 13149–51); Vipungos
Material Examined: Namibe Province: [-15.0439°, 12.4234°, 333 m] (AMB 13065); Baynes
[-17.0161°, 12.9314°, 349 m] (AMB 13119, 13120);
Iona National Park, Curoca River, Pediva Hot
Curoca river near Omauha [-16.2555°, 12.3327°, 259
Springs area [-16.2836°, 12.5611°, 250 m] (CAS m] (AMB 13266); Epupa [-16.9943°, 13.2515°, 649 m]
254851); Namibe Reserve [-15.7739°, 12.3331°, (AMB 13183, 13191, 13207).
263 m] (CAS 254852); Virei-Calundolo Distribution in Angola: In Angola, the
[-16.3102°, 12.7950°, 471 m] (CAS 263483). species is known only from central and southern
Additional Material: Namibe Province: Plage
Namibe Province (fig. 51). Baptista et al. (2018)
«das Conchas», Moçâmedes [-15.1333°, 12.1167°, 16
m] (MD 1932); Namibe road to Tambor, boulders on
recorded the species from the Tundavala region.
side of road, in dry river bed [-15.8761°, 12.2058°, 189 Global Distribution: Endemic to north-
m] (PEM R17951); Iona National Reserve [-16.9000°, western Namibia and southwestern Angola
12.5833°, 1000 m] (TM 40733–40737); Otchifengo (Branch, 1998).
FIGURE 52. Typical habitat of Trachylepis hoeschi, Pediva Hot Springs, Namibe Province. Photo by L.M.P.C.
Habitat and Natural History Notes: Mabuya bayoni huilensis: Ceríaco et al. (2020a:
This species is usually found among boulders 421).
and rocky outcrops in a mosaic of steppe, arid Trachylepis varia: Ceríaco et al. (2016b: 31).
savannah and mopane woodlands (Grandvaux- Trachylepis bayonii [part]: Marques et al. (2018:
Barbosa, 1970; fig. 52). 256); Ceríaco et al. (2020a: 422).
Trachylepis bayoni huilensis: Baptista et al. (2018:
Trachylepis huilensis (Laurent, 1964)— 400); Branch et al. (2019a: 318).
Huíla Skink Trachylepis huilensis: Butler et al. (2019: 234).
Figures 53, 54, plate 3 In the original description of Euprepes Bayo-

nii, Bocage (1872) noted the existence of two
Mabuya bayoni huilensis Laurent, 1964: 67. varieties: variety A (the “typical one”) from
Holotype: MD 1866 (collected by A. Bar- Duque de Bragança, and variety B from Huíla
ros Machado). Type locality: “Boca da Province. According to the author, the difference
Humpata, environs de Sá da Bandeira” [= between the two was exclusively based on color-
Lubango], Huíla Province, Angola. ation patterns on the dorsum and flanks (Bocage,
Euprepes Bayonii Var. B.: Bocage (1872: 75). 1872). In his major review of the Angolan herpe-
Mabuia Bayonii [part]: Bocage (1895: 38). tofauna, Bocage (1895) no longer referred to
FIGURE 53. Specimen of Trachylepis huilensis from Leba Pass, Huíla Province (CAS 254884). Photos by
L.M.P.C.
variety B, but still noted two coloration forms same taxon. Ceríaco et al. (2016b) confounded
corresponding to the two original varieties. the species with T. varia. Marques et al. (2018)
Laurent (1964) described Mabuya bayoni hui- took a conservative approach and lumped hui-
lensis based on a juvenile specimen that pre- lensis into the nominotypical form until further
sented several differences in terms of scalation research provided evidence regarding its validity,
from the original form, namely the presence of while Branch et al. (2019a) kept it as a subspe-
two frontoparietal scales (vs. a single fused fron- cies, noting, however, that further studies would
toparietal in the nominotypical form). As noted likely elevate it to full species. Based on prelimi-
by Ceríaco et al. (2020a) who reviewed the holo- nary molecular data from a recently collected
type of M. b. huilensis, it is uncertain whether juvenile specimen from Tundavala region, Huíla
Laurent (1964) considered his newly described Province, Butler et al. (2019) confirmed that hui-
subspecies conspecific with Bocage’s Variety B, lensis should be considered a full species, rather
because he did not make any reference to this than a subspecies. Based on newly collected
form, nor did he focus its diagnosis on any col- molecular and morphological data, we found
oration pattern as Bocage did. As the original evidence that the species is more widespread
“Var. B” material from Bocage was lost, it is than originally known.
impossible to recheck the frontoparietal condi- Diagnosis: A small skink (max. SVL 60.4
tion of those specimens. Nevertheless, we feel mm, MUNHAC/MB03-001470), with fully
that it likely that these two entities refer to the developed, pentadactyl limbs (figs. 53, 54);
FIGURE 54. Life photo of Trachylepis huilensis from Tundavala, Huíla Province. Photo by L.M.P.C.
dorsal scales usually pentacarinate; ventral labials cream white to brownish, usually with
scales smooth; 55–66 SAV; 51–56 SAD; 34–39 gray mottling. Upper flanks darker, with black
MSR; lamellae beneath fingers and toes smooth; spots that may overcome the brownish back-
plantar scales smooth; 16–18 LUFT, 12–14 ground; a white to bluish lateral stripe extends
LUFF; supranasals always in contact; parietals from the labials to the hind-limb insertion,
usually in contact; prefrontals always sepa- sometimes indistinct anteriorly; lower flanks
rated; frontoparietals always in contact; one and sides of the neck usually with extensive
pair of enlarged nuchal scales present; ear black speckling. Venter bluish to white; sub-
opening vertically ovoid and smaller than the digital lamellae and plantar scales brown.
eye; four subtriangular auricular scales (shorter Material Examined: Huíla Province: Tun-
than the diameter of ear) extend posteriorly davala [-14.8239°, 13.3811°, 1295 m] (CAS
and usually slightly upward from the anterior 263351); Leba Pass, between river and highway
margin of the ear opening. Eight supralabials, [-15.0703°, 13.2438°, 1670 m] (CAS 254874,
the sixth being subocular; supraciliaries usually 254884); Boca da Humpata [-14.9300°, 13.5200°,
three, the first notably longer; nostrils oriented 1782 m] (MD 1866). Namibe Province: Serra da
dorsally. Dorsum olive brown, with a pair of Neve base camp [-13.7770°, 13.2591°, 1488 m]
faint pale dorsolateral stripes; between these (CAS 263565; UF 187310); Serra da Neve
stripes there are often black-edged scales with [-13.7877°, 13.2572°, 1600 m] (UF 187311);
tendency to form dark longitudinal lines that Serra da Neve [-13.7865°, 13.2572°, 1594 m]
continue through the tail. Top of head uni- (CAS 263558); Serra da Neve [-13.7881°,
formly brownish or with light grayish stippling; 13.2571°, 1612 m] (CAS 263559); Serra da Neve,
FIGURE 55. Distribution map of Trachylepis huilensis in Angola. Black star denotes the type locality.
2016 base camp [-13.7770°, 13.259°, 1488 m] Neve and Serra da Leba (Grandvaux-Barbosa,
(MUNHAC/MB03-001388, 001516); Serra da 1970; fig. 56).
Neve, Catchi surroundings [-13.7619°, 13.2569°,
1585 m] (MUNHAC/MB03-001467, 001470, Trachylepis laevis (Boulenger, 1907)—
001471, 001517). Angolan Blue-Tailed Skink
Distribution in Angola: The species is Figures 57, 58, plate 3
known only from southwestern Angola, in the
highlands of Huíla and Namibe provinces (fig. 55). Mabuia laevis Boulenger, 1907: 212. Holotype:
Global Distribution: Endemic to Angola. BMNH 1946.8.15.31 (formerly BMNH
Habitat and Natural History Notes: 06.8.24.71, collected by W.J. Ansorge). Type
This species inhabits miombo woodlands and locality: “Maconjo, Benguella” [restricted
savannahs in the high-elevation areas of Serra da to “the vicinity of the streams Conjo, Conjo
FIGURE 56. Typical habitat of Trachylepis huilensis, Tundavala, Huíla Province. Photo by L.M.P.C.
Pequeno, and Cocumba (12°52′S, 13°21′E, guela Province, and collected the first topotypic
355 m asl)” by Vaz Pinto et al. (2019)], Ben- material since the original description. Hellmich
guela Province, Angola. (1957b) recorded one specimen from Piri-Dembos,
Mabuya laevis: Hellmich (1957b: 54); Laurent a considerable extension of the known natural
(1964: 76); Branch (1998: 155). range of this species. Laurent (1964) expressed
Oelofsia laevis: Steyn and Mitchell (1965: 2) doubts regarding the identity of this outlier speci-
Trachylepis laevis: Ceríaco et al. (2016b: 33, 57, men, and Marques et al. (2018) noted that this
2020a: 402); Paluh and Bauer (2017: 4); record is “highly improbable and surely represents
Marques et al. (2018: 263, 2019b: 518); a misidentification, although it is not clear to which
Branch et al. (2019a: 318); Vaz Pinto et al. species it might actually belong.” While reviewing
(2019: 37); Lobón-Rovira et al. (2022: 309). Hellmich’s (1957b) specimens in the ZMH, we were
not able to locate any representative of T. laevis,
This conspicuous skink was described by Bou- thus this record remains uncertain. However, con-
lenger (1907) from a specimen collected by W.J. sidering the continuous discovery of new species in
Ansorge in “Maconjo, Benguella.” Vaz Pinto et al. the country, the presence of a not-yet-described,
(2019) recently restricted the type locality to “the morphologically similar species in the forested
vicinity of the streams Conjo, Conjo Pequeno, and highlands of Piri-Dembos cannot be dismissed. We
Cocumba (12°52′S, 13°21′E, 355 m a.s.l.),” Ben- posit that the specimen locality was mislabeled, and
FIGURE 57. Specimen of Trachylepis laevis from N’Dolondolo, Namibe Province (CAS 263541). Photos by
L.M.P.C.
FIGURE 58. Life photo of Trachylepis laevis from N’Dolondolo, Namibe Province (UF 187308). Photo by
L.M.P.C.
FIGURE 59. Distribution of Trachylepis laevis in Angola. Black star denotes the type locality.
that the now lost specimen may have originated tact; one to two pairs of enlarged nuchal scales
from one of the localities in Benguela or Namibe present; ear opening vertically ovoid and smaller
provinces where parts of Hellmich’s (1957b) collec- than the eye; lacking subtriangular auricular scales
tion was obtained. on the anterior margin of the ear opening. Supra
Diagnosis: A medium-sized, flattened-body labials usually nine, the seventh subocular; supra-
skink (max. SVL 63 mm, CAS 263541), with fully ciliaries usually five; nostril oriented laterally.
developed, pentadactyl limbs (fig. 57, 58); dorsal Dorsum and flanks black, with a pair of bright blue
and ventral scales smooth; 56–57 SAV; 49–53 SAD; dorsolateral stripes; a series of irregular blue bars or
30–33 MSR; lamellae beneath fingers and toes spots running along the lower edge of the flanks;
keeled; plantar scales smooth; 19–22 LUFT; 15–16 tail bright blue. Belly grayish blue; arms and legs
LUFF; supranasals separated; parietals in contact; grayish blue below and bright blue above, with
prefrontals always separated; frontoparietals in con- black outlined scales forming a reticulated pattern.
FIGURE 60. Typical habitat of Trachylepis laevis in Mucungo, Namibe Province. Photo by L.M.P.C.
Head and throat orange in life, pale to reddish (MD 1918); Stop 2, Granite outcrops in sandy veld, 50
brown in preserved specimens; a black stripe starts Km East Namibe South West Angola [no exact location]
at the nostril, crosses the eye and continues through (PEM R17939); Assunção [-14.8667°, 13.1000°, 505 m]
(TM 40166–40171; NMZB 16415); Caraculo [-15.0167°,
the flanks; middorsal black band bifurcates behind
12.6667, 463 m] (TM 40231); Virei [-15.7333°, 12.9500°,
the head and extends to the supraoculars. 451 m] (TM 40991); Iona National Park, north of Tam-
Material Examined: Namibe Province: vic. bor [-15.9964°, 12.4071°, 297 m] (CAS 254838); 14 km
N’Dolondolo [-13.8105°, 13.1361°, 713 m] (CAS NE of Caraculo [-14.9021°, 12.7423°, 451 m] (TM 40195,
263541; UF 187308); Omauha-Chitundolo 40196); Tambor [-16.0667°, 12.4500°, 401 m] (TM
[-15.8706°, 12.9030°, 551 m] (CAS 263515); Serra 40499); Rio Coroca – Iona [-16.3000°, 12.4200°, 224 m]
da Neve [-13.7881°, 13.2571°, 1612 m] (CAS (TM 40593); Chapeau Armado Turnoff [-14.5119°,
263582); Omauha [-16.1986°, 12.4007°, 340 m] (UF 12.5019°, 459 m] (TM 41129); Lucira rd 5 KM S of Cat-
ara River [-13.6019°, 12.6332°, 412 m] (TM 41193,
187309); Mucungo farm, rocky area [-14.7781°,
41194); Serra Cafema [-17.1269°, 12.5126°, 2030 m]
12.4888°, 325 m] (CAS 264662); Bentiaba river (P1.131 fide Lobón-Rovira et al., 2022); Vipungos
near Maungo [-14.5106°, 12.8391°, 4174 m] (CAS [-15.0439°, 12.4234°, 333 m] (AMB 13007).
264722); Dirt road to Chingo [-14.3934°, 12.8290°, Historical Localities (no extant speci-
618 m] (MUNHAC/MB03-001389); Serra da Neve, mens): Kwanza Norte Province: Piri-Dembos
boulders near base camp [-13.7646º, 13.2601º, 1603 (questionable record) [-8.5299°, 14.4377°, 712 m]
m] (MUHNAC/MB03-001463–001465). (Hellmich, 1957b).
Additional Material (* denotes type material): Distribution in Angola: This species
Benguela Province: 53 km SE of Benguela [-12.9324°,
occurs in the low-elevation areas of Namibe and
13.5526°, 656 m] (TM 41273); Maconjo [-12.8667°,
13.3500°, 355 m] (BMNH 1946.8.15.31*); 35 km S of Benguela provinces (fig. 59).
Dombe Grande [-13.2098°, 12.9802°, 326 m] (TM 41252, Global Distribution: Endemic to south-
41253; NMZB 16416). Namibe Province: Muninho 50 western Angola and northwestern Namibia
km west Sá da Bandeira [-14.9667°, 12.9667°, 393 m] (Steyn and Mitchell, 1965; Bauer et al., 1993).
Habitat and Natural History Notes: ranging from West to East Africa (Mausfeld-
Trachylepis laevis is usually found on granite Lafdhiya et al., 2004; Ceríaco et al., 2016a; Allen
outcrops in a mosaic of steppe and arid savan- et al., 2019). It was originally described by Gray
nah (Grandvaux-Barbosa, 1970; fig. 60). This (1845) from a specimen from “West Africa.” Its
rupiculous skink is dorsoventrally depressed, taxonomic and nomenclatural history is compli-
with several morphological adaptations to its cated, with more than a dozen different nomina
crevice-dwelling habits (Paluh and Bauer, 2017). proposed as subspecies and junior synonyms
(Broadley, 1974; Hoogmoed, 1974; Ceríaco et al.,
Trachylepis maculilabris (Gray, 1845)— 2016a). It has long been known that the Trachy-
Speckled-Lipped Skink lepis maculilabris species complex comprises at
least two clearly distinct species—the nomino-
Figures 61–63, plate 4 typical form from West Africa, and a putatively
undescribed species in East Africa (Mausfeld-
Euprepis maculilabris Gray, 1845: 114. Holo-
Lafdhiya et al., 2004; Allen et al., 2019). Despite
type: BMNH 1946.8.18.17 (formerly
this fact, the nomenclatural complexity of the
BMNH xv.109a, collected by W. Raddon).
group has prevented taxonomists from formally
Type locality: “W. Africa.”
describing the East African form due to the need
Euprepes Anchietae: Bocage 1866a: 44 Neotype
for a careful revision of the extant type material
[here designated]: An unsexed adult
pertaining to the numerous nomina associated
(MUNHAC/MB03-001390, field number with the complex. Allen et al. (2019) showed that
AMB 11108; fig. 63) collected by S. Ban- within the West-African (nominotypical) form,
deira in June 2018. Type locality: Sanctu- there are considerable levels of phylogenetic
ary of the Catholic Mission of Cabinda, structure. A review of the several names cur-
Cabinda Province, northwestern Angola rently in the synonymy of T. maculilabris was
[-5.5614°, 12.1779°, 7 m]. presented by Ceríaco et al. (2016a). While some
Euprepes Anchietae: Bocage (1866b: 62). of these synonyms may prove in the future to
Euprepes (Eupr.) Perrotetii: Peters (1877: 614); represent valid taxa, their status does not affect
Bocage (1895: 39). the Angolan population.
Mabuia maculilabris: Boulenger (1887: 164); The Angolan populations are conspecific with
Bocage (1895: 40); Ferreira (1906: 170); the nominotypical form, as shown by our molecular
Monard (1937: 85). and morphological data. Nevertheless, the nomen-
Mabuya maculilabris: Loveridge (1933: 312); clatural history of the species within Angolan bor-
Parker (1936: 138); Laurent (1950: 12, 1954: ders also reflects its nomenclatural complexity.
65); Broadley (2000: 94). Bocage (1866a) coined the name Euprepes anchietae
Mabuya maculilabris maculilabris: Hellmich as a nomen nudum for a specimen brought from
(1957a: 61); Loveridge (1957: 209); Laurent “Zaire,” Angola, by José d’Anchieta. A valid descrip-
(1964: 65). tion appeared in a subsequent paper (Bocage,
Euprepes maculilabris: Mausfeld-Lafdhiya et al. 1866b) of the same issue of the journal. The holo-
(2004: 160). type of this species was destroyed in the fire that
Trachylepis maculilabris: Ceríaco et al. (2014: 671, consumed Museu Bocage in 1978. Years later, Peters
2020a: 402); Branch and Conradie (2015: (1879) described Euprepes notabilis based on speci-
200); Marques et al. (2018: 263); Branch et al. mens collected in “Chinxoxo,” Cabinda Province
and another from “Pungo Andongo,” Malanje Prov-
(2019: 318); Santos et al. (2021: 24).
ince. Subsequent authors who have dealt with
Trachylepis maculilabris is one of the most Angolan populations of T. maculilabris considered
widespread species of the genus in the continent, both anchietae and notabilis as junior synonyms of
FIGURE 61. Specimen of Trachylepis maculilabris from Serra Mucaba, Uíge Province (MUNHAC/MB03-
FIGURE 62. Life photo of Trachylepis maculilabris from Serra Mucaba, Uíge Province (MUNHAC/MB03-
FIGURE 63. Neotype of Euprepes anchietae from Cabinda, Cabinda Province (MUNHAC/MB03-001390).
Photos by L.M.P.C.
the nominotypical form (Boulenger, 1887; Bocage, Together with T. albilabris, T. maculilabris is a
1895; Schmidt, 1919; Bauer et al., 2003; Marques et representative of the central/western African clade
al., 2018). While destruction of the complete type of the genus (Weinell et al., 2019). It has histori-
series of Euprepes anchietae impedes its correct tax- cally been recorded in the northwestern areas of
onomic allocation, the existence of type material of the country by several authors (Bocage, 1895; Fer-
Euprepes notabilis confidently allows us to consider reira, 1903; Parker, 1936; Hellmich, 1957b) and
notabilis as a valid species (see account below). has recently been collected in Bengo (this paper)
Given that both notabilis and maculilabris are sym- and Uíge provinces (Ernst et al., 2020; this paper),
patric in the Cabinda region from where anchietae where its habitat presents a southern continuation
was described, and given the overall similarity of the Congolese habitats and biomes.
between both species, the type material of anchietae Diagnosis: A medium-sized and robust skink
could correspond to either one of the species. (max. SVL of Angolan specimens 91.9 mm, MUN-
Herein, we opt to follow the opinion of most authors HAC/MB03-001403; max. SVL of extralimital
who have dealt with this nomen and consider anchi- populations 95 mm; see Allen et al., 2017), with
etae as a junior synonym of maculilabris. To stabilize fully developed, pentadactyl limbs (fig. 61–63);
the taxonomy of the group, we designate a neotype dorsal scales usually pentacarinate; ventral scales
for anchietae, which corresponds both genetically smooth; 50–66 SAV; 47–62 SAD; 32–40 MSR;
and morphologically to T. maculilabris. lamellae beneath fingers and toes smooth; plantar
FIGURE 64. Distribution of Trachylepis maculilabris in Angola. White star denotes the locality of the neotype
of Euprepes anchietae.
scales smooth; 12–19 LUFT, 10–15 LUFF; suprana- usually with scattered black speckles; upper flanks
sals usually in contact or touching at a single point; often darker; lower flanks lighter, grayish to orange;
parietals usually in contact; prefrontals usually in scattered white speckles on the flanks, sometimes
contact; frontoparietals always in contact; one pair extending to the limbs. Supralabials with series of
of enlarged nuchal scales present; ear opening verti- white spots and dashes, usually forming a white
cally ovoid and smaller than the eye, lacking subtri- stripe extending to the ear opening or the forelimb
angular auricular scales on the anterior margin. insertion; auricular and orbicular regions often yel-
Supralabials usually seven, the fifth is subocular lowish to orange. Venter, throat and infralabials
(sometimes third or fourth); usually five supracili- whitish to yellow in life, white in preserved speci-
aries, but may vary from four to seven; nostril ori- mens, uniformly colored; occasionally scattered
ented laterally. Dorsum olive gray to dark brown, black punctuations on the underside of tail.
FIGURE 65. Typical habitat of Trachylepis maculilabris in Serra Mucaba, Uíge Province. Photo by L.M.P.C.
Neotype [for Euprepes anchietae Bocage, Pingano [-7.6829, 14.9340, 744 m] (MUNHAC/
1866]: Here designated, an unsexed adult (MUN- MB03-001403).
HAC/MB03-001390, field number AMB 11108; Additional Material: Benguela Province: Entre
fig. 63) collected at the Sanctuary of the Catholic Rios, Dep. De Benguela [-13.0167°, 14.6333°, 1267 m]
Mission of Cabinda, Cabinda Province, north- (ZSM 99/1953). Cabinda Province: near Tafe Beach
[-5.5578°, 12.1796°, 5 m] (CAS 264215, CAS 264216);
western Angola [-5.5614°, 12.1779°, 7 m], by S.
Fortaleza, landfill [-5.6528°, 12.2481°, 114 m] (CAS
Bandeira in June 2018. 264217). Kwanza Sul Province: Congulu, Amboim
Material Examined (* denotes type mate- [-10.8667°, 14.2833°, 640 m] (BMNH 1936.8.1.612–
rial): Cabinda Province: Cabinda, at the Catho- 617); Quirimbo, 75 km E of Amboim [-10.6833°,
lic Mission Sanctuary [-5.5614°, 12.1779°, 7 m] 14.2667°, 302 m] (BMNH 1936.8.1.618–619); Cada
(MUNHAC/MB03-001390*); Municipal beach Amboim, abandoned plantation [-10.8672°, 14.3244°,
(Tafi’s Beach) [-5.5578°, 12.1796°, 5 m] (CAS 930 m] (CAS 263098). Lunda Norte Province: Dundo
264215, 264216); Fortaleza [-5.6528°, 12.2481°, [-7.3667°, 20.8333°, 663 m] (BMNH 1966.250–255;
MD 226, 2259, 5085, 5088, 5090, 5092, 5094, 5099,
114 m] (CAS 264217). Huíla Province: Gungue,
5103, 5119, 5130, 5142, 5150, 5155, 5164, 5200, 5209);
Rio Kwando [-13.7363º, 15.4216º, 1514 m] (AMB Muita [-7.8000°, 21.4100°, 725 m] (MD 807, 1076); De
12713, 12714). Malanje Province: Laúca Dam, Beers, Lucapa [-8.4228°, 20.7392°, 949 m] (PEM
flooded area [-9.7627°, 15.1438°, 750 m] (MUN- R19459, 19463); Headwaters of Lovua, North of village
HAC/MB03-001352–001361); swamp on the Capaia [-8.3385°, 20.2425°, 953 m] (PEM R19459),
edge of corn field, Kalandula [-9.0755°, 16.0117°, Mucoso bei Dondo [-9.5333°, 14.6500°, 280 m] (ZSM
1063 m] (CAS 263587). Uíge Province: Forested 100/1953); Carumbo, Lucapa [-8.4228°, 20.7392°, 949
m] (Branch and Conradie, 2015). Lunda Sul Province:
lagoon in Mucaba mountains [-7.2430°, 15.1673°,
Alto Cuíla (= Alto Cuílo) [-10.0500°, 19.5170°, 1260 m]
1185 m] (MUNHAC/MB03-001401, 001402); (MCZ R-74111; MD 5308, 5309). Malanje Province:
Camp site near water pump, east of Ponte, Serra Ndalla Tando (currently N’dalatando) [-9.3000°,
14.9167°, 782 m] (BMNH 1909.10.29.99); Duque de 8485 (collected by J.G. Falkenstein, lost fide
Bragança (currently Kalandula) [-9.1333°, 16.0667°, Bauer et al., 2003). Type locality: “Chin-
1010 m] (TM 45468); Capanda [-9.7284°, 15.3459°, 859 choxo,” Cabinda Province (ZMB 8485) and
m] (MNHNL Rep/A/Sc 1–36); Quifangondo [-8.7667°,
13.4333°, 9 m] (MNHNL Rep/A/Sc no number); Kalan-
“Pungo Andongo,” Malanje Province (ZMB
dula Falls [-9.0740°, 16.0030°, 1069 m] (CAS 263586). 9204), Angola.
Zaire Province: Ambriz [-7.8574°, 13.1182°, 36 m] Mabuya maculilabris: Bauer et al. (2003: 275).
(BMNH 1851.1.19.8–11). Undetermined locality:
This long-forgotten species was described by
Unknown locality (AMNH 48710–48726; FMNH
74301; MHNG 1545.015; MNHN 1907.200, 1907.256; Peters (1879) as Euprepes notabilis based on
MHNC-UP/REP 272). specimens from “Chinchoxo,” Cabinda Prov-
Historical Localities (no extant speci- ince and another from “Pungo Andongo,”
mens): Cabinda Province: Cabinda [-5.5500°, Malanje Province (the latter still extant in the
12.1833°, 6 m] (Bocage, 1895). Kwanza Norte collections of ZMB, accession number ZMB
Province: Dondo (Quanza edges) [-9.6833°, 9204, fig. 66). All subsequent authors consid-
14.4333°, 33 m] (Bocage, 1895); Golungo ered E. notabilis as a junior synonym of T. mac-
[-9.1333°, 14.7667°, 666 m] (Ferreira, 1906); Cam- ulilabris (Boulenger, 1887; Bocage, 1895;
bondo [-9.1596, 14.6577, 374 m] (Ferreira, 1906). Schmidt, 1919; Bauer et al., 2003; Marques et
Distribution in Angola: The distribution of al., 2018), and the taxon was apparently forgot-
this species in Angola is restricted to the northern ten. Only recently, during field surveys in Lauca
areas of the country, from the Cabinda enclave region in the vicinity of Pungo Andongo (ca. 50
eastward to Lunda Norte and Lunda Sul prov- km east), new specimens were collected in sym-
inces, and southward along the escarpment to Bié, patry with nominotypical T. maculilabris (figs.
Kwanza Sul, and Benguela provinces (fig. 64). 67, 68). Although T. notabilis differs morpho-
Global Distribution: The species has a logically from T. maculilabris, the two species
wide distribution from West Africa, through are very similar (fig. 68). The main characters
Central Africa, reaching its southern distribution that appear to differ in these species are the
in western Angola. East African populations general appearance of the male’s body, with T.
have long been known to represent a different notabilis being much bulkier than T. maculila-
taxon (Mausfeld-Lafdhiya et al., 2004; Allen et bris, and ventral coloration, with T. notabilis
al., 2019). having a strikingly blue neck and yellow venter
Habitat and Natural History Notes: versus a homogeneously whitish or yellow ven-
This species occupies a variety of habitats in a ter in T. maculilabris (fig. 68). Both our mor-
forest-savannah mosaic, as well as anthropogenic phological and molecular data support its
habitats such as human settlements (Grandvaux- recognition as a valid species. Since the speci-
Barbosa, 1970; fig. 65). It is diurnal and arboreal, men from “Chinchoxo” was lost (see Bauer et
frequently found in open habitats such as forest al., 2003), we designate the remaining syntype
clearings, often climbing trees and bushes, or from Pungo Andongo (ZMB 9204) as the lecto-
even buildings and walls. type of the species in order to stabilize its
nomenclature and avoid potential confusions
with other species occurring in the region.
Trachylepis notabilis (Peters, 1879)— Diagnosis: A large, robust skink (max. SVL
Notable Skink 97.9 mm, MUNHAC/MB03-001355) with fully
Figures 66–69, plate 4 developed, pentadactyl limbs (figs. 66–69);
dorsal scales usually quadricarinate or pentac-
Euprepes notabilis Peters, 1879: 36. Syntypes: arinate; ventral scales smooth; 50–60 SAV;
ZMB 9204 (collected by A.V. Homeyer) and 49–59 SAD; 31–38 MSR; lamellae beneath fin-
FIGURE 66. Lectotype of Trachylepis notabilis from Pungo Andongo, Malanje Province (ZMB 9204). Photos
by L.M.P.C.
FIGURE 67. Recently collected specimen of Trachylepis notabilis from Lauca, Malanje Province (MUNHAC/
MB03-001355). Photos by L.M.P.C.
FIGURE 68. Comparison between two adult specimens of Trachylepis maculilabris (left; MUHNAC/MB03-
001359 and 001360) and one juvenile and one adult of Trachylepis notabilis (right; MUHNAC/MB03-001351
and 001355). All specimens from Lauca, collected in the same day. Photo by L.M.P.C.
FIGURE 69. Life photo of Trachylepis notabilis from Cassumbi, Bié Province (MUHNAC/MB03-001404).
Photo by L.M.P.C.
FIGURE 70. Distribution of Trachylepis notabilis in Angola. Black stars denote the type locality.
gers and toes smooth; plantar scales smooth; grayish to orange; scattered white speckles on
13–18 LUFT; 11–15 LUFF; supranasals usually the flanks, sometimes extending to the limbs.
in contact; parietals usually separated; pre- Top of head uniform brown; labials, auricular
frontals always in contact; frontoparietals and orbicular regions often yellowish to
always in contact; one pair of enlarged nuchal orange. Supralabials often with series of white
scales present; ear opening vertically ovoid spots and dashes, usually forming a white
and smaller than the eye, lacking subtriangular stripe extending to the ear opening or the fore-
auricular scales on the anterior margin. Supra- limb insertion. Venter and underside of tail
labials usually seven, the fourth or fifth suboc- uniform yellowish in life, white in preserved
ular; supraciliaries 5–6; nostril oriented specimens; gular region white anteriorly and
laterally. Dorsum olive gray to dark or golden bluish posteriorly in males.
brown, usually with scattered black speckles; Material Examined (* denotes type mate-
upper flanks often darker; lower flanks lighter, rial): Bié Province: Cassumbi [-11.08088°,
FIGURE 71. Typical habitat of Trachylepis notabilis in Lauca, Malanje Province. Photo by L.M.P.C.
16.66593°, 1223 m] (MUHNAC/MB03- Trachylepis occidentalis (Peters, 1867)—

001404). Malanje Province: Pungo Andongo Western Three-Striped Skink
[-9.6695°, 15.5893°, 1149 m] (ZMB 9204*);
Laúca Dam [-9.76275°, 15.14380°, 750 m]
(MUNHAC/MB03-001349, 001350, 001351, Euprepes vittatus var. australis Peters, 1862: 19
001355). [preoccupied by Euprepes australis (Gray,
Historical Localities (no extant speci- 1839)]. Syntypes: ZMB 4212, 64401-02
mens): Cabinda Province: Chinchoxo [-5.1000°, (formerly ZMB 4212A-B) (collected by
12.1000°, 45 m] (Peters, 1879). C.H. Hahn). Type locality: “Neu-Bar-
Distribution in Angola: The species is only men” [= Gross Barmen], Otjozondjupa
known from northern and central Angola, from Region, Namibia fide Bauer et al. (2003).
Cabinda to Bié Province (fig. 70). Euprepes occidentalis Peters, 1867: 20 [replace-
Global Distribution: Currently known ment name for Euprepes vittatus var. autralis
only from material from Angola, but must also Peters, 1862].
be present in the neighboring areas of the Repub- Euprepes occidentalis: Bocage (1870: 68).
lic of Congo and the Democratic Republic of the Mabuia occidentalis: Bocage (1895: 42).
Congo, given its occurrence in Cabinda. Mabuya occidentalis: Laurent (1964: 73); Branch
Habitat and Natural History Notes: The (1998: 156); Broadley (2000: 97).
species seems to be associated with riparian areas Trachylepis occidentalis: Masterson et al. (2014a:
with rocky outcrops (fig. 71). 263); Ceríaco et al. (2016b: 33, 57; 2020a:
FIGURE 72. Specimen of Trachylepis occidentalis from Pico Azevedo, Namibe Province (CAS 254931). Photos
by L.M.P.C.
FIGURE 73. Life photo of Trachylepis occidentalis from Espinheira, Namibe Province. Photo by William R.
Branch.
FIGURE 74. Distribution of Trachylepis occidentalis in Angola.
402); Marques et al. (2018: 266); Branch et name Euprepes occidentalis (Bauer et al., 2003).
al. (2019a: 318); Santos et al. (2021: 24); The first record of this species from Angola con-
Lobón-Rovira et al. (2022: 309). sisted of two specimens collected in 1867 by José
Trachylepis acutilabris [part]: Ceríaco et al. d’Anchieta in “Rio Coroca,” Namibe Province
(2016b: 34). (Bocage, 1895). Additional specimens were col-
lected by Francisco Newton in 1905 from Baía
Peters (1862) originally described this large dos Tigres, Namibe Province (Santos et al.,
skink as Euprepes vittatus var. australis. However, 2021), but subsequent published records were
this name was preoccupied by Euprepes australis only provided by Laurent (1964), and more
(Gray, 1839) [= Ctenotus australis (Gray, 1839)], recently by Ceríaco et al. (2016b) and Lobón-
and Peters (1867) later provided the replacement Rovira et al. (2022).
FIGURE 75. Typical habitat of Trachylepis occidentalis in Espinheira, Namibe Province. Photo by L.M.P.C.
Diagnosis: A larger-sized skink (max. SVL reddish brown, with five pale, dark-edged, lon-
110.7 mm, MHNC-UP/REP 274), with fully gitudinal stripes. Dorsal stripes start behind
developed, pentadactyl limbs (fig. 72, 73); dor- the head and may extend beyond the base of
sal scales tricarinate; ventral scales smooth; the tail; dorsolateral stripes narrower than ver-
56–58 SAV; 45–50 SAD; 30–34 MSR; lamellae tebral and lateral ones. A pale stripe covers the
beneath fingers and toes keeled and feebly spi- supralabials and extends through the flanks,
nose; plantar scales smoothly keeled; 18–24 followed below by a brownish stripe that may
LUFT; 13–16 LUFF; supranasals always in con- be faint and indistinct in preserved specimens.
tact; parietals usually separated or touching at Underparts uniform white.
a single point; prefrontals usually separated; Material Examined: Namibe Province:
frontoparietals always in contact; one pair of Curoca river [-16.2653°, 12.3210°, 188 m] (UF
enlarged nuchal scales present; ear opening 187307); Pico Azevedo [-15.5340°, 12.4920°, 372
vertically ovoid and smaller than the eye; two m] (CAS 254931); Baía dos Tigres [-16.5856°,
subtriangular auricular scales (shorter than 11.8271°, 98 m] (MHNC-UP/REP 273, 274).
the diameter of ear) extend posteriorly and Undetermined locality: Unknown locality
usually slightly upward from the anterior mar- (MHNC-UP/REP 275).
gin of the ear opening. Supralabials usually Additional Material: Namibe Province: Espin-
seven, the fifth subocular; five supraciliaries, heira [-16.7500°, 12.3667°, 437 m] (PEM R17979,
the second slightly larger than others; nostrils 17980, 20356); Moçâmedes [-15.1667°, 12.1667°, 6 m]
oriented dorsally. Dorsum and flanks olive to (TM 24456; MD 1945); Rio Coroca Mouth [-15.7300°,
11.9200°, 6 m] (TM 40401, 40402); Kondundo encompasses eastern Zimbabwe, the southern
[-16.8788°, 12.3720°, 515 m] (JLRZC0117 fide Lobón- parts of lake Malawi (in Malawi and Mozam-
Rovira et al., 2022). bique), and the western and isolated southwest-
Historical Localities (no extant speci- ern Angolan population (Broadley, 2000;
mens): Namibe Province: Rio Coroca (Bocage, Pietersen et al., 2021). When referring to the
1895). isolated Angolan population, Broadley (1974b)
Distribution in Angola: The range of this suggested that it might represent a distinct taxon.
species in Angola is restricted to the arid regions Our molecular results show that these popula-
of Namibe Province (fig. 74). tions belong to an undescribed lineage, sister to
Global Distribution: Occurs from south- a clade comprising T. hilariae, T. variegata, and
western Angola southward through Namibia to T. vunongue (fig. 1). The specimens cited by
South Africa and southwestern Botswana Broadley are still in the collections of the TM,
(Auerbach, 1986; Branch, 1998; Masterson, although in a poor state of preservation.
2014a). Holotype: An unsexed adult (CAS 263493,
Habitat and Natural History Notes: field number AMB 10630; fig. 76) collected at
This species is found in the arid habitats of Virulundo, Namibe Province [-16.2852°,
Namibe Province, from steppes to mobile sand 12.9419°, 718 m], by Luis M.P. Ceríaco, Suzana
dunes (Grandvaux-Barbosa, 1970; fig. 75). It is A. Bandeira, and Ishan Agarwal, on 2 Decem-
terrestrial, taking refuge in holes dug in the ber 2016.
sand near the base of spiny shrubs (Branch, Additional Material: Namibe Province: Cainde,
1998; Broadley, 2000; Ceríaco et al., 2016b). Mossamedes [-15.4833°, 13.3667°, 631 m] (TM 40959,
This skink is known to be oviparous in the 40960); 14 km NE of Caraculo, Mossamedes [-14.9021°,
Kalahari region, but viviparous in the Namib 12.7423°, 451 m] (TM 40192).
Desert, Namibia, and Free State Province, Historical Localities (no extant speci-
South Africa (Branch, 1998; Broadley, 2000). mens): Cunene Province: Otchinjau [-16.5025°,
13.9240°, 363 m] (TM 40805—lost). Namibe
Province: Cainde, Mossamedes [-15.4833°,
Trachylepis ovahelelo, sp. nov.— 13.3667°, 631 m] (TM 40958—lost).
Ovahelelo Skink Diagnosis: A small-sized skink (max. SVL
37.3 mm, CAS 263493), with fully developed,
Figure 76, plate 5
pentadactyl limbs (fig. 76); dorsal scales pen-
Mabuya lacertiformis: Broadley (1974b: 12, 2000: tacarinate; ventral scales smooth; 53 SAV; 43
SAD; 34 MSR; lamellae beneath fingers and
101); Branch (1998: 154).
toes spinose; plantar scales spinose; 18 LUFT,
Trachylepis lacertiformis: Portik and Bauer (2012:
14 LUFF; supranasals in contact; parietals in
128); Ceríaco et al. (2016a: 57).
contact; prefrontals separated; frontoparietals
Trachylepis cf. lacertiformis: Marques et al. (2018:
in contact; one pair of enlarged nuchal scales
262); Branch et al. (2019a: 318).
present; ear opening vertically ovoid and
Broadley (1974b) was the first to report the smaller than the eye; two subtriangular auricu-
existence of an isolated population of Mabuya lar scales (shorter than the diameter of ear)
lacertiformis in southern Angola, based on speci- extend posteriorly and usually slightly upward
mens collected by Wulf Haacke in Namibe and from the anterior margin of the ear opening.
Cunene provinces (specimens currently housed Seven supralabials, the fifth subocular; five
in TM). Trachylepis lacertiformis (Peters, 1854) supraciliaries, the second notably longer than
was originally described from the Tete region in others; nostrils oriented dorsally. Dorsum
Mozambique, and its current known distribution grayish to golden brown, uniform or with
TABLE 7 series of transversely aligned dark speckles;

Mensural and Meristic Data for the Holotype of
there may be a pair of indistinct pale dorso
Trachylepis ovahelelo, sp. nov. lateral stripes, especially on the tail; flanks
Abbreviations are listed in the Materials and Methods. uniformly colored. An incomplete dark stripe
Measurements are presented in millimeters starts at the nostril and extends to the ear
and ratios as percentages opening; labials white. Venter white, with light
grayish speckling that may form longitudinal
CAS 263493
lines on the chin, throat, and tail.
Holotype Description of the Holotype: A well-
preserved, unsexed adult. Body cylindrical and
Sex unsexed
robust with a poorly defined neck and well-
SVL 37.3
developed pentadactyl limbs; tail long, its
TL 70.9
length almost twice the SVL, smoothly taper-
TL/SVL 190 ing. Fore- and hind limbs overlap when
HL 8 adpressed against the body. SVL 37.3 mm, TL
HL/SVL 20 70.9 mm. HL 3.3 mm, with relatively long and
SVL/HL 470 thin snout. Additional measurements are pre-
HW 5.8 sented in table 7. Ear opening medium sized.
HW/HL 70
Two subtriangular auricular scales extend pos-
teriorly from the anterior margin of the ear
HH 3.3
opening. Rostral visible from above. Nostrils
IN 0.9
oriented dorsally and set posteriorly, so that
EN 2.5 postnasal effectively borders nostril. Suprana-
ES 3.2 sals in contact. Frontonasal broader than long,
MSR 34 in contact with loreal scale. Prefrontals subtri-
SAD 43 angular, without contact, each in contact with
SAV 53 the following head shields: frontonasal, loreals,
LUFF 14 first and second supraocular, and frontal. Two
LUFT 18
loreals. Frontal length similar to the distance
between anterior tip of frontal and tip of snout.
SC 3
Frontal in contact with two supraoculars on
SL (SO) 7 (5)
each side. Two frontoparietals, in contact with
CP C each other and the frontal, third and fourth
CFP C supraoculars, parietal and interparietal. Fron-
CSN C toparietal plus interparietal length similar to
CPF S frontal length. Interparietal twice as long as
KDS 5 broad, with a visible parietal foramen. Pari-
Plantar scales spinose etals greater than frontoparietals. Parietals in
contact in a single point with each other. Five
supraciliaries, second largest. Seven supralabi-
als, fifth subocular. Seven infralabials. Post-
mental bordering seven scales (mental, two
infralabials on each side and two primary chin
shields). Transparent scale present in lower
eyelid, as is usual for Trachylepis. Tympanum
visible, at same level as mouth. Dorsal scales
FIGURE 76. Holotype of Trachylepis ovahelelo, sp. nov., from Virulundo, Namibe Province (CAS 263493).
Photos by L.M.P.C.
each with five smooth keels. Ventral scales T. bayonii, T. binotata, T. hoeschi, T. laevis, T.
smooth. MSR 34, SAD 43, SAV 53. Limbs with maculilabris, T. notabilis, T. occidentalis, and
five digits; plantar scales spinose. Relative T. raymondlaurenti, by having spinose plantar
length of fingers IV > III > V > II > I, relative scales. It differs from T. attenboroughi, and
length of toes IV > III > V > II > I. Finger-IV T. wahlbergii by the presence of subtriangu-
lamellae 14, Toe-IV lamellae 18. lar auricular scales on the anterior margin of
Coloration in Ethanol: Dorsum brown, the ear opening. T. ovahelelo can be readily
with 11 series of transversely aligned dark speck- distinguished from T. suzanae and T. wilsoni
les between the forelimbs and the base of the tail; by absence of wedge-shaped snout. It can be
a pair of barely distinct, pale dorsolateral stripes; readily distinguished from T. albopunctata, T.
flanks uniform brown. There is an irregular and damarana, T. ansorgii, and T. sulcata by hav-
faint dark stripe from the nostril to the ear opening its nostrils situated more dorsally, directed
ing; labials white. Ventral side white, with light upward (vs. nostrils situated more laterally,
grayish speckling laterally that forms faint longi- directed sideward in the latter). T. ovahelelo
tudinal lines on the tail. can be distinguished from T. hilariae by hav-
Comparison with Other Angolan and ing 34 MSR (vs. 29–30 in the latter), 53 SAV
Southwest African Trachylepis: Trachylepis (vs. 58), 43 SAD (vs. 46–48), 14 LUFF (vs.
ovahelelo, sp. nov., differs from T. albilabris, 15–16) and 18 LUFT (vs. 21–22). It differs
FIGURE 77. Distribution of Trachylepis ovahelelo in Angola. Black star denotes the type locality.
from T. punctulata by having 18 LUFT (vs. of the variegata subgroup sensu Weinell et al.
21–24 in the latter). It is distinguished from (2019), T. ovahelelo is readily differentiated
T. vunongue by the presence of transversely from T. variegata by having five keels on dor-
aligned dark speckles (vs. dorsum uniform or sal scales (vs. three in T. variegata). Similar
with irregularly scattered dark and pale spots to Broadley (1974b), we were unable to find
in the latter) and a longer tail (TL/SVL 190% any major morphological difference between
vs. 111%–152% in T. vunongue). It can be dis- T. ovahelelo and T. lacertiformis. Misidentifi-
tinguished from T. bouri and T. chimbana by cations are, however, very unlikely, as the two
having only 43 SAD (vs. >48 in the latter). T. species are separated by a broad (more than
ovahelelo can be readily distinguished from T. 1500 km) band of aelion soils, with T. ova-
bocagii by the absence of dorsal bands (vs. five helelo in southwestern Angola and the clos-
yellowish dorsal bands in T. bocagii). Regard- est population of T. lacertiformis in western
ing the other southwestern African congeners Zimbabwe.
FIGURE 78. Type locality of Trachylepis ovahelelo in Virulundo, Namibe Province. Photo by L.M.P.C.
Distribution in Angola: This species is Trachylepis punctulata (Bocage, 1872)—

known only from southwestern Angola, in Speckled Sand Skink
Namibe and Cunene provinces (fig. 77).
Global Distribution: Currently known only Figures 79, 80, plate 5
from material from Angola, but almost certainly Euprepes punctulatus Bocage, 1872: 76. Syn-
present in the neighboring areas of Namibia, given types: MBL 742 (2 specimens) (collected by
its close proximity to the border. J.A. Anchieta, destroyed by fire on March
Habitat and Natural History Notes: This
18, 1978); ZMB 6478 (collected by J.A.
is a rupiculous species that is usually found on
Anchieta; see Bauer et al., 2003: 276). Type
weathered rocky outcrops with fissures (usually
locality: “Rio Coroca, dans le littoral au
granite) or loose rocks in mopane woodlands and
sud de Mossamedes,” Namibe Province,
steppe-savannah mosaic (Grandvaux-Barbosa,
Angola.
1970; Broadley, 1974b; fig. 78).
Mabuia punctulata: Boulenger (1887: 204);
Etymology: The specific epithet “ovahelelo” is
a noun in apposition and is given in honor of the Bocage (1895: 44, 1897a: 195).
dominant ethnolinguistic group of the region Mabuya punctulata [part]: Schmidt (1933: 12);
where the species exists. We acknowledge the Frade (1963: 253); Broadley (2000: 100).
Mucubal people, a subgroup of the Ovahelelo (also Mabuya longiloba longiloba: Laurent (1964: 73).
known as Herero) ethnolinguistic group, for allow- Mabuya variegata punctulata [part]: Broadley
ing our team to camp and study the herpetofauna (1974b: 7); Branch (1998: 158).
of their lands and for their welcoming attitude and Trachylepis punctulata [part]: Portik (2009:
support. We suggest “Ovahelelo Skink” and “Lagar- 136); Portik and Bauer (2012: 128);
tixa Ovahelelo” as the English and Portuguese com- Ceríaco et al. (2016b: 31, 57); Marques et
mon names for this species, respectively. al. (2018: 267); Branch et al. (2019: 319);
Santos et al. (2021: 25); Lobón-Rovira et al. conspecific with the southwestern Angolan form,
(2022: 310). and we interpret them instead as representing
Trachylepis cf. triebneri, while the southeastern
Described by Bocage based on two specimens Angolan and western Zambian populations rep-
from “Rio Curoca,” Namibe Province, this taxon resent a new species, Trachylepis vunongue (see
has experienced a convoluted taxonomic and account below). Our present sample size does
nomenclatural history—see Broadley (1974b). not allow us to confidently assess whether true
The Angolan populations were considered either punctulata is endemic to Angola or it also
as a valid species (Bocage, 1872, 1895, 1897a; extends to northern Namibia (but has yet to be
Boulenger, 1887; Schmidt, 1933; Frade, 1963), as genetically sampled). The role of the Cunene
Mabuya (= Trachylepis) longiloba (Laurent, River as a potential barrier for this group needs
1964), or as a subspecies of Mabuya (= Trachyl- to be further investigated. One outcome of our
epis) variegata (Broadley, 1974b, Branch, 1998). interpretation is that the previously accepted
The work of Broadley (1974b) was of particular range of T. punctulata (see Branch, 1998) is now
importance to resolve the taxonomic chaos into much smaller and wholly or mostly restricted to
which T. punctulata, as a member of the “Mabuya southwestern Angola.
lacertiformis complex” was involved. Broadley Diagnosis: A small-sized skink (max. SVL
considered that T. punctulata has a broad distri- 48.5 mm, MUNHAC/MB03-1409), with fully
bution, from southern Angola through a large developed, pentadactyl limbs (fig. 79, 80);
extent of northern and central Namibia, dorsal scales usually pentacarinate; ventral
Botswana, Zimbabwe, Zambia, northern parts of scales smooth; 46–60 SAV; 42–50 SAD; 28–35
South Africa, and Mozambique. This interpreta- MSR; lamellae beneath fingers and toes keeled
tion was followed by Branch (1998). However, and spinose; plantar scales spinose; 21–24
Broadley (1974b) noted that the populations LUFT; 12–18 LUFF; supranasals in contact;
from Botswana, Zimbabwe, Zambia, South parietals usually in contact; prefrontals always
Africa, and Mozambique have considerable mor- separated; frontoparietals in contact; one pair
phological differences from those of Angola and of enlarged nuchal scales present; ear opening
Namibia, which may reflect either a clinal varia- vertically ovoid and smaller than the eye; 2–3
tion or even a different taxon (see the account of subtriangular auricular scales (shorter than
Trachylepys vunongue). Broadley (2000), Portik the diameter of ear) extend posteriorly and
(2009), and Portik and Bauer (2012) had pro- usually slightly upward from the anterior mar-
vided, respectively, morphological and molecular gin of the ear opening. Supralabials usually
evidence in support of T. punctulata as a full spe- eight, the sixth subocular; usually five supra-
cies, rather than a subspecies of T. variegata. ciliaries, the second the longest; nostril ori-
Subsequently, all recent authors who have dealt ented dorsally. Dorsum bronze to golden
with the species in Angola considered it a valid brown, uniform or with scattered black speck-
species (Ceríaco et al., 2016a; Marques et al., les; sometimes with a pair of weakly defined
2018; Branch et al., 2019a; Santos et al., 2021; pale dorsolateral stripes; flanks grayish or
Lobón-Rovira et al., 2022). The present work mottled with black and white. Head uniform
presents a considerable change to the concept of or with irregular black speckling; labials
T. punctulata. We have molecular evidence sup- white, sometimes with black spots or vertical
porting the monophyly of all southwestern bars; a subocular white stripe may extend to
Angolan populations, including topotypical the tympanum or forelimb insertion. Venter
material from Curoca River. Available samples uniformly white.
from Namibia indicate that the Namibian popu- Material Examined (* denotes type mate-
lation of what was known as T. punctulata is not rial): Namibe Province: Iona National Park,
FIGURE 79. Specimen of Trachylepis punctulata from Chipumpo, Namibe Province (CAS 263502). Photos by
L.M.P.C.
FIGURE 80. Life photo of Trachylepis punctulata from Virei camp, Namibe Province (CAS 263508). Photo by
Ishan Agarwal.
FIGURE 81. Distribution of Trachylepis punctulata in Angola. Black star denotes the type locality.
Espinheira [-16.7891°, 12.3576°, 452 m] (CAS way to Espinheira (INP) [-16.4777°, 12.4418°,
254793); Virei camp [-16.1196°, 12.8346°, 409 m] (MUNHAC/MB03-001411).
522 m] (CAS 263508, 263509, 263510); Virei- Additional Material: Huíla Province: 50 km W
Chipumpo [-16.2229°, 12.9038°, 593 m] (CAS of Humpata, by roadside in boulders [-15.0507°,
263497, 263498); Virei-Calundolo [-16.3022°, 13.0279°, 464 m] (PEM R1800); Leba Pass [-15.0700°,
12.8008°, 470 m] (CAS 263488); Chipumpo 13.2434°, 1667 m] (TM 46760). Namibe Province: 41
mi NE of Moçamedes [-14.7705°, 12.5942°, 518 m]
[-16.1923°, 12.8565°, 542 m] (CAS 26502); Rio
(CAS 85961); Namibe-Lubango road, marker 59, 1.8
Curoca [-16.2653°, 12.3210°, 188 m] (ZMB
km W of Caraculo, N side of road [-15.0153°, 12.6424°,
6478*); Moçâmedes [-15.2000°, 12.1500°, 24 m] 497 m] (CAS 254903, 255057); Cunene 8 km SW of
(MHNC-UP/REP 263, 264, 265); Baía dos Tigres Oncocua [-16.8583°, 12.6128°, 808 m] (PEM R18004);
[-16.5856°, 11.8271°, 98 m] (MHNC-UP/REP Namibe 16 km E of Iona [-16.7980°, 12.6806°, 783 m]
278); Mucungo farm [-14.7823°, 12.4859°, 286 (PEM R18003); Namibe 9 km South of Camp at the
m] (MUNHAC/MB03-001405–001410); on the Red Canyon [-15.7460°, 12.1399°, 81 m] (PEM R17998);
FIGURE 82. Typical habitat of Trachylepis punctulata in Curoca, Namibe Province. Photo by L.M.P.C.
Mucungu [-14.7824°, 12.4907°, 304 m] (CM S5648); [undetermined locality] (TM 44052); São Nicolau, 17
Cima [-15.0667°, 12.1500°, 57 m] (TM 41243); Porto km [undetermined locality] (TM 41152); Rio Caroca,
Alexandre (currently Tombwa) [-15.8046°, 11.8449°, 8 between Mossamedes and Port Alexandre (currently,
m] (TM 40400); Inamangando River on Lucira Road Tombwa) [undetermined locality] (TM 22554–22556).
[-14.0500°, 12.3833°, 6 m] (TM 41153); 23 km W of Historical Localities (no extant speci-
Virei [-15.6011°, 13.1777°, 476 m] (TM 41020); Namibe mens): Namibe Province: Rio Coroca [-15.7833°,
Omauha Lodge, N of entrance to Iona National Park 12.0667°, 45 m] (Bocage, 1872, 1895, 1897a).
[-15.9968°, 12.4068°, 301 m] (PEM R17999); Namibe Distribution in Angola: This species is
road from Lake Arco to Espinheira rocky valley
known from southwestern areas of the country,
[-15.7460°, 12.1399°, 81 m] (PEM R18001); Espinheira
in Namibe Province and western Huíla Province
[-16.7500°, 12.3667°, 437 m] (PEM R18002); Caraculo
[-15.0167°, 12.6667°, 463 m] (TM 24532); Caraculo, 16 (fig. 81).
km E [-15.5000°, 13.5000°, 1288 m] (TM 23905); Rio Global Distribution: Confirmed records
Curoca mouth [-15.7233°, 11.9139°, 4 m] (TM 40395– are restricted to southwestern Angola. Further
40397, 40399); 30 km N of Tambor [-16.1356°, 12.4297°, work is needed to establish the status of Namib-
379 m] (TM 40492); Pico Azevedo [-15.5500°, 12.5167°, ian populations (labelled as T. cf. triebneri in fig.
347 m] (TM 41062–41069); São Nicolau [-14.2582°, 1). The distribution of the non-Angolan popula-
12.3967°, 17 m] (TM 43963, 43964); Otchifengo river tion has been noted by Portik (2009), Portik and
[-16.6167°, 12.8833°, 560 m] (TM 40773); Foz do Bauer (2012), and Branch (1998).
Cunene [-17.2693°, 11.7961°, 13 m] (TM 40671); 6 km Habitat and Natural History Notes: This
S of Rio Coroca on Iona road [-15.7833°, 12.0667°, 45
species occupies a variety of habitats in the arid
m] (TM 40590); Road to Omauha [-16.1843°, 12.4189°,
365 m] (FKH 0781 fide Lobón-Rovira et al., 2022);
regions of southwestern Angola, from mopane
Tonombe [-17.0964°, 12.3035°, 615 m] (FKH 0791 fide woodlands to steppe-savannah mosaic, or even
Lobón-Rovira et al., 2022); Curoca river at Ialutala the mobile sand dunes of the Kaokoveld desert
[-16.2641°, 12.3277°, 172 m] (AMB 13265). Undeter- (Grandvaux-Barbosa, 1970; fig. 82). It is com-
mined locality: between Mossamedes and Benguela monly found associated with Welwitschia plants.
Trachylepis raymondlaurenti Marques et al., SAV; 48–53 SAD; 24–28 MSR; lamellae beneath
2019—Laurent’s Long-tailed Skink fingers and toes smooth; plantar scales smooth;
20–27 LUFT; 16–21 LUFF; supranasals sepa-
Figures 83, 84, plate 5 rated; parietals usually separated (rarely in
contact); prefrontals separated; frontoparietals
Trachylepis raymondlaurenti Marques et al., always in contact; one pair of enlarged nuchal
2019: 59. Holotype: CAS 258401 (collected scales present; ear opening vertically ovoid
by M.P. Marques, L.M.P. Ceríaco, S. Ban- and smaller than the eye; lacking subtriangular
deira, E. Stanley and J. Vindum). Type auricular scales on the anterior margin of the
locality: “Giant Sable Sanctuary of Can- ear opening. Supralabials usually seven, the
gandala National Park (-9.84606° N, fifth subocular; five supraciliaries, second
16.7223° E, WGS-84; elevation 1101 m),” largest; nostril oriented laterally. Dorsum uni-
Malanje Province, Angola. formly brown in preserved specimens, and
Mabuya megalura: Laurent (1964: 74). greenish brown in live specimens, with some
Trachylepis cf. megalura: Ceríaco et al. (2016c: white speckles running from the labials
71, 2018a: 423); Marques et al. (2018: 264); through the anterior half of the tail. Ventral
Branch et al. (2019a: 318). surfaces uniformly whitish with some scat-
Trachylepis raymondlaurenti: Marques et al. tered dark speckles near the flanks and under
(2019a: 59); Ceríaco et al. (2020a: 402, 423). the tail.
Material Examined (* denotes type mate-
Laurent (1964) provided the first record of rial): Lunda Sul Province: Alto Cuílo
this species for Angola based on two speci- [-10.0167°, 19.5500°, 1264 m] (MD 5309*; MCZ
mens from Alto Cuilo, Lunda Sul Province R-67627). Malanje Province: Cangandala
(specimens still extant in the collections of National Park [-9.8461°, 16.7223°, 1101 m]
Museu do Dundo and Museum of Compara- (CAS 258401*).
tive Zoology; see Ceríaco et al., 2020a). Lau- Distribution in Angola: This species
rent (1964) assigned these specimens to a appears to be restricted to northern regions of
putative new subspecies of Mabuya megalura the country, with records for Malanje and
from Angola and the Katanga region of the Lunda-Norte provinces. This skink may have a
Democratic Republic of the Congo, but wider distribution and potentially occur in the
refrained from providing a description of this neighboring provinces of Bié, Lunda-Sul, and
new “race,” leaving this task for his colleague Moxico (fig. 85).
Gaston-F. de Witte, who had collected and Global Distribution: Trachylepis raymond-
studied a larger series of the same taxon from laurenti is known only from northern Angola
Upemba National Park, Democratic Republic and southeastern Democratic Republic of Congo,
of the Congo (de Witte 1953). However, a although it may extend into neighboring north-
description was never published by de Witte, ern Zambia (Marques et al., 2019a).
and the species was only recently described by Habitat and Natural History Notes:
Marques et al. (2019a) based on a comprehen- This species is usually found in savannahs and
sive review of historical and recent material. miombo woodlands (Grandvaux-Barbosa,
Diagnosis: A medium-sized slender skink 1970; fig. 86). Marques et al. (2019a) noted
(max. SVL 80.0 mm, MD 5309), with fully that the holotype (collected in September
developed pentadactyl limbs (fig. 83, 84); tail 2015) contained 10 well-developed fetuses,
more than twice as long as body; dorsal and suggesting that this viviparous species gives
ventral scales smooth, rarely keeled; 50–58 birth in late September or October.
FIGURE 83. Holotype of Trachylepis raymondlaurenti from Cangandala National Park, Malanje Province (CAS
FIGURE 84. Life photo of the holotype of Trachylepis raymondlaurenti from Cangandala National Park,
Malanje Province (CAS 258401). Photos by L.M.P.C.
FIGURE 85. Distribution of Trachylepis raymondlaurenti in Angola. Black star denotes the type locality.
Trachylepis sulcata (Peters, 1867)— Euprepes sulcatus Peters, 1867: 20 [replacement

Western Rock Skink name for Euprepes olivaceus Peters, 1862].
Figures 87–90, plate 6 Euprepes olivaceus [part] [?]: Bocage (1870: 68)
Mabuia sulcata [part]: Bocage (1895: 41)
Euprepes olivaceus Peters, 1862: 21 [preoccupied Mabuya sulcata ansorgii [part]: Mertens (1938:
by Euprepes olivaceus Gray, 1839]. Syn- 438); Hellmich (1957b: 55); Branch (1998:
types: ZMB 4209, 4210, 64329-30 (formerly 157)
ZMB 4210 part), 64346-47 (formerly ZMB Mabuya sulcata ansorgei: Laurent (1964: 74)
4209), MNHN 1470 (collected by C.H. Mabuya sulcata: Broadley (2000:102)
Hahn). Type locality: “Neu-Bramen” [= Trachylepis sulcata [part]: Portik (2009: 23); Por-
Gross Barmen], Otjozondjupa Region, tik et al. (2010: 147; 2011: 1744); Masterson
Namibia. et al. (2014b: 267); Marques et al. (2018:
FIGURE 86. Typical habitat of Trachylepis raymondlaurenti in Cangandala National Park, Malanje Province.
Photo by L.M.P.C.
269); Ceríaco et al. (2020a: 403); Lobón- patterns in T. sulcata from South Africa and
Rovira et al. (2022: 310). Namibia. Angolan samples were unavailable at
Trachylepis sulcata sulcata: Butler et al. (2019: the time. The results showed that the melanistic
235); Baptista et al. (2019: 110). form T. s. nigra from Namibia is nested within
Trachylepis sulcata ansorgii [part]: Branch et al. the nominal T. s. sulcata, the population struc-
ture of which displayed three distinct groups: a
(2019a: 319).
northern clade from northern Namibia, a cen-
Trachylepis sulcata is a commonly observed tral clade from southern Namibia and western
viviparous skink distributed across the western South Africa, and a southern clade with the
parts of South Africa, Namibia, and into south- remaining South African individuals (Portik et
western Angola (Branch, 1998; Masterson et al., al., 2010, 2011).
2014b; Marques et al., 2018). Two subspecies Until recently, the taxonomy of the Angolan
have been historically recognized: T. sulcata populations of T. sulcata was unresolved due to
nigra (Werner, 1915), a melanistic form from the uncertainties of the taxonomic validity of its
Lüderitz Bay, Namibia, and T. sulcata ansorgii subspecies T. s. ansorgii. This was exacerbated by
(Boulenger, 1907), from the southwestern high- the confusion created by Mertens (1938, 1955,
lands of Angola and Namibia (Branch, 1998; 1971) who referred to the existence of sympatric
Masterson, 2014b; Marques et al., 2018). Previ- populations of T. s. sulcata and T. s. ansorgii in
ous works by Portik et al. (2010, 2011) looked northern Namibia. Branch (1998) perpetuated
at population structure and phylogeographic this confusion. With some preliminary molecu-
FIGURE 87. Male specimen of Trachylepis sulcata from dirt road to Chingo, Namibe Province (CAS 264711).
Photos by L.M.P.C.
FIGURE 88. Female specimen of Trachylepis sulcata from dirt road to Chingo, Namibe Province (CAS 24731).
Photos by L.M.P.C.
FIGURE 89. Life photo of a male Trachylepis sulcata from Tundavala, Huíla Province (CAS 263338). Photo
by L.M.P.C.
FIGURE 90. Life photo of a female Trachylepis sulcata from Tundavala, Huíla Province (CAS 263344). Photo
by L.M.P.C.
FIGURE 91. Distribution map of Trachylepis sulcata in Angola.
lar data, Butler et al. (2019) were the first to pro- province and ranges to Benguela, as well as to the
vide evidence that both taxa were valid, and higher elevation areas of the Angolan Plateau
subsequent authors followed this arrangement (see T. ansorgii account) (Butler et al., in review).
(Baptista et al., 2019; Vaz Pinto et al., 2019; Diagnosis: A medium-sized robust skink
Ceríaco et al., 2020a; Lobón-Rovira et al., 2022). (max. SVL 87.9 mm, CAS 263350), with fully
Branch et al. (2019a) considered all populations developed pentadactyl limbs (figs. 87–90);
of the T. sulcata group in Angola as T. sulcata dorsal scales usually pentacarinate; ventral
ansorgii. Our morphological and molecular data scales smooth; 57–68 SAV; 48–58 SAD; 33–42
suggests that T. sulcata occurs in the southwest- MSR; lamellae beneath fingers and toes spi-
ern lowlands of Angola, especially in the south- nose; plantar scales spinose; 20–27 LUFT;
ern half of the Namibe Province. Trachylepis 16–21 LUFF; supranasals usually in contact;
ansorgii replaces it in the northern half of the parietals usually in contact or touching at a
FIGURE 92. Typical habitat of Trachylepis sulcata in Omauha, Namibe Province. Photo by L.M.P.C.
single point; prefrontals usually separated; irregular dark line or series of spots. Dorsum
frontoparietals in contact; one pair of enlarged uniform golden brown to reddish brown in
nuchal scales present; ear opening vertically adult males; flanks and limbs light grayish; tail
ovoid and smaller than the eye; two to three often with remnants of dark stripes. Ventral
subtriangular auricular scales (shorter than parts whitish; labials, throat, and chin some-
the diameter of ear) extend posteriorly and times yellowish to orange in live specimens;
usually slightly upward from the anterior mar- usually with heavy black speckling, in some
gin of the ear opening. Supralabials usually 7 cases restricted to the labials and in others
or 8, the sixth is subocular; usually five supra- extending to the sides and top of the head.
ciliaries, second largest; nostril oriented later- Material Examined: Huíla Province:
ally. Color pattern exhibits pronounced sexual Cristo Rei, Lubango [-14.9401°, 13.5117°, 2124
dimorphism. Adult females and juveniles are m] (CAS 263333, 263336, 263338–263341);
pale to golden brown above, with six dark lon- Tundavala, Lubango [-14.8239°, 13.38114°,
gitudinal stripes (a pair of paravertebral 1295 m] (CAS 263342–263345, 263348–
stripes, a pair of dorsolateral stripes, and 263350); Leba Pass [-15.0700°, 13.2434°, 1667
another pair on the flanks) that usually con- m] (CAS 254875); Mangueiras near Leba
tinue on the tail as irregular lines or series of [-15.0446°, 13.1586°, 653 m] (CAS 254886,
spots; lower flanks whitish, sometimes with an 254888); Capelongo, near Bicuar National Park
entrance [-14.8443°, 15.0356°, 1220 m] savannah-steppe mosaic (Grandvaux-Barbosa,

(MHNC-UP/REP 460, 461). Namibe Province: 1970; fig. 92), but occupies more heavily vege-
Virei-Calundolo [-16.3102°, 12.7960°, 471 m] tated areas in parts of inland Namibia and
(CAS 263484, 263485); Curoca-Omauha South Africa (A.M.B., personal obs.). Females
[-15.9503°, 12.4953°, 394 m] (CAS 263520); give birth to three to five young and may have
Virulundo [-16.2949°, 12.9408°, 759 m] (CAS two broods per year (Broadley, 2000). Butler et
263489–263491); Iona National Park, Curoca al. (2019) noted that in Huíla Province many
river, Pediva Hot Springs Area [-16.2836°, females were found gravid in August, suggest-
12.5611°, 250 m] (CAS 254853); Omauha-Chi- ing that the reproductive season may be in win-
tundolo [-15.8706°, 12.9030°, 553 m] (CAS ter and spring.
263516); dirt road to Chingo (approx. 31 km N
of Chingo) [-14.4816°, 12.8046°, 449 m] (CAS
Trachylepis suzanae, sp. nov.—
264711, 2674712); dirt road to Chingo (approx.
Suzana’s Wedge-Snouted Skink
18 km N to Chingo) [-14.3934°, 12.8289°, 620
m] (CAS 264731); Bentiaba River near Maungo Figures 93–95, plate 5
[-14.5106°, 12.8391°, 417 m] (CAS 264723);
Maungo houses [-14.8443°, 15.0356°, 1220 m] Euprepes acutilabris [part]: Bocage (1870: 68);
(CAS 264717). Peters (1877: 614).
Additional Material: Cunene Province: Otsch- Mabuia acutilabris [part]: Boulenger (1887: 208);
injau [-16.5790°, 13.9052°, 1218 m] (ZSM 247/1953, Bocage (1895: 46); Monard (1937: 94).
253/1953, 273/1953, 276/1953). Huíla Province: 11 Mabuya acutilabris [part]: Schmidt (1919: 551);
km south of Chibemba [-15.8439°, 14.1107°, 1275 m]
Parker (1936: 138); Laurent (1947: 8, 1954:
(Baptista et al. 2019). Namibe Province: Munhino
[-14.9667°, 12.9667°, 393 m] (MD 1918); Otchifengo 65, 1964: 75); Hellmich (1957a: 58); Branch
River [-16.6167°, 12.8833°, 560 m] (TM 40775, 40776); (1998: 151).
Espinheira [-16.7885°, 12.6127°, 455 m] (PEM Trachylepis acutilabris [part]: Marques et al.
R17975); Tchamalindi [-16.9757°, 12.8856°, 1436 m] (2018: 252); Branch et al. (2019a: 318);
(P1.97–8, fide Lobón-Rovira et al., 2022); Epupa Ceríaco et al. (2020a: 402).
[-16.9968°, 13.2486°, 649 m] (AMB 13222, 13199,
13208, 13030). This species has previously been confounded
Historical Localities (no extant speci- with the phenotypically similar T. acutilabris.
mens): Huíla Province: Capangombe (Rio Bocage (1870) first cited Mabuia acutilabris for
Chimba et Biballa) (Bocage, 1895). Angola and was followed by several authors who
Distribution in Angola: This species reported specimens from most of the Angolan
appears to be restricted to the southwestern coastal regions (Peters, 1877; Bocage, 1895;
regions of the country, with records for southern Schmidt, 1919; Monard, 1937; Laurent, 1947,
Namibe, western Cunene, and southwestern 1954, 1964; Hellmich, 1957a; 1957b; Haacke,
Huíla provinces (fig. 91). 2008; Ceríaco et al., 2016b; Vaz Pinto et al.,
Global Distribution: Trachylepis sulcata is 2019). Recently collected material allowed us to
widely distributed from southwestern Angola identify two divergent lineages in Angolan popu-
through Namibia to western South Africa lations, forming a clade sister to true Trachylepis
(Branch, 1998; Masterson, 2014b). acutilabris from Namibia (fig. 1). Populations
Habitat and Natural History Notes: from northwestern Angola previously assigned
This rupiculous species is usually found as to T. acutilabris are here described as T. suzanae,
groups of several individuals on granite out- sp. nov., based on specimens collected at the
crops in the arid habitats of southwestern mouth of the Kwanza River, Luanda Province.
Angola, including dry mopane woodlands and Historical records from Cabinda (Peters, 1877),
FIGURE 93. Holotype of Trachylepis suzanae, sp. nov., from Kissama National Park, Luanda Province (CAS
FIGURE 94. Life photo of the holotype of Trachylepis suzanae, sp. nov., from Kissama National Park, Luanda
Province (CAS 263357). Photo by John Cavagnaro.
Expedition in 1925, are deposited in the AMNH

and remained unpublished until now, as well as
a small series from Luanda Province deposited at
the USNM, collected at the end of the 19th cen-
tury by both Héli Chatelain and the U.S. Eclipse
Expedition. The now lost specimens from
“Duque de Bragança,” Malanje Province (Bocage,
1895) represent a dubious record, as it is the only
inland locality cited for the species.
Diagnosis: A medium-sized skink (max. SVL
62.3 cm), with fully developed, pentadactyl limbs
and a wedge-shaped snout (figs. 93–95). Dorsal
scales tricarinate or quadricarinate; ventral scales
smooth; 48–56 SAV; 43–49 SAD; 26–32 MSR;
lamellae beneath fingers and toes spinose; plantar
scales spinose; 19–26 LUFT; 12–15 LUFF; suprana-
sals in contact or separated, with frequent occur-
rence of intrusive scales and scale fusion (see
Schmidt, 1919 for further detail on the variation of
nasal scales arrangement); parietals in contact; pre-
frontals usually separated; frontoparietals usually in
contact; one pair of enlarged nuchal scales present;
ear opening vertically ovoid and smaller than the
eye; three subtriangular auricular scales (shorter
than the diameter of ear) extend posteriorly and
usually slightly upward from the anterior margin of
the ear opening. Dorsum olive to grayish brown,
with three white to yellowish longitudinal stripes;
between dorsal stripes usually scattered pale spots,
and dark, irregular transverse bands that extend to
the upper flanks. A white lateral stripe starts behind
FIGURE 95. Details of head morphology of Trachy-
lepis suzanae. Drawings by A.T. the eye and extends to the hind-limb insertion;
lower flanks white, usually with light grayish speck-
Luanda (Hellmich, 1957a), and Benguela prov- ling. Limbs brown above; hind limbs with black-
inces (Boulenger, 1887; Parker, 1936; Monard, edged, white circles or irregular shapes posteriorly.
1937; Laurent, 1947, 1954, 1964) are also referred Top of head uniform brown; labials white, usually
to this species. Schmidt (1919) recorded “Mabuya with light grayish speckling. Venter white, with
acutilabris” from several localities along the light grayish speckling near the flanks, under the
Congo River in northern Angola and southern tail, hands and feet.
Democratic Republic of the Congo, but only Holotype: An adult female (CAS 263357,
those from Santo António do Zaire, Zaire Prov- field number MCZ-A 36480; fig. 93, 94) collected
ince represent Trachylepis suzanae the remaining at Kissama National Park, Kwanza River mouth,
specimens being representatives of T. albopunc- southern bank, Luanda Province [-9.3510°,
tata (see respective account). Additional speci- 13.1532°, 5 m], by Mariana P. Marques, Luis M.P.
mens of this species from Benguela Province, Ceríaco, John Cavagnaro, and Phillip Pastor, on
collected by the Lang-Boulton Vernay Angola 3 June 2016.
TABLE 8
2024
Mensural and Meristic Data for the Type Series of Trachylepis suzanae, sp. nov.
Abbreviations are listed in the Materials and Methods. Measurements are presented in millimeters and ratios as percentages
CAS AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH
263357 11101 11103 11106 11124 11128 11137 11151 11153 11162 11163 48542 48553 48555 48605 48644
Holo- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para-
type type type type type type type type type type type type type type type type
Sex ♀ ♂ unsexed unsexed unsexed ♂ unsexed ♀ ♂ ♀ unsexed unsexed ♂ ♂ unsexed ♂
SVL 41.6 56.9 52.4 54.3 55.5 56.6 57.4 52.6 55.9 54.8 54.9 62.3 57.5 57.4 58.3 50.0
TL 60.8 97.5 87.4 95.1 102.9 103.4 107.8 83.1 88.3 103.6 77.2 – 106.6 109.9 – –
TL/SVL 150 170 170 170 180 180 180 160 170 180 174 – 180 190 – –
HL 10.6 14.9 12.3 13.2 13.5 13.4 13.6 13.2 12.4 13.6 12.5 14.1 14.1 14.1 13 12.9
HL/SVL 30 30 20 20 20 20 20 20 20 20 20 20 20 20 20 20
SVL/HL 390 380 420 410 410 420 420 390 450 400 440 410 410 410 450 390
HW 6.2 9.6 8.1 8.6 8.6 9.1 8.8 8.3 8.9 8.5 8.7 8.5 9.1 9.1 8.4 8.1
HW/HL 60 60 60 60 60 60 60 60 70 60 70 60 60 60 60 60
HH 4.5 6.8 5.7 6.4 6.9 6.6 6.5 6.5 6.4 6.6 6.4 7.1 6.5 7.0 6.4 6.7
IN 1.5 2.3 1.5 1.8 2.1 1.9 2.1 1.9 2.3 2.1 2.1 2.2 2.0 2.2 2.2 1.9
EN 3.5 4.5 3.7 4.2 3.9 3.9 3.9 3.8 4.1 3.7 4.0 4.2 4.6 4.8 4.1 4.0
ES 4.6 6.7 5.2 6.0 5.7 5.9 5.8 5.4 6.2 5.5 5.6 6.1 6.2 6.1 5.8 5.6
MSR 29 29 28 29 29 29 29 28 29 29 30 30 29 30 32 31
SAD 49 44 43 45 47 47 44 45 45 47 47 44 48 49 45 43
SAV 56 51 51 52 55 52 53 53 52 53 48 54 49 57 51 53
LUFF 15 13 13 14 14 – 13 14 14 13 14 12 15 13 14 15
LUFT 26 21 22 20 19 22 20 22 20 20 20 26 24 23 22 22
SC 5 6 5 5 5 5 5 5 7 6 5 5 5 5 5 5
SL (SO) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 7 (5) 6 (5) 7 (5)
CP C SPC C SPC SPC C C C C C S C C C SPC C
CFP C C C C C C C C C C C C C C C C
CSN C Sa C Sa Sa Sa C C C Sa Sa C C C C C
CPF C S S S S S S S S S S S SPC S S S
KDS 3–4 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3
Plantar spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose
scales
115
aSeparated by small internasal scale.

FIGURE 96. Distribution of Trachylepis suzanae in Angola. Black star denotes the type locality.
Paratypes: Two adult females (AMNH [-12.2506°, 13.7116°, 83 m] by Herbert Lang and
11151, 11162), three adult males (AMNH Rudyerd Boulton on April 1925.
11101, 11128, 11153) and five unsexed adults Additional Material: Benguela Province: Lobito
(AMNH 11103, 11106, 11124, 11137, 11163) Bay [-12.3500°, 13.5500°, 7 m] (AMNH 48645, 147566–
collected at “St. Antonio” [do Zaire], Zaire 147579); Hanha [-12.2506°, 13.7116°, 83 m] (AMNH
Province [-6.1333°, 12.3667°, 4 m] by Herbert 40655, 41548–41561, 48530–48541, 48543–48552, 48554,
Lang and James P. Chapin on August 1915; one 48556–48604, 48606–48643); Benguela [-12.5833°,
13.4167°, 15 m] (BMNH 1906.8.24.65–67); Lobito, Rest-
adult male (AMNH 48644) collected at “Lobito
inga [-12.3292°, 13.5661°, 6 m] (MD 1265.22); Lobito
Bay,” Benguela Province [-12.3500°, 13.5500°, 7
[-12.3709°, 13.5542°, 3 m] (BMNH 1936.8.1.593–596,
m] by Herbert Lang and Rudyerd Boulton on MHNC 91.0496); Chimalavera Nature Reserve, vic main
April 1925; two adult males (AMNH 48553, camp [-12.8338°, 13.1699°, 250 m] (CAS 263306–263309;
48555) and two unsexed adults (AMNH 48542, UF 187301, 187302). Luanda Province: Kissama
48605) collected at “Hanha,” Benguela Province National Park, Cabo Ledo [-9.6777°, 13.3881°, 26 m]
FIGURE 97. Typical habitat of Trachylepis suzanae at the mouth of Kwanza River, Luanda Province. Photo by
Luis Querido.
(AMB 12428–38, 12440–42); Kissama National Park, with a poorly defined neck and well-developed
Kwanza river mouth [-9.3510°, 13.1532°, 14 m] (AMB pentadactyl limbs; tail long, its length almost
12466–73); Luanda [-8.7949°, 13.2661°, 1 m] (USNM twice the SVL, smoothly tapering. Fore- and hind
20045, 20046); Cabo Ledo [-9.6556°, 13.2392°, 101 m]
limbs overlap when adpressed against the body.
(USNM 16105, 16106). Zaire Province: “St. Antonio”
[do Zaire], Zaire Province [-6.1333°, 12.3667°, 4 m] SVL 41.6 mm, TL 60.8 mm. HL 10.6 mm, with
(AMNH 11099, 11100, 11102, 11104, 11105, 11107– wedge-shaped snout. Additional measurements
11123, 11125–11127, 11129–11136, 11138–11150, 11152, are presented in table 8. Four subtriangular auric-
11154–11161, 11164–11193). ular scales extend posteriorly from the anterior
Historical Localities (no extant speci- margin of the ear opening, the lowermost dis-
mens): Benguela Province: Benguela [-12.5833°, tinctly smaller. Rostral visible from above. Nostrils
13.4167°, 15 m] (Bocage, 1895); Catumbela oriented dorsally, set posteriorly, so that postnasal
[-12.4333°, 13.5000°, 7 m] (Bocage, 1895); “Plage effectively borders nostril. Supranasals in contact
de Lobito” (Laurent, 1964); “Baía dos Elefantes” at a single point. Frontonasal broader than long,
[-13.2323°, 12.7317°, 5 m] (Laurent, 1947: 8); in contact with loreal scales. Prefrontals pentago-
“Mullet Bay” [-13.4051°, 12.5735°, 138 m] (Lau- nal, in contact, each in contact with the following
rent, 1947: 8); “Baía Farta” [-12.6000°, 13.2000°, head shields: frontonasal, posterior loreal, first
4 m] (Laurent, 1947: 8); “Baía de St. Bras, près supraocular and frontal. Two loreals. Frontal
Lobito” (Laurent, 1947: 8). Cabinda Province: length similar to the distance between anterior tip
Chinchoxo [-5.1000°, 12.1000°, 45 m] (Peters, of frontal and tip of snout. Frontal in contact with
1877). Luanda Province: Ilha de Luanda three supraoculars on each side. Two frontopari-
[-8.7789°, 13.2437°, 0 m] (Hellmich, 1957a). etals, in contact with each other and the frontal,
Description of the Holotype: A well-pre- third and fourth supraoculars (left frontoparietal
served adult female. Body cylindrical and robust in contact only with third supraocular), parietal,
and interparietal. Frontoparietal plus interparietal suzanae differs from all other species of Trachyl-
length shorter than frontal length. Interparietal epis known to occur in Angola, with the excep-
longer than broad, with a visible parietal foramen. tion of T. wilsoni by having a wedge-shaped
Parietals greater than frontoparietals, in narrow snout. It differs from its sister taxon, T. wilsoni,
contact with each other. Five supraciliaries, sec- by having a more acuminate head and less prom-
ond largest. Seven supralabials; subocular narrow, inent canthus rostralis (fig. 95). This difference
not reaching the lip. Seven infralabials. Postmen- also applies to true T. acutilabris from Namibia,
tal and right primary chin shield fused into a large which also has a higher number of MSR than T.
and irregular scale, in contact with the mental, suzanae (28–32 in T. suzanae vs. 32–39 in T.
first two left infralabials, first three right infralabi- acutilabris).
als, left primary chin shield, right secondary chin Distribution in Angola: This species
shield, and two gular scales. Transparent scale occurs in the coastal regions of northern and
present in lower eyelid, as is usual for Trachylepis. central Angola, from Benguela Province north-
Tympanum visible, at same level as mouth. Dorsal ward to Zaire Province (fig. 96).
scales each with three smooth keels. Ventral scales Global Distribution: Confirmed records of
smooth. MSR 29, SAD 49, SAV 56. Limbs with this species are limited to Angola, but its pres-
five digits; scales on palms and soles spinose. Rela- ence at the mouth of the Congo River and the
tive length of fingers IV > III > V > II > I, relative historical record from Chinchoxo, Cabinda
length of toes IV > III > V > II > I. Finger-IV Province (Peters, 1877) suggest that it is present
lamellae 15, Toe-IV lamellae 26. Color in life is in adjacent Democratic Republic of Congo and
homogenously light brown on the flanks, upper possibly in the Republic of Congo as well.
side of head, neck, dorsum, legs and tail brown Habitat and Natural History Notes: This
with three pale dorsal stripes; vertebral stripe psammophilous species is found in a forest-savan-
starts at the nape and extends to the base of the nah mosaic or miombo woodlands, in habitats
tail; a pair of dorsolateral stripes starts behind the with sandy substrate, especially near the shore and
eye and extend to the base of the tail. Scattered along riverbanks (Grandvaux-Barbosa, 1970; fig.
pale spots and irregular black spots between dor- 97). These skinks have been recorded on the sandy
sal stripes; the latter form transverse bands beaches of Luanda Island (Hellmich, 1957a) and
extending to the flanks. Hind limbs covered above Lobito (Laurent, 1964), and are known to dig bur-
by black-edged, pale circles or irregular shapes. A rows in the sand at the mouth of the Congo River,
white lateral stripe starts below the eye and sometimes so close to the shore that they are in
extends to the hind limb insertion; lower flanks reach of the waves, although slightly vegetated
white, with grayish speckling. Top of head uni- areas are preferred (Schmidt, 1919). Laurent
formly brown; labials, subocular and lower half of (1954) recorded the species under the layer of
loreals white, with light grayish speckling. Ventral fallen needles in coniferous woods. Schmidt
surfaces uniformly whitish with some scattered (1919) also noted that a fishery at the mouth of
dark speckles near flanks, under tail, hands, and the Congo River formed a great attraction for
feet. these lizards, with fish spoils and waste which in
Coloration in Ethanol: Same as in life (see turn attracted a variety of insects.
above). Etymology: The specific epithet “suzanae” is
Variation: Variation in scalation and mea- formed in the genitive singular and is feminine. It
surements among the type series is reported in is given in honor of Suzana A. Bandeira (1991–),
table 8. All paratypes generally agree with the Angolan herpetologist and conservationist.
holotype in terms of coloration. Suzana has been a fundamental part of our team
Comparison with Other Angolan and since the beginning of our joint herpetological
Southwest African Trachylepis: Trachylepis studies in the country. Suzana initiated her scien-
tific career collecting Trachylepis and other rep- from subocular to anterior border of ear open-
tiles in the Namibe Province in 2013 and is now ing; the ones from Botswana, which are “uniform
an established and respected member of the apart from the white lateral stripe”; and finally
Angolan scientific community. We suggest “Suza- the topotypical Angolan population (see T.
na’s Wedge-snouted Skink” and “Lagartixa de punctulata account). A recent visit to the collec-
Suzana Bandeira” as the English and Portuguese tions of the NMZB allowed us to review Broad-
common names, respectively, for this species. ley’s (1974b, 2000) specimens from Botswana,
Zambia, Zimbabwe, and Mozambique and con-
firm these morphological data. Portik and Bauer
Trachylepis vunongue, sp. nov.— (2012) supported the existence of cryptic specia-
Mwene Vunongue Skink tion within the T. punctulata species complex,
noting that part of the populations from North-
ern Cape province in South Africa were geneti-
Mabuya variegata punctulata [part]: Broadley cally similar to those from Namibia but separate
(1974b: 7). from those from Limpopo and Zimbabwe.
Mabuya punctulata [part]: Broadley (2000: 100). The available data allow us to confirm some
Trachylepis punctulata: Pietersen et al. (2021: of Broadley’s (1974b) observations and to start
276). solving some of the taxonomic uncertainties
Trachylepis punctulata [part]: Marques et al. surrounding the difficult T. punctulata species
(2018: 267); Branch et al. (2019: 319). complex. Sequenced specimens from Namibia
Trachylepis cf. punctulata: Conradie et al. (2022: morphologically agree with the coloration pat-
203). tern (presence of well-defined, pale lateral and
vertebral stripes) and morphological characters
The presence in Angola of this taxon has only presented by Mertens (1954) in the description
very recently been confirmed by Conradie et al. of Mabuya longiloba triebneri. This taxon, con-
(2022), as Trachylepis cf. punctulata, and by a sidered by Broadley (1974b, 2000) as a synonym
recently collected specimen from Cuatir, Cuando of T. punctulata, was described from “Osona bei
Cubango Province (this study). According to our Okahandja” in central Namibia (Mertens 1954).
molecular results, these animals belong to a The geographic origin of our sequenced speci-
group containing T. hilariae, T. ovahelelo and T. mens includes areas surrounding the triebneri
variegata, which is sister to a clade comprising type locality and the overall distribution of the
true T. punctulata and T. cf. triebneri from Namibian population noted in Broadley
Namibia, and it presents clear morphological (1974b). Additional material available in the
and coloration differences from its closest rela- collections of CM (CM 115685, 115686) also
tives. These differences had already been noted morphologically agree with both the sequenced
by Broadley (1974b), who considered that T. specimens and the descriptions provided by
punctulata sensu lato comprised different, mor- Mertens (1954) and Broadley (1974b). These
phologically distinctive populations: those in data lead us to consider that triebneri may rep-
central Namibia, whose specimens presented resent a valid species. Research is underway to
“well defined pale lateral and (sometimes) verte- review the taxonomic status and distribution of
bral stripes”; those in Gordonia District in the the T. punctulata species complex in Namibia,
Northern Cape Province, South Africa, with and therefore we refrain from elevating trieb-
fused paravertebral black streaks forming con- neri to a full species status yet, preferring to list
tinuous stripes; the Zambian and northwestern it as T. cf. triebneri (fig. 1).
Zimbabwe populations that are uniform gray- Specimens from western Zambia and north-
brown above, apart from a white stripe extending western Zimbabwe, studied by Broadley (1974b)
FIGURE 98. Preserved holotype of Trachylepis vunongue, sp. nov., from Cuatir main camp, Cuando Cubango
Province (MUNHAC/MB03-001520). Photos by L.M.P.C.
FIGURE 99. Life photo of the holotype of Trachylepis vunongue from Cuatir main camp, Cuando Cubango
Province (MUNHAC/MB03-001520). Photo by D.P.
FIGURE 100. Distribution of Trachylepis vunongue in Angola.
and reviewed by us (tables 1, 9), agree entirely Angola 46.0 mm, PEM R 23255 (fide Conradie
with the morphological data provided for the et al., 2022), with fully developed pentadactyl
Angolan populations by Conradie et al. (2022) limbs (figs. 98, 99); dorsal scales pentacarinate;
and for the western Zambian populations pro- ventral scales smooth; 45–55 SAV; 40–52 SAD;
vided by Pietersen et al. (2021), as well as our 30–35 (36 fide Broadley, 2000) MSR; lamellae
recently collected specimen from Cuatir, which beneath fingers and toes keeled and spinose;
is uniform gray-brown above, apart from a white plantar scales spinose; 15–22 (28 fide Broadley,
stripe extending from subocular to anterior bor- 2000) LUFT; 9–14 LUFF; supranasals in contact;
der of ear opening. parietals in contact, at least at a single point; pre-
Diagnosis: (Based on Conradie et al., 2022, frontals separated; frontoparietals in contact; one
and the type series, including specimens from pair of enlarged nuchal scales present; ear open-
Zambia and Zimbabwe partly studied by Broad- ing vertically ovoid and smaller than the eye,
ley, 2000) A medium-sized skink (max. SVL in usually with two (2–4 fide Broadley, 2000) lan-
FIGURE 101. Typical habitat of Trachylepis vunongue at Cuatir main camp, Cuando Cubango Province. Photo
by D.P.
ceolate auricular scales on the anterior margin of Holotype: An unsexed adult (MUNHAC/
the ear opening. Supralabials seven to eight MB03-001520, field number LMPC 1452; fig. 98,
(rarely nine); supraciliaries usually five, some- 99) collected at Cuatir main camp, Cuando
times four or six; nostril oriented dorsally. Dor- Cubango Province [-16.4852°, 18.2030°, 1148
sum light gray-brown. There may be black and m], Angola, by Diogo Parrinha on 21 February
pale spots irregularly scattered on the dorsum, 2023.
flanks, limbs, and tail. Top of head uniformly Paratypes: An unsexed adult (NMZB 16016,
colored or with irregular dark vermiculation; field number BFP/R 114) and two adult females
labials dirty white, uniform or with diffuse gray- (NMZB 16017–16018, field numbers BFP/R 114)
ish stippling; a dark stripe runs from nostril, collected at Ndau School, Western Province
through the eye, to ear opening; immediately [-15.4415°, 23.2227°, 1044 m], Zambia, by Don-
below, a white stripe starts at subocular position ald G. Broadley and Shiela Broadley on 27 March
and stops at ear opening, rarely extending 1999; an unsexed adult (NMZB 16006, field
beyond ear opening to forelimb insertion. Ven- number BFP/R 111) collected at Ndau School,
tral parts white to bluish white, sometimes with Western Province [-15.4415°, 23.2227°, 1044 m],
irregular dark spotting, especially on subcaudal Zambia, by Peter van Daele on 26 March 1999;
and gular region, and near flanks. an unsexed adult (NMZB-UM 21008) collected
at Kalabo, Barotseland [-14.9928°, 22.6785°, 1026 with loreal scale. Prefrontals subquadrangular,
m], Zambia, by R.G. Japp on unknown date; an separated, each in contact with the following
unsexed adult (NMZB 13382, field number GR head shields: frontonasal, loreals, first and sec-
83) collected at Makona Pan, Hwange National ond supraocular, and frontal. Two loreals. Fron-
Park [-19.3167°, 26.9167°, 993 m], Zimbabwe, by tal length slightly greater than the distance
G.S.A. Rasmussen on 9 March 1994; an unsexed between anterior tip of frontal and tip of snout.
adult (NMZB 13589, field number GSA 279) col- Frontal in contact with two supraoculars on
lected at Shaba Shaba, Hwange National Park each side, plus an intrusive scale between sec-
[-19.3167°, 26.1667°, 976 m], Zimbabwe, by ond and third supraoculars on right side. Two
G.S.A. Rasmussen on 9 May 1994; two unsexed frontoparietals, in contact with each other and
adults (NMZB 13563, 13564) collected at Shak- the frontal, third and fourth supraoculars, pari-
wanki Pan 0,5km NW, Hwange National Park etal, and interparietal. Frontoparietal plus inter-
[-19.1297°, 26.2303°, 976 m], Zimbabwe, by parietal length slightly shorter than frontal
G.S.A. Rasmussen on 6 May 1994. length. Interparietal longer than broad, with a
Additional Material: Cuando Cubango Prov- visible parietal foramen. Parietals greater than
ince: Cuanavale River source [-14.8547°, 19.2864°, frontoparietals, in contact with each other at a
1201 m] (PEM R23371). Moxico Province: single point. Four supraciliaries, second largest.
Cacundu falls [-13.7739°, 18.7552°, 1281 m] (PEM
Eight supralabials, sixth subocular. Eight infral-
R23255); Cuanavale River [-13.9948°, 19.1492°,
1251 m] (PEM R23372); Cuando River source abials. Postmental bordering seven scales (men-
[-13.0035°, 19.1275°, 1343 m] (PEM R23425–6); tal, two infralabials on each side, and two
Quembo River, trap 4 [-13.1359°, 19.0471°, 1368 primary chin shields). Transparent scale present
m] (PEM R23461–2); Sombanana village river in lower eyelid, as is usual for Trachylepis. Tym-
[-12.3071°, 18.6235°, 1408 m] (PEM R23504); panum visible, at same level as mouth. Dorsal
Quembo River source camp [-13.1456°, 19.0457°, scales each with five smooth keels. Ventral
1422 m] (PEM R23550–2); Lungwebungu River scales smooth. MSR 32, SAD 47, SAV 54. Limbs
trap 3 [-12.5806°, 18.6642°, 1302 m] (PEM
with five digits; scales on palms and soles spi-
R23981–2); Quembo River bridge camp
[-13.5275°, 19.2806°, 1241 m] (PEM R27438–40; nose. Relative length of fingers IV > III > II > V
WC-6769 fide Conradie et al. 2022); Lake Hundo > I, relative length of toes IV > III > V > II > I.
[-14.9743°, 21.6297°, 1100 m] (WC-6942 fide Finger-IV lamellae 14, Toe-IV lamellae 19. Col-
Conradie et al. 2022). oration in life homogeneously grayish brown on
Description of the Holotype: A well- dorsum, flanks, dorsal aspect of head, limbs,
preserved, unsexed adult. Body cylindrical and and tail; few white and black speckles on flanks,
robust with a poorly defined neck and well- barely distinct. Top of head with irregular dark
developed pentadactyl limbs; tail moderate, its stippling; a black stripe runs from through the
length slightly greater than the SVL, smoothly eye to ear opening; below, a white stripe starts
tapering. Fore- and hind limbs overlap when at subocular position and stops at ear opening;
adpressed against the body. SVL 41.1 mm, TL labials creamy white, with diffuse grayish stip-
45.5 mm. HL 9.2 mm, with relatively long and pling. Ventral parts creamy white with diffuse
thin snout. Additional measurements are pre- grayish stippling, especially near flanks, under
sented in table 9. Ear opening medium. Two tail and gular region.
subtriangular auricular scales extend posteri- Coloration in Ethanol: Same as in life (see
orly from the anterior margin of the ear open- above).
ing. Rostral visible from above. Nostrils oriented Variation: Variation in scalation and measure-
dorsally and set posteriorly, so that postnasal ments among the type series is reported in table 9.
effectively borders nostril. Supranasals in con- All paratypes generally agree with the holotype in
tact. Frontonasal broader than long, in contact terms of scalation and coloration. In NMZB 14564
and 16016 there is a pair of barely distinct, pale Distribution in Angola: Trachylepis
dorsolateral stripes, while in NMZB 16016 and vunongue is know from the southeastern parts of
13563 the pale subocular stripe extends beyond the the country, in the Cuanavale, Cuanda, and
ear opening to the forelimb insertion. Quembo river sources, as well as the Cuatir basin
Comparison with Other Angolan and (fig. 100).
Southwest African Trachylepis: Trachylepis Global Distribution: The global distribu-
vunongue, sp. nov., differs from all other species tion of this taxon is poorly known, but appears
of Trachylepis known to occur in Angola, with to be restricted to the Kalahari basin, from
the exception of T. albopunctata, T. damarana, southeastern Angola to southwestern Zambia
T. chimbana, T. bouri, T. bocagii, T. attenbor- and northwestern parts of Zimbabwe (Broadley
oughi, T. wahlbergii, T. sulcata, T. ansorgi, and 1974b; Pietersen et al. 2021). The species also
T. hilariae by having spinose plantar scales. It likely occurs in northeastern Namibia and
differs from T. attenboroughi and T. wahlbergii Botswana, but these populations need to be criti-
by the presence of subtriangular auricular scales cally reviewed to confirm their taxonomic iden-
on the anterior margin of the ear opening (vs. tity, as already noted by Conradie et al. (2022).
absence in the latter). Trachylepis vunongue is Habitat and Natural History Notes:
readily distinguished from T. suzanae and T. According to Conradie et al. (2022), the species
wilsoni by not having a wedge-shaped snout (vs. is common where it occurs and often found
wedge shaped in the latter two). The new spe- moving around on the sandier regions, in close
cies can be readily distinguished from T. albo- proximity to water sources. This is concordant
punctata, T. damarana, T. ansorgii, and T. with our own observations at Cuatir, where the
sulcata by having its nostrils situated more dor- species was found on sandy substrate close to
sally, directed upward (vs. nostrils situated the river floodplains (D.P., personal obs.; fig.
more laterally, directed sideward in the latter). 101). Pietersen et al. (2021) noted that the spe-
It can be distinguished from T. bouri and T. cies occurs in arid and mesic savannahs, and
bocagii by having <36 MSR (vs. >35 in the lat- forages between grass tussocks in sandy regions,
ter). It can be distinguished from T. ovahelelo by feeding on insects and spiders. Females give
having the dorsum uniform or with irregularly birth to 2–4 young in March (Pietersen et al.
scattered black and pale speckles (vs. trans- 2021). The female paratype NMZB 16017, col-
versely aligned series of black speckles in the lected on 17 March 1999, contained a fully
latter) and a shorter tail (TL/SVL 111–152% vs. developed embryo.
190% in T. ovahelelo). It can be distinguished Etymology: The specific epithet
from T. chimbana by having 45–55 SAV and “vunongue” is a noun in apposition and is given
9–14 LUFF (vs. 55–63 and 15–17, respectively, in honor of the Mwene (= King) Vunongue
in T. chimbana). In comparison with its sister (1800–1886), king of the N’Ganguelas and
taxon T. punctulata, T. vunongue presents a one of the most important Angolan chieftains,
light gray-brown dorsum (vs. bronze to golden- whose kingdom spanned approximately the
brown in T. punctulata). It differs from T. hilar- same region as the known distribution of the
iae by having 30–35 MSR (vs. 29–30 in the new species. As a curiosity, the name of the
latter), 45–55 SAV (vs. 58), and 9–14 LUFF (vs. capital city of Cuando Cubango Province—
15–16 in T. hilariae). Regarding the other Menongue—derivates from the admixture of
Southwestern African congeners of the varie- the words “Mwene” and “Vunongue.” We pro-
gata subgroup sensu Weinell et al. (2019), T. pose the Portuguese common name of “Lagar-
vunongue is readily differentiated from T. varie- tixa de Mwene Vunongue,” and the English
gata by having five keels on dorsal scales (vs. common name of “Mwene Vunongue Skink”
three in T. variegata). for this species.
TABLE 9
Mensural and Meristic Data for the Type Series of Trachylepis vunongue, sp. nov.
MUHNAC/ NMZB–
NMZB NMZB NMZB NMZB NMZB NMZB NMZB NMZB
MB03– UM
16016 16017 16018 13382 13589 16006 13563 13564
001520 21008
Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
Sex unsexed unsexed female female unsexed unsexed unsexed unsexed unsexed unsexed
SVL 41.1 39.5 40.7 44.8 43.5 35.1 43.3 37.5 40.0 38.7
TL 45.5 – 57.4 63.0 – 53.3 53.7 – – 47.0
TL/SVL 1.1 – 1.4 1.4 – 1.5 1.2 – – 1.2
HL 9.2 9.4 8.4 9.1 9.0 8.1 9.0 8.3 8.8 8.7
HL/SVL 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.2
SVL/HL 4.5 4.2 4.9 4.9 4.8 4.4 4.8 4.5 4.5 4.4
HW 6.0 6.1 5.9 6.2 6.1 5.4 6.0 5.4 5.6 5.6
HW/HL 0.7 0.7 0.7 0.7 0.7 0.7 0.7 0.7 0.6 0.6
HH 4.4 4.7 4.6 4.8 5.0 4.0 4.4 3.9 4.2 4.4
IN 1.0 1.5 1.3 1.4 1.2 0.8 1.2 1.1 1.3 1.1
EN 3.0 2.3 2.3 2.0 2.8 2.0 2.5 2.3 2.8 2.4
ES 3.9 3.4 3.5 3.6 3.9 3.4 3.4 3.4 3.8 3.8
MSR 32 31 32 31 32 32 30 30 32 31
SAD 47 45 48 48 45 48 50 50 46 52
SAV 54 48 52 50 52 54 54 53 49 52
LUFF 14 12 – 14 – 9 9 – 11 12
LUFT 19 19 – 15 18 16 19 17 17 20
SC 4 5 5 6 5 4 5 5 6 6
SL (SO) 8 (6) 9 (7) 8 (6) 8 (6) 7 (5) 8 (6) 8 (6) 8 (6) 8 (6) 7 (6)
CP SPC SPC SPC C SPC SPC SPC C C SPC
CFP C C C C C C C C C C
CSN C C C C C C C C C C
CPF S S S S S S S S S S
KDS 5 5 5 5 5 5 5 5 5 5
Plantar
spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose
scales
Trachylepis wahlbergii (Peters, “1869” 1870)— provided the first records of the species for
Wahlberg’s Striped Skink Angola as “Euprepes punctatissimus” from
“Duque de Bragança,” “Biballa,” and “Caconda.”
Figures 102–104, plate 6
Additional material was later recorded as
Euprepes Wahlbergii Peters, “1869” 1870: 661. “Mabuya striata” by several authors (Bocage,
Lectotype: ZMB 6155 (collected by J.A. 1895, 1896, 1897b; Boulenger, 1905; Angel, 1923;
Wahlberg), designated by Boulenger (1887). Schmidt, 1933; Monard, 1937). Based on mor-
Type locality: “Damaraland,” Namibia. phological and coloration differences, Monard
(1937) described specimens from “Kuvangu” and
Euprepes punctatissimus: Bocage (1866a: 44,
“Bimbi” as “Mabuya striata angolensis” (fig. 102).
1870: 68, 1872: 80).
Specimens conforming to T. wahlbergii were also
Mabuia striata: Bocage (1895: 41, 1896: 111,
recorded by Laurent (1964) as “Mabuya striata
1897b: 211); Boulenger (1905: 111); Angel
chimbana” and “Mabuya angolensis.” Consider-
(1923: 160); Monard (1937: 88); Themido
ing that the specific epithet angolensis was preoc-
(1941: 8).
cupied by Bocage’s (1872) Euprepis angolensis,
Maubya striata: Schmidt (1933: 12); Parker Marques et al. (2018) provided a replacement
(1936: 136). name for Mabuya striata angolensis, Trachylepis
Mabuia striata angolensis: Monard (1937: 89); monardi. Branch and McCartney (1992) were the
Schätti and Perret (1997: 366). first to refer Angolan specimens to wahlbergii,
Mabuya striata striata: Manaças (1963: 235). and were followed by most authors (Branch,
Mabuya striata chimbana: Laurent (1964: 69). 1998; Broadley, 2000; Branch and Conradie,
Mabuya angolensis: Laurent (1964: 72). 2015; Conradie et al., 2016; Marques et al., 2018).
Mabuya striata angolensis: Mahnert (1976: 488); In a revision of the Trachylepis from southeastern
Ortiz (1989: 56). Africa, Broadley (2000) treated both wahlbergii
Mabuya striata wahlbergi: Branch and McCart- and striata as valid species, while Castiglia et al.
ney (1992: 1); Branch (1998: 157) (2006), based on molecular and karyological
Mabuya wahlbergii: Broadley (2000: 106). data, suggested these two taxa should be consid-
Trachylepis wahlbergi: Branch and Conradie ered conspecific. However, the taxonomic status
(2015: 200); Conradie et al. (2016: 26); Bap- and relationships within the striata complex
tista et al. (2019: 110); Branch et al. (2019a: remained unresolved, with some authors accept-
319). ing the presence of striata, wahlbergii, and
Trachylepis striata: Ceríaco et al. (2016c: 69); monardi in Angola (Branch et al., 2019a). In a
Branch et al. (2019a: 319). species-level phylogeny of the genus, Weinell et
Trachylepis wahlbergii: Marques et al. (2018: al. (2019) confirmed the validity of both T. stri-
270); Ceríaco et al. (2021: 115); Conradie et ata and T. wahlbergii, with Angolan populations
al. (2022: 205). assigned to the latter. Our data support this deci-
Trachylepis monardi: Marques et al. (2018: 265); sion, and previous records of striata from Angola
Branch et al. (2019a: 318); Ceríaco et al. should be assigned to this species, as well as
(2020a: 402). those recorded as T. angolensis and T. monardi.
Trachylepis chimbana: Ceríaco et al. (2020a: Although not based on Angolan specimens, Ste-
phens et al. (2022) showed that misidentification
402).
of this species in museums and collections is
This species is a member of the Trachylepis quite common.
striata species complex, a complex whose taxo- Diagnosis: A large-sized skink (max. SVL
nomic status and phylogenetic relationships are 85.8 mm, CAS 258394, max. SVL of extralimital
not yet fully resolved. Bocage (1866a, 1870, 1872) populations 107 mm, see Pietersen et al., 2021),
FIGURE 102. Syntype of Trachylepis monardi from Angola (presumably from Kuvangu or Bimbi, Cunene and
Huíla provinces, respectively) (MHNC 91.0610). Photos by L.M.P.C.
with fully developed pentadactyl limbs (figs. the anterior half of the body and sometimes
102–104); dorsal scales usually tricarinate; ven- completely absent. Middorsal scales sometimes
tral scales smooth; 53–63 SAV; 46–55 SAD; edged with black to form irregular longitudinal
35–40 MSR (32–43 in extralimital populations, lines. A black lateral stripe starts at the eye and
see Branch, 1998); lamellae beneath fingers and becomes fainter posteriorly, usually extending
toes keeled and spinose; plantar scales spinose; through the anterior half of the flank, rarely
15–18 LUFT; 12–15 LUFF; supranasals in con- reaching the hind-limb insertion. There may be
tact; parietals usually separated; prefrontals usu- black and pale spots irregularly scattered on the
ally separated; frontoparietals in contact; one dorsum, flanks, limbs, and tail. Top of head uni-
pair of enlarged nuchal scales present; ear open- formly colored or with irregular dark vermicula-
ing vertically ovoid and smaller than the eye; tion; labials dirty white without markings; gular
lacking subtriangular auricular scales on the region sometimes with irregular dark spotting.
anterior margin of the ear opening. Supralabials Breeding males acquire a vibrant yellow to
usually eight, sixth subocular; supraciliaries usu- orange coloration on the head and throat, some-
ally five, second largest; nostril oriented laterally. times extending to the neck and flanks. Ventrum
Dorsum grayish to olive brown; a pair of pale white to bluish white.
dorsolateral stripes starting above the eye and Material Examined: Bié Province: Chi-
becoming fainter posteriorly, usually restricted to tau [-11.4333°, 17.1500°, 1510 m] (CM 5863,
FIGURE 103. Specimen of Trachylepis wahlbergii Cangandala National Park, Malanje Province (CAS 258400).
Photos by L.M.P.C.
5873, 5879, 5882, 5887); Pavalange, Luando (CAS 258397); Cangandala National Park,
Natural Reserve [-10.9698°, 17.6124°, 1127 m] park headquarters [-9.8194°, 16.6539°, 1090
(ANSP 32190, 32197, 32198, 32200); 14 km m] (CAS 258394); Cangandala National Park
ENE from Mumbue village, near camp in small [-9.8178°, 16.6553°, 1119 m] (MUNHAC/
farm [-13.8577°, 17.4260°, 1569 m] (CAS MB03-001412); Cangandala National Park,
266043); camp established near Cassumbi vil- park guesthouse, around buildings [-9.8194,
lage, 14km dirt road, village near banana 16.6539, 1090 m] (MUNHAC/MB03-001413).
[-11.0839°, 16.6637°, 1242 m] (CAS 266027); Moxico Province. Cameia lake [-11.7167°,
camp surroundings, 14 km dirt road from Cas- 20.8000°, 1110 m] (IICT/R 23/1958, 26/1958,
sumbi village [-11.0832°, 16.6666°, 1241 m] 232/1959); Calombe [-11.8333°, 19.9333°, 1345
(MUNHAC/MB03-001414). Huíla Province: m] (IICT/R 61–67/1959, 253/1959, 256–
Jau [-15.2000°, 13.5200°, 1753 m] (MD 1890). 257/1959, 259/1959, 280/1959, 283/1959,
Malanje Province: Cangandala National Park, 326/1959, 335/1959, 405/1959, 426/1959);
vicinity of park headquarters [-9.8177°, Dilolo lake [-11.5000°, 22.0167°, 1088 m]
16.6546°, 10961 m] (CAS 258399); Cangandala (IICT/R 79–84/1959, 124–125/1958); Luena
National Park, vicinity of park headquarters [-11.7833°, 19.9167°, 1337 m] (IICT/R 234–
[-9.8181°, 16.6554°, 1091 m] (CAS 258395); 235/1959, 238/1959); Santa Cruz farm, Luena
Cangandala National Park, vicinity of park [-11.78333°, 19.9167°, 1337 m] (IICT/R
headquarters [-9.8194°, 16.6538°, 1090 m] 138/1958); environs du lac Calundo [-11.8000°,
FIGURE 104. Life photo of Trachylepis wahlbergii from 14 km dirt road from Cassumbi village, Bié Province
(MUNHAC/MB03-001414). Photo by L.M.P.C.
20.867°, 1119 m] (MD 5603, 5672, 5703, 5746, 13349, 13350, MHNC 91.0606*); Cubango River source
5752); Cazombo [-11.8800°, 22.9200°, 1110 m] site [-12.6605°, 16.089°, 1777 m] (PEM R23401). Huíla
(MD 5783); Calunda [-12.1200°, 23.4600°, Province: Kuvangu (= Kuvango) [-14.4667°, 16.3000°,
1473 m] (MD 5857). 1453 m] (NMB 13344–13347; MHNC 91.0561*,
Additional Material (* denotes type material): 91.0565*, 91.0599–91.0604*); Kalukembé [-13.7833°,
Bié Province: Silva Porto (currently Kuito) [-12.3975°, 14.6830°, 1699 m] (MHNC 91.0605); “Sangévé”
16.9377°, 1727 m] (MD 1806-5); Kuito [-12.3919°, [-13.8833°, 15.8333°, 1634 m] (MHNC 91.0546,
16.9387°, 1716 m] (IICT/R 77/1957, 81/1957); Chitau 91.0547); “Kapelongo” [-14.8833°, 15.0833°, 1192 m]
[-11.4333°, 17.1500°, 1510 m] (NMZB/UM 16356); (MHNC 91.0596); “Kambisa” [-15.3167°, 16.2167°,
HALO Cuito [-12.3958°, 16.9607°, 1700 m (PEM 1377 m] (MHNC 91.0554); “Kului” [-15.4167°,
R23363–6); Munhango village [-12.1607°, 18.5504°, 15.7333°, 1228 m] (MHNC 91.0562); “Mulondo”
1428 m] (PEM R23559). Cuando Cubango: 75 km [-15.6333°, 15.2000°, 1133 m] (MHNC 91.0555,
West of Mavinga [-15.6667°, 19.7000°, 1204 m] (PEM 91.0563). Lunda Sul Province: Alto Cuílo, Lunda
R4821); Cuando River Basin (site 43) [-17.5350°, [-10.0500°, 19.5170°, 1260 m] (MD 5307); Alto Chi-
23.1892°, 981 m] (PEM R20520‒1). Cunene Province: capa, Lunda [-10.9333°, 19.1500°, 1373 m] (MD 5402);
“Kuvelai” [-15.6500°, 15.8000°, 1217 m] (MHNC “Lunda” [-10.9667°, 20.0667°, 1238 m] (MHNC
91.0568); “Humbi” [-16.6833°, 14.9000°, 1105 m] 91.0551). Malanje Province: Cangandala National
(MHNC 91.0548, 91.0550, 91.0542); “Mupa” [-16.1833°, Park, vicinity of park headquarters [-9.8191°, 16.6547°,
15.7500°, 1166 m] (MHNC 91.0556–91.0558, 91.0560); 1094 m] (CAS 258400), [-9.8177°, 16.6546°, 1091 m]
“Mupanda” [-17.1333°, 15.7667°, 1114 m] (MHNC (CAS 258398), [-9.8181°, 16.6554°, 1091 m] (CAS
91.0559, 91.0570, 91.0597, 91.0598). Huambo Prov- 258396); Palavange [-11.3662°, 17.7216°, 1142 m]
ince: Bimbi [-11.8167°, 15.8333°, 1777 m] (NMB (USNM 85169, 85170). Moxico Povince: en route to
FIGURE 105. Distribution of Trachylepis wahlbergii in Angola. White star denotes the type locality of the
syntype of Trachylepis monardi.
Cuanavale River source [-12.6368°, 18.6598°, 1316 lake [-12.6886°, 18.3603°, 1426 m] (PEM R23513);
m] (PEM R23259); PEM R23289–95, Cuanavale Quembo River bridge camp [-13.5274°, 19.2806°,
River source [-13.0933°, 18.8940°, 1356 m] (PEM 1241 m] (PEM R27442); Luio River camp flood-
R23289–95); Cuito River source lake [-12.6894°, plains [-13.1971°, 20.2219°, 1181 m] (PEM
18.3601°, 1435 m] (PEM R23339–41); outlet of R27443; INBAC no number fide Conradie et al.,
Cuito River source lake [-12.7045°, 18.3545°, 1429 2022); Lake Hundo, trap 1 [-14.9916 °, 21.6310°,
m] (PEM R23376); Lungwebungu River camp 1100 m] (PEM R27444; INBAC/WC-6919). Unde-
bridge crossing [-12.5835°, 18.6660°, 1304 m] termined locality: Angola [undetermined locality]
(PEM R23412); Cuando River source [-13.0035°, (IICT/R Angola10–16); Angola [presumably Kuvangu
19.1275°, 1343 m] (PEM R23427; INBAC no or Bimbi] (MHNC 91.0607–91.0613*; MHNG 858.95*;
number fide Conradie et al., 2022); Cuando River MHNN 2130*); Angola [undetermined locality]
source trap 4 [-13.0016°, 19.1296°, 1372 m (PEM (MHNC 91.0542–91.0545, 91.0549, 91.0553, 91.0564,
R23484–5; INBAC/WC-4776); Cuito River source 91.0566, 91.0567, 91.0569, 91.0571–91.0595).
FIGURE 106. Typical habitat of Trachylepis wahlbergii in Cassumbi, Bié Province, Angola. Photo by L.M.P.C.
Historical Localities (no extant speci- Distribution in Angola: Trachylepis wahlber-

mens): Benguela Province: Hanha [-13.4941°, gii is widespread in most of the country, but appar-
14.5276°, 1669 m] (Bocage, 1896, 1897b). ently absent from the coastal lowlands (fig. 105).
Cuando Cubango Province: “region du Kwito, Global Distribution: Widespread in
affluent du Kubango” [-15.1667°, 19.1833°, Angola, eastward through Zambia to western
1185 m] (Angel, 1923). Cunene Province: Malawi, and southward through Namibia and
Humbe [-16.6833°, 14.9000°, 1105 m] (Bocage, Botswana to western South Africa (Masterson,
1895). Huambo Province: Galanga [-12.0667°, 2014c; Pietersen et al., 2021).
15.1500°, 1573 m] (Bocage, 1895). Huíla Prov- Habitat and Natural History Notes:
ince: Rio Cuce [-13.5167°, 15.2000°, 1752 m] This species occupies a variety of habitats,
(Bocage, 1895); Caconda [-13.7333°, 15.0667°, most notably miombo woodlands and savan-
1674 m] (Bocage, 1872); Huilla [-15.0500°, nahs (Grandvaux-Barbosa, 1970; fig. 106),
13.5000°, 1901 m] (Bocage, 1895; Angel, 1923); where it is frequently found basking on tree
Gambos [-15.7667°, 14.1000°, 1329 m] (Bocage, trunks or hiding under loose bark (Loveridge,
1895). Malanje Province: Duque de Bragança 1953; Broadley, 2000). Although mostly arbo-
[-9.1000°, 15.9500°, 1010 m] (Bocage, 1866a; real, it can also be found in rocky outcrops
1872; 1895). Namibe Province: Biballa (Laurent, 1964) and manmade structures,
[-14.7667°, 13.3667°, 955 m] (Bocage, 1872). such as buildings and bridges (Broadley,
Undetermined locality: Rio Quando (Bocage, 2000). This species is known to be viviparous
1895); Between Benguella and Bihé (Boulenger, (Broadley, 2000; Weinell et al., 2019), how-
1905). ever, gravid females containing eggs have
been reported from Angola (Manaças, 1963). material of this species was collected during the
Manaças (1963) examined four sexually Vernay Angola Expedition in 1925 in Pico Aze-
mature females collected between August and vedo and 101 km east of Moçamedes (specimens
October, containing three to seven eggs, those in the American Museum of Natural History).
collected in October with fully developed Holotype: An unsexed adult (CAS 254930,
embryos. A female collected by Monard field number JVV 8681; fig. 107) collected at Pico
(MHNC 91.0581, date unknown) contained Azevedo [-15.5340°, 12.49197°, 372 m], by Luis
two fully developed embryos. Stomachal con- M.P. Ceríaco, Edward L. Stanley, Arianna Kuhn,
tents reported by Manaças (1963) included Jens V. Vindum, Sango de Sá, Suzana A. Bandeira,
mostly termites and beetles. and Hilária Valério, on 7 December 2013.
Paratypes: All specimens from Angola. 21
specimens: Two unsexed adults (CAS 254927,
Trachylepis wilsoni, sp. nov.—
254928) Pico Azevedo [-15.5340°, 12.4919°, 372
Wilson’s Wedge-Snouted Skink
m] same data as holotype; an unsexed adult
Figures 107–109, plate 6 (CAS 254751), collected at Iona National Park,
3.4 km SW (by air) of Espinheira, vicinity of
Euprepes acutilabris [part]: Bocage (1870: 68). “Lion Cave” [-16.8151°, 12.337°, 453 m] by Luis
Mabuia acutilabris [part]: Bocage (1895: 46). M.P. Ceríaco, Edward L. Stanley, Arianna Kuhn,
Mabuya acutilabis [part]: Hellmich (1957b: 53), Jens V. Vindum, Sango de Sá, Suzana A. Ban-
Branch (1998: 151). deira, and Hilária Valério, on 30 November 2013;
Trachylepis acutilabris [part]: Haacke (2008: 90), two unsexed adults (CAS 254899, 254907)
Ceríaco et al. (2016b: 34, 57); Marques et al. Namibe-Lubango road, marker 59, 1.8 km W of
(2018: 252); Branch et al. (2019a: 318); Caraculo, N side of road [-15.0165°, 12.643°, 487
Ceríaco et al. (2020a: 402); Lobón-Rovira et m] by Luis M.P. Ceríaco, Edward L. Stanley, Ari-
al. (2022: 308). anna Kuhn, Jens V. Vindum, Sango de Sá, Suzana
A. Bandeira, and Hilária Valério, on 6 December
This new species has previously been mis- 2016; one unsexed adult (UF 187318) Virei camp
taken for T. acutilabris due to their remarkable [-16.1196°, 12.8346°, 522 m] by Luis M.P.
morphological similarities. Bocage (1870) was Ceríaco, Suzana A. Bandeira, and Ishan Agarwal,
the first to cite Mabuia acutilabris for Angola and on 29 November 2016; one unsexed adult (UF
was followed by several authors who reported 187317) Omauha-Chitundolo [-16.0006°,
specimens from most of the Angolan coastal 12.8381°, 583 m] by Luis M.P. Ceríaco, Suzana A.
regions (Peters, 1877; Bocage, 1895; Schmidt, Bandeira, and Ishan Agarwal, on 29 November
1919; Monard, 1937; Laurent, 1947, 1954, 1964; 2016; one unsexed adult (CAS 263487) Virei-
Hellmich, 1957a; 1957b; Haacke, 2008; Ceríaco Calundolo [-16.3102°, 12.7960°, 471 m] by Luis
et al., 2016b; Vaz Pinto et al., 2019). Recently col- M.P. Ceríaco, Suzana A. Bandeira, and Ishan
lected material allowed us to identify two diver- Agarwal, on 30 November 2016; two unsexed
gent lineages in Angolan populations, forming a adults (CAS 263492, UF 187298) Virulundo
clade sister to true T. acutilabris from Namibia. [-16.2852°, 12.9419°, 718 m] by Luis M.P.
Based on material recently collected in Namibe Ceríaco, Suzana A. Bandeira, and Ishan Agarwal,
Province, we here describe populations from on 2 December 2016; three unsexed adults (CAS
southwestern Angola previously assigned to T. 263494, 263495; UF 187297) Virei-Chipumpo
acutilabris as a new species, Trachylepis wilsoni, [-16.2793°, 12.9584°, 742 m] by Luis M.P.
sp. nov. Historical records of T. acutilabris from Ceríaco, Suzana A. Bandeira, and Ishan Agarwal,
Namibe Province (Bocage, 1870, 1895; Hellmich, on 1 December 2016; one adult female (MUN-
1957b) are referred to this species. Additional HAC/MB03-001415) Mucungu farm, rocky area
FIGURE 107. Holotype of Trachylepis wilsoni, sp. nov., from Pico Azevedo, Namibe Province (CAS 254930).
Photos by L.M.P.C.
[-14.7799°, 12.4878°, 305 m] by Mariana P. 12.4920°, 372 m] (CAS 254929, AMNH 48646, 48647,
Marques, Luis M.P. Ceríaco, and Joyce M. Janota, 48648); Espinheira [-16.7856°, 12.3589°, 454 m] (TM
on 2 August 2018; two adult females (MUN- 40628, 40629; PEM R18019, 18022, 18035, 18036);
Otchifengo River [-16.6167°, 12.8833°, 560 m] (TM
HAC/MB03-001416, 001417) Bentiaba river near
40774); Iona 6 km S of Rio Curoca [-16.3400°, 12.4400°,
Maungo [-14.5106°, 12.8391°, 417 m] by Mari- 287 m] (TM 40581–40589); 101 km east of Moçamedes
ana P. Marques, Luis M.P. Ceríaco, and Joyce M. [location not georeferenced] (AMNH 32799); Caraculo
Janota, on 11 August 2018; (MUNHAC/MB03- [-15.0165°, 12.6425°, 464 m] (AMB 13030–32, 13041–
001418–001420) base camp, in a dry river line at 43); Munhino [-14.9788°, 12.9781°, 415 m] (AMB
Maungo farm [-14.3934°, 12.8289°, 367 m] by 13050, 13052, 13053, 13062, 13063); Omauha
Mariana P. Marques, Luis M.P. Ceríaco, and [-16.1986°, 12.4007°, 337 m] (AMB 13226); Otchifengo
Joyce M. Janota, on 12 August 2018; one adult [-16.6849°, 12.8413°, 584 m] (AMB 13084, 13135,
13159, 13160, 13103–13); Vipungos [-15.0439°,
female (MUNHAC/MB03-001421) on the road
12.4234°, 333 m] (AMB 13018, 13019, 12993–95).
to Sangaia [-16.3382°, 12.2525°, 242 m] by Luis
Historical Localities (no extant speci-
M.P. Ceríaco on 13 November 2019; one adult
mens): Namibe Province: Rio Coroca [-15.7833°,
female (MUNHAC/MB03-001422) on Iona
12.0667°, 45 m] (Bocage, 1895: 46); Cahinde-
National Park [-16.6896°, 12.8128°, 623 m] by
Ougueiria [-15.4833°, 13.3667°, 631 m]
Luis M.P. Ceríaco on 15 November 2019.
Additional Material: Namibe Province: Deserto (Hellmich, 1957b: 53).
de Moçâmedes (MD 1945, 1946); Iona National Park, Diagnosis: A medium-sized skink (max. SVL
20 km SSW (by air) of Espinheira [-16.9316°, 12.2460°, 57.3 mm, CAS 254899), with fully developed,
615 m] (CAS 254789); Pico Azevedo [-15.5340°, pentadactyl limbs, and wedge-shaped snout (figs.
FIGURE 108. Life photo of the paratype of Trachylepis wilsoni from Iona National Park, Namibe Province
(MUNHAC/MB03-001422). Photo by L.M.P.C.
107, 108). Dorsal scales usually tricarinate or irregular, dark transverse bars that extend to the
quadricarinate; ventral scales smooth; 52–57 upper flanks. Limbs brownish above; hind limbs
SAV; 48–52 SAD; 29–33 MSR; lamellae beneath with dark-edged, pale circles posteriorly. A white
fingers and toes spinose; plantar scales spinose; lateral stripe may be evident, starting below the
22–26 LUFT; 15–17 LUFF; supranasals always in eye and extending to the hind limb insertion;
contact; parietals usually in contact; prefrontals lower flanks usually grayish. There is often heavy
always separated; frontoparietals always in con- grayish speckling, especially on top of the head,
tact; one pair of enlarged nuchal scales present; labials, and along the flanks. Venter white.
ear opening vertically ovoid and smaller than the Description of the Holotype: A well-pre-
eye; three to four subtriangular auricular scales served, unsexed adult. Body cylindrical with a
(shorter than the diameter of ear) extend poste- poorly defined neck and well-developed penta-
riorly and usually slightly upward from the ante- dactyl limbs; tail long, its length greater than the
rior margin of the ear opening. Supralabials SVL, smoothly tapering. Fore- and hind limbs
usually seven; subocular does not reach the lip; overlap when adpressed against the body. SVL
supraciliaries usually five, second largest; nostril 50.6 mm, TL 69.2 mm. HL 10.8 mm, with
oriented dorsally. Dorsum and upper flanks pale wedge-shaped snout. Additional measurements
to grayish brown, uniform or with pale longitu- are presented in table 10. Three subtriangular
dinal stripes; when present, dorsal stripes are auricular scales extend posteriorly from the ante-
often faint, especially vertebral one; there are rior margin of the ear opening. Rostral visible
usually scattered white spots on the back and from above. Nostrils oriented dorsally and set
TABLE 10
Mensural and Meristic Data for the Type Series of Trachylepis wilsoni, sp. nov. 2024
Abbreviations are listed in the Materials and Methods. Measurements are presented in millimeters and ratios as percentages.
CAS CAS CAS CAS CAS CAS CAS CAS CAS CAS UF UF UF UF MUH- MUH- MUH- MUH- MUH- MUHN MUH- MUH-
254930 254927 254928 254751 254899 254907 263487 263492 263494 263495 187317 187318 187297 187298 NAC/ NAC/ NAC/ NAC/ NAC/ AC/ NAC/ NAC/
MB03- MB03- MB03- MB03- MB03- MB03- MB03- MB03-
001415 001416 001417 001418 001419 001420 001421 001422
Holo- Paratype Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para- Para-
type type type type type type type type type type type type type type type type type type type type type
Sex ♀ unsexed unsexed unsexed unsexed unsexed unsexed unsexed unsexed unsexed unsexed unsexed unsexed unsexed ♀ ♀ ♀ ♀ ♀ ♀ ♀ ♀
SVL 50.6 51.8 56.3 51.9 57.3 53.8 56.3 56.8 50.5 44.8 46 54.6 49.6 51.7 42.4 49.1 40.5 40.8 40 43.2 46.2 38.6
TL 69.2 60.3 61.2 – 72.9 83.6 87 63 83 81.2 69 – 67.7 83 69 – – 73.4 – 81.5 91.1 –
TL/ 140 120 110 – 130 150 150 110 160 180 150 – 160 160 160 – – 180 – 190 200 –
SVL
HL 10.8 11.4 11.7 12 11.8 12.6 12.1 12.2 10.9 10.5 10 11.9 12.5 11.2 9.6 10.9 9 10.3 9.3 9.5 10.5 8.2
HL/ 20 20 20 20 20 20 20 20 20 20 20 20 30 20 20 20 20 30 20 20 20 20
SVL
SVL/ 437 450 480 430 480 430 470 470 460 430 460 450 400 460 440 450 450 400 430 450 440 470
HL
HW 7.8 8.1 8.2 7.8 8.2 8.4 8.7 8 7.2 6.7 7,1 8 7.6 7.7 6.5 7.4 6.7 6 6.4 6.5 6.8 6.2
HW/ 70 70 70 70 70 70 70 70 70 60 70 70 60 70 70 70 70 60 70 70 60 80
HL
HH 5.6 5.5 5.9 5.6 5.9 5.9 6.1 5.5 5 4.9 5,1 5.7 5.1 5.6 4.6 6 5.1 4.4 4.6 4.5 4.5 4.3
IN 2 2 1.8 1.9 2 1.7 2.2 1.9 2 1.7 1,7 1.9 1.6 1.6 2.3 2.5 1.2 1.3 1.8 1.7 2.3 1.8
EN 3.5 3.5 3.5 3.7 3.5 3.8 3.7 4.8 3.5 3.5 3 3.4 3.4 3.2 2.8 3.6 2.2 3.3 3 2.9 3.4 2.5
ES 4.9 5 5.2 5.3 5.3 5.1 5.3 5.2 4.8 5.1 4.3 4.9 4.8 4.7 4.1 4.6 3.5 4.1 3.9 4.3 4.6 4
MSR 31 29 30 30 31 29 30 33 29 32 30 30 30 31 31 34 32 34 32 21 30 33
SAD 51 50 48 52 50 50 50 50 51 51 49 50 49 49 48 48 48 53 49 48 50 44
SAV 56 53 55 55 57 54 54 53 55 54 54 52 54 55 52 52 54 56 52 55 52 45
LUFF 16 16 17 16 15 16 16 16 16 17 17 15 16 16 17 16 14 14 14 15 15 14
LUFT 26 23 23 24 22 23 25 23 23 26 25 24 25 25 22 22 22 22 22 22 26 23
SC 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5
SL 8 (6) 8 (6 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6) 8 (6)
(SO)
CP C C C SPC C S C SPC C C C SPC C C C C C C C C C C
CFP C C C C C C C C C C C C C C C C C C C C C C
CSN C C C C C C C C C C C C C C C C C C C C C C
CPF S S S S S S S S S S S S S S S S S S S S S S
KDS 3–4 3–4 3–4 3–4 3–4 3–4 3 2–3 3 3–4 3 2–3 3–4 3 3–4 3–4 3–4 3–4 3–4 3–4 3–4 3–4
Plantar spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose spinose
scales
135
posteriorly so that postnasal effectively borders

nostril. Supranasals in single point contact. Fron-
tonasal broader than long, in contact with ante-
rior loreal scale. Prefrontals subquadrangular,
without contact, each in contact with the follow-
ing head shields: frontonasal, loreals, first supra-
ocular and frontal. Two loreals. Frontal tapering
posteriorly, longer than the distance between
anterior tip of frontal and tip of snout. Frontal in
contact with three supraoculars on each side.
Two frontoparietals, in contact with each other
and the frontal, third and fourth supraoculars,
parietal, and interparietal. Frontoparietal plus
interparietal length smaller than frontal length.
Interparietal twice as long as broad, with a visible
parietal foramen. Parietals greater than frontopa-
rietals. Parietals in contact in a single point with
each other. Five supraciliaries, second largest.
Seven supralabials (six on left side); subocular
elongated, above fifth supralabial, not reaching
the lip. Six infralabials. Postmental bordering
seven scales (mental, two infralabials on each
side and two primary chin shields). Transparent
scale present in lower eyelid, as is usual for Tra-
chylepis. Tympanum visible, at same level as
mouth. Dorsal scales each with three to four
smooth keels. Ventral scales smooth. MSR 31,
SAD 51, SAV 56. Limbs with five digits; scales on
palms and soles spinose. Relative length of fin-
gers IV > III > V > II > I, relative length of toes
IV > III > V > II > I. Finger-IV lamellae 16, Toe- FIGURE 109. Details of head morphology of Trachy-
IV lamellae 26. Color in life is homogenously lepis wilsoni. Drawings by A.T.
light brown on the flanks, upper side of head,
neck, dorsum, legs, and tail brown with three with light grayish speckling. Ventral surfaces
pale dorsal stripes; vertebral stripe starts at the uniformly whitish with some scattered dark
nape and extends to the base of the tail; a pair of speckles near flanks, under tail, hands and feet.
dorsolateral stripes start behind the eye and Coloration in Ethanol: Same as in life (see
extend to the base of the tail. Scattered pale spots above).
and irregular black spots between dorsal stripes; Variation: Variation in mensural and meris-
the latter form transverse bands extending to the tic characters among the type series is presented
flanks. Hind limbs covered above by black- in table 10.
edged, pale circles or irregular shapes. A white Comparison with Other Angolan and
lateral stripe starts below the eye and extends to Southwest African Trachylepis: T. wilsoni dif-
the hind limb insertion; lower flanks white, with fers from all other species of Trachylepis known
grayish speckling. Top of head uniformly brown; to occur in Angola, with the exception of T. suza-
labials, subocular and lower half of loreals white, nae, by having a wedge-shaped snout. It differs
FIGURE 110. Distribution of Trachylepis wilsoni in Angola. Black star denotes the type locality.
from its sister taxon, T. suzanae, in having a establish the distribution limits of T. wilsoni and
more round, less acuminate head and more true T. acutilabris, and the putative role of the
prominent canthus rostralis (fig. 109). The new Cunene River as a barrier between these taxa.
species differs from T. acutilabris from Namibia, Habitat and Natural History Notes: This
by having 29–32 midbody scale rows (vs. 32–39 species is usually found in mopane woodlands
in T. acutilabris). and mosaic of steppe and savannah (Grandvaux-
Distribution in Angola: This species is Barbosa, 1970; fig. 111). It is associated with sandy
known only from inland areas of central and substrates, where it digs burrows near the base of
southern Namibe Province (fig. 110). vegetation (Ceríaco et al., 2016b).
Global Distribution: Confirmed records of Etymology: The specific epithet “wilsoni” is
this species are restricted to southwestern Angola. masculine in the genitive singular. It is given in
Further sampling in northern Namibia and a honor of Edward O. Wilson (1929–2021), North
review of Namibian species is needed to better American biologist, naturalist and author, in rec-
FIGURE 111. Typical habitat of Trachylepis wilsoni in Pico Azevedo, Namibe Province. Photo by L.M.P.C.
ognition of the importance and influence of his agreed that the country presents two main zoo-
work and legacy to all field biologists, taxono- geographic areas—one in the northern regions of
mists, and naturalists. We suggest “Wilson’s the country, coming south through the escarp-
Wedge-snouted Skink” and “Lagartixa de Edward ment with strong links to the Congolese/Central
Wilson” as the English and Portuguese common African fauna, and another one in the eastern
names, respectively, for this species. and southern regions, that has strong biogeo-
graphic links to the southern African and Zam-
bezian faunas (Monard, 1937; Frade, 1963;
DISCUSSION
Crawford-Cabral, 1991, unpubl. report; Marques
The considerable taxonomic diversity of Tra- et al., 2018).
chylepis occurring in Angola, combined with its While a biogreographical study of Angola is
almost complete geographic coverage of the being prepared for publication in the near future
country’s territory, makes the genus an interest- (Ceríaco et al., in prep.), the distribution of the
ing proxy to investigate the main biogeographic main lineages in the genus serves as a very inter-
patterns present in the region. The complexity of esting example of this north-south division. Fol-
the biogeography of Angola has been the subject lowing the whole-genus phylogeny of Weinell et
of many studies (Monard, 1937; Frade, 1963; al. (2019), Angola has representatives of two of
Crawford-Cabral, 1991, unpubl. report; Romei- the three main phylogenetic groups (sensu Wei-
ras et al., 2014; Rodrigues et al., 2015; Marques nell et al. (2019)) comprising the genus—Group
et al., 2018; Krásová et al.; 2021). B, mostly representing Central/Western African
As outlined as early as the 19th century by taxa, and Group C, mostly representing southern
Bocage (1895), Angola presents a north-south African taxa.
biogeographic division. Other authors deepened Within Group B sensu Weinell et al. (2019),
and refined this division, but as a whole they all Angola has representatives of the T. maculilabris
group (sensu Weinell et al., 2019)—T. maculila- example, the T. depressa group, represented in
bris and T. notabilis—and from the T. affinis Angola by two new species (formerly consid-
group (sensu Weinell et al., 2019)—T. albigularis. ered as part of T. acutilabris), T. wilsoni and
While Weinell et al.’s (2019) Group B is com- T. suzanae, contains sandy substrate adapted
posed of groups that are not exclusively Central/ specialists whose distribution in Angola fol-
West African (e.g., the Seychelles group endemic lows the coastal areas from Namibe Province
to the Seychelles archipelago and the T. quinqu- northward to Zaire Province and adjoining
etaeniata group, which contains T. margaritifera, Democratic Republic of Congo, reaching the
a species endemic to southeastern Africa), both northern distributional limit of the whole T.
the affinis and maculilabris groups are exclusively depressa group. The T. sulcata group, contain-
composed of species that have their distribution ing Namib specialists, occurs in southwestern
range in Central and West Africa. These three Angola (T. sulcata and T. laevis), reaching
species, T. affinis, T. maculilabris, and T. notabi- central Angola (with T. ansorgii), while T.
lis, can therefore be viewed as representatives of binotata, T. occidentalis, and T. hoeschi are all
the Central/West African fauna that finds its southwestern African species that also occur
southern distribution limit in northern Angola in Angola. The most diverse group represented
and following the escarpment north to south. in Angola, however, is the T. variegata group,
This distribution pattern agrees with the distri- containing species that occur both in more
bution of other co-occurring Central/West Afri- central areas of Angola—such as T. bayonii
can reptiles in the country, such as Panaspis or T. bocagii—and in the southern areas of
cabindae and Panaspis breviceps (Ceríaco et al., the country. The latter contain a large diver-
2018b), Bitis heraldica (Ceríaco et al., 2020c), sity, including four new species—T. hilariae,
Hemidactylus nzingae (Ceríaco et al., 2020b), T. ovahelelo, T. vunongue, and T. bouri. These
etc., as well as rodents (Krásová et al., 2021). observations sufficiently demonstrate the need
Within Group C sensu Weinell et al. (2019), to pursue a more detailed revision of the bio-
Angola has representatives of all but the Mala- geographic patterns of the genus in Angola.
gasy group, which is endemic to Madagascar. There is also the possibility that taxa occurring
Angolan representatives of the T. varia group— in neighboring countries, but not yet recorded
T. albopunctata, T. raymondlaurenti, and T. dam- in Angola, can be recorded in the country in
arana—as well as of the T. striata subgroup—T. the future, augmenting the current numbers
wahlbergii and T. attenboroughi—present a more and providing additional data to such biogeo-
Zambezian distribution, with the majority of the graphic analysis. Taxa such as T. variegata and
group being restricted to the Zambezi region, and T. cf. triebneri have been recorded near the
the Angola representatives distributed mostly in Angolan border at the Cunene River mouth
the central and eastern highlands of the coun- region in Namibia (Broadley, 1974b), while
try. This pattern is similar to that described by Pietersen et al. (2021) reported the presence of
Krásová et al. (2021) for rodent taxa with affini- an unknown Trachylepis species—T. sp. “Cen-
ties to the Zambezian region. tral Africa”—in northwestern Zambia, right at
The remaining groups contained within the border with Angola.
Group C sensu Weinell et al. (2019) have their Regarding the conservation status of Trachyl-
distribution in southern Africa, in an area epis species, all species occurring in Angola that
that roughly extends from southern Angola, have previously been assessed by the IUCN have
Namibia, South Africa, southern Botswana, a conservation status of Least Concern. This is
and Zimbabwe to Mozambique. Still under this mostly due to large ranges of occurrence of all
“southern African” sensu lato description, dif- these species, which, in almost all cases, fall
ferent groups have particular distributions. For within at least one conservation area, combined
with presumed large populations, based on their ACKNOWLEDGMENTS

ease of detection. The same conservation cate-
gory is recommended for the newly described The present work is a result of the ongoing col-
species, all of which have large distributions that laboration between the Instituto Nacional de Biodi-
occur in at least one conservation area. Some versidade e Áreas de Conservação (INBAC) from
species are rarer than others, however, this is the Ministry of Environment of Angola and its
most likely a sampling artifact. In the case of T. international partners. Angolan specimens were
notabilis, numbers and records may be underes- collected and exported under permits issued by
timated due to its common confusion with T. INBAC (155/INBAC.MINAMB/2017; 28/INBAC.
maculilabris. MINAMB/2019). We also thank the provincial and
With this study, we deepen our understanding local authorities for their support and cooperation
of the diversity and taxonomy of the genus Trachy- during our fieldwork. We thank Adam Fergunson,
lepis in Angola. While it is likely that more taxa will Amanda Picelli, António Martins, Arianna Kuhn,
be recognized within the country (even potentially Ben Marks, Calum Devaney, David C. Blackburn,
undescribed species), the current contribution rep- David Elizalde, Edward L. Stanley, Francisco Maiato
resents an important step not only to support the Gonçalves, Greg Jongsma, Hilária Valério, Ilola
ongoing revision of the country’s vast biodiversity, Jorge, Ishan Agarwal, Jens V. Vindum, John Cavagn-
but also to better understand the region as both a aro, Jorge Paiva, Joyce Janota, Luis Querido, Mat-
hotspot and cradle of herpetological diversity. Simi- thew P. Heinicke, Natasha Stepanova, Pedro
lar revisions are due for Trachylepis taxa occurring Baptista, Philip Pastor, Rogério Ferreira (RIP),
in the neighboring countries, such as the Demo- Sango de Sá, Sara Elizalde, Stuart Nielsen, Suzana
cratic Republic of Congo, Zambia, and Namibia, Bandeira and Timóteo Júlio for their support during
and will benefit from this study as a base for com- fieldwork and collecting valuable material. Stefan
parisons and data. Van Wyk is kindly acknowledged for his hospitality
Angola is a key to solving many taxonomic at the Cuatir Private Reserve. Special thanks to
issues surrounding several of the recognized Álvaro (“Varito”) Baptista and his team from
groups within Trachylepis. This is mostly due to Omauha Lodge, for all the assistance, great support,
the fact that many of the early taxa descriptions and friendship during the fieldwork. A special
were based on Angolan material, which has been thanks is also owed to António “Muloge” Lopes,
lacking from modern revisions, leading many António Francisco dos Santos, and Victor “Paka” for
authors to make assumptions regarding the distri- their help, dedication, and enthusiasm with field-
bution ranges and taxonomic identity of extralim- work. As in any major taxonomic revision, the
ital populations. Some of the extralimital access to natural history collections is critical.
taxonomic problems in the genus, such as the sta- Therefore, the authors want to present a special
tus and distribution of T. triebneri, are also acknowledgment to the collection managers and
informed by Angolan populations. In this case, curators who provided access and data from the col-
clarifying the status of Angolan members of the T. lections under their care: David Kizirian and Lauren
variegata group provides a better standpoint from Vonnahme (AMNH); Ned Gilmore (ANSP); Pat-
which to start dealing with the Namibian popula- rick Campbell and Jeffrey Streicher (BMNH); Lau-
tions, which historically have been considered as ren Scheinberg and Erica Ely (CAS) ; Steve Rogers
conspecific with those from Angola, an interpreta- (CM); Alan Resetar and Joshua Mata (FMNH); José
tion shown herein as not always warranted. Rosado (MCZ); Ilunga André (MD); Arnaud
Further studies are needed to clarify remaining Maeder (MHNC); Andreas Schmitz (MHNG);
questions surrounding some taxa within the genus, Belmira Gumbe and Ana Lavres (MNHNL);
namely more complete distribution areas, ecologi- Annemarie Ohler and Roger Bour (MNHN); Raf-
cal preferences, and basic natural history. fael Ernst (MTD); Silke Schweiger (NHMW); Moira
FitzPatrick and Shiela Broadley (NHZB); Michael F. Allen, K.E., W.P. Tapondjou, E. Greenbaum, L.J. Welton,
Bates (NMB); William R. Branch and Werner Con- and A.M. Bauer. 2019. High levels of hidden phyloge-
radie (PEM); Uwe Fritz (SMF); Adriaan Jordaan netic structure within Central and West African Tra-
chylepis skinks. Salamandra 55 (4): 231–241.
and Moleboheng Mohapi (TM); Coleman Sheehy
Angel, M.F. 1923. Reptiles. In J. Rohan-Chabot, (editor),
III (UF); Esther Langan (USNM); Mark-Oliver
Mission Rohan-Chabot, Angola et Rhodesia (1912–
Rödel and Frank Tillack (ZMB); Jakob Hallermann 1914), Tome IV, Histoire Naturelle, Fascicule 1 (Mam-
(ZMH); and Frank Glaw and Michael Franzen mifères—Oiseaux—Reptiles—Poissons): 157–169, 1
(ZSM). William R. Branch, Luis Querido, Ishan pl. Paris: Imprimerie Nationale.
Agarwal, and Werner Conradie provided photos of Auerbach, R. 1986. First steps in Setswana herpetology.
specimens and habitats and are acknowledged here. Botswana Notes and Records 18: 71–90.
The authors want to thank Mary Knight, managing Baptista, N., T. António, and W.R. Branch. 2018. Amphib-
editor of the Bulletin of the American Museum of ians and reptiles of the Tundavala region of the Ango-
lan escarpment. Biodiversity and Ecology 6: 397–403.
Natural History, for her editorial work, which
Baptista, N.L., T. António, and W.R. Branch. 2019. The
greatly improved this manuscript. Miguel Trefaut herpetofauna of Bicuar National Park and sur-
Rodrigues and Olivier Pauwels provided critical and roundings, southwestern Angola: a preliminary
important comments during the revision process, checklist. Amphibian and Reptile Conservation 13
and are also acknowledged here. (2): 96–130.
This work was funded by U.S. National Science Bauer, A.M., W.R. Branch, and W.D. Haacke. 1993. The
Foundation grants (DEB 1556255, 1556585, herpetofauna of the Kamanjab area and adjacent
1556559, 1657527, 2146654); a grant from JRS Bio- Damaraland, Namibia. Madoqua 2: 117–145.
diversity Foundation to A.M.B., Matthew P. Bauer, A.M., G. Shea, and R. Günther. 2003. An annotated
catalogue of the types of scincid lizards (Reptilia, Squa-
Heinicke, and David C. Blackburn. L.M.P.C. was
mata, Scincidae) in the collections of the Museum für
supported by the National Geographic Society Naturkunde der Humboldt-Universität zu Berlin
Explorer Grant (NGS-73084R-20); D.P. is supported (ZMB). Mitteilungen aus dem Museum für
by Fundação para a Ciência e Tecnologia (FCT) Naturkunde zu Berlin 79 (2): 253–321.
grant (2021.05238.BD); and M.P.M. was supported Bocage, J.V.B. 1866a. Lista dos reptis das possessões Por-
by FCT grants (SFRH/BD/129924/2017, COVID/ tuguezas d’Africa ocidental que existem no Museu de
BD/152155/2022). Work was cofunded by the proj- Lisboa. Jornal de Sciencias Mathematicas, Physicas e
ect NORTE-01-0246-FEDER-000063, supported by Naturaes 1: 37–56.
Norte Portugal Regional Operational Programme Bocage, J.V.B. 1866b. Reptiles nouveaux ou peu connus
recueillis dans les possessions portugaises de l’Afrique
(NORTE2020), under the PORTUGAL 2020 Part-
occidentale, qui se trouvent au Museum de Lisbonne.
nership Agreement, through the European Regional Jornal de Sciencias Mathematicas, Physicas e Naturaes
Development Fund (ERDF). The late William (Bill) 1: 57–78.
R. Branch, who collaborated during the early stages Bocage, J.V.B. 1867a. Segunda lista dos reptis das posses-
of this project, is fondly remembered here. sões portuguezas d’Africa ocidental que existem no
Museu de Lisboa. Jornal de Sciencias Mathematicas,
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PLATES
PLATE 1. 1. Trachylepis albilabris; 2. T. albopunctata; 3. T. ansorgii. Drawings by A.T.

PLATE 2. 4. Trachylepis attenboroughi, sp. nov.; 5. T. bayonii; 6. T. binotata; 7. T. bocagii. Drawings by A.T.
PLATE 3. 8. Trachylepis bouri, sp. nov.; 9, T. chimbana; 10. T. damarana; 11. T. hoeschi; 12. T. huilensis; 13. T.
hilariae, sp. nov.; 14. T. laevis. Drawings by A.T.
PLATE 4. 15. Trachylepis maculilabris; 16. T. notabilis. Drawings by A.T.

PLATE 5. 17. Trachylepis occidentalis; 18. T. ovahelelo, sp. nov.; 19. T. suzanae, sp. nov.; 20. T. punctulata; 21.
T. raymondlaurenti. Drawings by A.T.
PLATE 6. 22. T. vunongue, sp. nov.; 23. T. sulcata; 24. T. wahlbergi; 25. T. wilsoni, sp. nov. Drawings by A.T.

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THE TRACHYLEPIS (SQUAMATA: SCINCIDAE)

OF ANGOLA: AN INTEGRATIVE TAXONOMIC REVIEW

LUIS M.P. CERÍACO

Copyright © American Museum of Natural History 2024 ISSN 0003-0090

Trachylepis of Angola. We provide a phylogeny of Natural, Luanda, Angola; MNHN-RA, Muséum

Taxon 16S ND2 RAG1

Trachylepis gravenhorstii 1 MAB 182 Madagascar: Tulear: Ifaty: Ranobe Village

Trachylepis spilogaster MB 20874 South Africa: Northern Cape: Farm Boseekoebaard

In our ML phylogeny (fig. 1), most lineages Species Concept

SC 6–7 5–6 5 5 4–5 5–6 4–6

EN 3.5–4.6 2.4–3.7 3.4–6.8 2.4–2.7 4.9–5.6 2.5–3.9 2.4–3.9

HW 6.1–15.0 7.6–16.3 7.7–17.7 5.8 4.5–7.0 5.9–8.6 6.6–14.7

EN 2.3–5.2 3.0–6.5 3.6–6.2 2.5 2.5–3.4 2.5–3.7 2.6–6.1

CSN Usually C (N = 7) or Usually C (N = 4) Always C C Always C Usually S (N = 16), Usually C, rarely S

CSN Usually C C Always C Always C

FIGURE 4. Distribution of Trachylepis albilabris in Angola.

Trachylepis albopunctata (Bocage, 1867)— National Park, Malanje Province, central

Trachylepis Trachylepis Trachylepis Trachylepis Trachylepis

Sex unsexed unsexed unsexed unsexed unsexed

Material Examined: Bié Province: Chitau 13.2569°, 1585 m] (MUNHAC/MB03-001468,

1890 m] (MUNHAC/MB03-001385). Malanje (=N’dalatando) [-9.3000°, 14.9167°, 782 m] (BMHN

Duque de Bragança (currently Kalandula) [-9.1000°, [-9.6833°, 14.4333°, 33 m] (Bocage, 1872,

CAS 263534 CAS 254903 CAS 85961

lid, as is usual for Trachylepis. Tympanum visible, Coloration in Ethanol: Background

Trachylepis damarana (Peters, 1870)— Cuando Cubango province as T. varia, but

FIGURE 44. Distribution map of Trachylepis damarana in Angola.

CAS 254775 CAS 254769 CAS 254770 CAS 254771

Habitat and Natural History Notes: Trachylepis hoeschi (Mertens, 1954)—

FIGURE 51. Distribution of Trachylepis hoeschi in Angola.

Figures 53, 54, plate 3 In the original description of Euprepes Bayo-

16.66593°, 1223 m] (MUHNAC/MB03- Trachylepis occidentalis (Peters, 1867)—

FIGURE 74. Distribution of Trachylepis occidentalis in Angola.

TABLE 7 series of transversely aligned dark speckles;

Distribution in Angola: This species is Trachylepis punctulata (Bocage, 1872)—

Trachylepis sulcata (Peters, 1867)— Euprepes sulcatus Peters, 1867: 20 [replacement

FIGURE 91. Distribution map of Trachylepis sulcata in Angola.

entrance [-14.8443°, 15.0356°, 1220 m] savannah-steppe mosaic (Grandvaux-Barbosa,

Expedition in 1925, are deposited in the AMNH

aSeparated by small internasal scale.

FIGURE 100. Distribution of Trachylepis vunongue in Angola.

Historical Localities (no extant speci- Distribution in Angola: Trachylepis wahlber-

posteriorly so that postnasal effectively borders

with presumed large populations, based on their ACKNOWLEDGMENTS

PLATE 1. 1. Trachylepis albilabris; 2. T. albopunctata; 3. T. ansorgii. Drawings by A.T.

PLATE 4. 15. Trachylepis maculilabris; 16. T. notabilis. Drawings by A.T.

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