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Herpetological Survey of Iona National Park and Namibe Regional Natural Park, with a Synoptic List of the Amphibians and Reptiles of Namibe Province, Southwestern Angola

Authors:
Herpetological Survey of Iona National Park and Namibe
Regional Natural Park, with a Synoptic List of the Amphibians
and Reptiles of Namibe Province, Southwestern Angola
Luis M. P. Ceríaco 1,2,8, Sango dos Anjos Carlos de Sá 3, Suzana Bandeira 3, Hilária Valério 3,
Edward L. Stanley 2, Arianna L. Kuhn 4,5, Mariana P. Marques 1, Jens V. Vindum 6,
David C. Blackburn 2, and Aaron M. Bauer 7
1Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Rua da Escola Politécnica,
58, 1269-102 Lisbon, Portugal. 2Department of Natural History, Florida Museum of Natural History,
Gainesville, FL, 32611, USA. 3Instituto Nacional da Biodiversidade e Áreas de Conservação, Ministério
do Ambiente de Angola, Centralidade do Kilamba, Rua 26 de Fevereiro, quarteirão Nimi ya Lukemi,
edíficio Q11, 3° andar, Angola. 4American Museum of Natural History, Central Park West at 79th Street,
New York, New York 10024, USA. 5 City University of New York, Graduate Center, 365 5th Ave.,
New York, New York, 10016, USA. 6 Department of Herpetology, California Academy of Sciences,
55 Music Concourse Drive, San Francisco, California 94118, USA. 7 Department of Biology,
Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1699, USA.
8Corresponding author: Luis M. P. Ceríaco, Email address: luisceriaco@netcabo.pt
Namibe Province is the southernmost province of Angola and, as the result of sever-
al expeditions from the nineteenth century to the present, it is one of the most her-
petofaunally well-known areas of the country. The Province harbors a high diversi-
ty of amphibians and reptiles, including roughly one-third of the reptile taxa report-
ed for Angola as a whole. In this paper we present the results of a joint herpetologi-
cal expedition to Namibe Province in 2013 by the California Academy of Sciences
and the Instituto Nacionalda Biodiversidade e Áreas de Conservação, as well as a
synoptic list of all the herpetological bibliographic records for the taxa known from
the Province. A total of 37 herpetological taxa was collected, including at least three
(then) undescribed species, two new country records, and new records for rarely
cited taxa in Angola. These taxa belong to four amphibian genera and 15 reptile gen-
era. Species accounts are provided for each of the species collected. We also highlight
biogeographic patterns, conservation issues, and possible future paths for the explo-
ration and knowledge of the herpetofauna of Namibe.
A província do Namibie situa-se no sudoeste de Angola e é uma das mais conhecidas
relativamente à sua herpetofauna. Este conhecimento é resultado de várias expe-
dições realizadas desde o século XIX até aos dias de hoje. A província alberga uma
espetacular diversidade de anfíbios e répteis, que para estes ultímos representa
aproximadamente a um terço dos taxa que ocorrem no país. Neste artigo apresenta-
mos os resultados da expedição herpetológica levada a cabo pela California Acade-
my of Sciences e o Instituto Nacional da Biodiversidade e Áreas de Conservação em
2013, bem como uma lista sinóptica de todos os registos bibliográficos para os taxa
conhecidos na província. Um total de 37 taxa de anfíbios e répteis foram colectados,
incluíndo pelo menos três espécies novas (uma já descrita e as outras em processo de
descrição), dois novos registos de espécies para o país, bem como o registo de espé-
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2, pp. 15–61, 19 figs., Appendix April 29, 2016
15
Reprinted frorm Proceedings of the California Academy of Sciences, ser. 4, vol. 63, pp. 15-61. © CAS 2016
cies raramente citadas para o país. Estes taxa pertencem a 4 géneros de anfíbios e a
15 géneros de répteis. Todos os resultados são apresentados em fichas taxonómicas.
São ainda apresentados algums comentários relativos a padrões biogeográficos e
questões ligadas à conservação, e futuros caminhos para a exploração e conheci-
mento da herpetofauna da província do Namibe.
KEYWORDS: amphibians, Angola, biogeography, conservation, geographic distribution,
Namibe Province, reptiles.
The current knowledge of
Angola’s herpetofauna is incom-
plete in contrast to neighboring
countries such as Namibia (Her-
rmann and Branch 2013, Mar-
ques 2015). Namibe is Angola’s
southwesternmost province and
is one of the better explored
provinces in terms of herpetolog-
ical diversity (Branch et al.
2014). Namibe Province occu-
pies an area of 57,097 km2and is
bordered by Huíla Province to
the northeast, Cunene Province
to the southeast, Benguela
Province to the north, Namibia to
the south, and the Atlantic Ocean
to the west. The province is geo-
graphically separated from Huíla
by the great escarpment of Serra
da Leba and Chela, which sepa-
rates the lower elevation areas of
the Namib Desert from the Huíla
Plateau. Topographically, the
majority of the province has an
elevation lower than 500 m, ris-
ing to 1500 m at the escarpment
in the east. The highest elevation
is at the Serra da Neve inselberg
(2403 m) in north of the
province, almost at the border
with Benguela. The province has two main conservation areas – Iona National Park (INP), the
largest conservation area in the country with an area 15,150 km2, and the smaller Namibe Region-
al Natural Park (NNP), with an area of 4,450 km2. Namibe lies within the African southwest arid
biome, mainly comprising Kaokoveld desert, Namibian savanna, miombo woodlands, and mopane
forest. The Kaokoveld desert, which extends along the coastal regions from southern Benguela
Province to the Skeleton Coast in Namibia, is mostly dominated by sandy dunes and the occasion-
al presence of Odyssea paucinervis, Sporobolus spicatus, and Acanthosicyos horridus dominated
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FIGURE 1. Map of the main sampled localities: A) Morro do Soba; B)
Omauha Lodge; C) Rio Curoca; D) Entrada do Iona; E) Campo das espin-
heiras; F) Iona; G) Ford car Wreck; H) Pediva; I) Tambor; J) Serra da Leba;
K) Pico do Azevedo; L) Caraculo; M) Praia do navio.
vegetation. The Namibian savanna woodlands in the central areas of the province are dominated
by herbaceous plants of the genera Aristida and Eragrostis, dispersed shrubs of Acacia, Com-
miphora and Combretum, and, towards the southwest, extensive populations of Welwitschia
mirabilis. The eastern portions of the province support smaller areas of Angolan mopane wood-
lands, which are dominated by the deciduous tree Colophospermum mopane, and Angolan miom-
bo woodlands, which are dominated by Brachystegia trees, but also typified by Isobertlinia
angolensis, Julbernardia paniculata and Baikiaea plurijuga (Grandvaux-Barbosa 1970). The
province is bounded in the south by the Cunene (Kunene) River, and crossed by the Curoca and
Giraul Rivers. Geologicaly, the south of the province is mostly dominated by schists, sometimes
interspersed with granites, while the north of the province is mainly comprises granites (Anony-
mous 1963).
Approximately 16 species of amphibians and 95 species of reptiles are known from Namibe
Province (Table 1). In separate works we are preparing a complete review of the diversity and dis-
tribution (including an annotated checklist) of the Angolan herpetofauna based published biblio-
graphic records prior to 2014 (Marques et al., in prep.), and an annotated checklist of the herpeto-
fauna of Namibe Province, based on bibliographic material, unpublished museum records, and
recent collections (Branch et al., in prep). This paper presents the results of an expedition con-
ducted by a team from the California Academy of Sciences (CAS), San Francisco (USA), Villano-
va University (VU), Villanova (USA), and the Instituto Nacional da Biodiversidade e Áreas de
Conservação (INBAC), Kilamba-Kiaxi (Angola). A total of 37 herpetological taxa were collected,
including at least three new species, one of which has recently been described (Stanley et al. 2016),
two new country records and new records for taxa rarely cited for the country. We then provide a
brief discussion of the present status and future prospects for the study of the herpetofauna of
Namibe.
HISTORY OF THE HERPETOLOGICAL EXPLORATION OF THE PROVINCE
Namibe Province was explored by several well documented expeditions in the nineteenth and
twentieth centuries. The first herpetological surveys conducted were those of the Portuguese
explorer José de Anchieta (1832–1897), who visited the region in the late nineteenth century.
Anchieta’s specimens were deposited in the Natural History Museum of Lisbon and largely stud-
ied by the Portuguese zoologist José Vicente Barbosa du Bocage (1823–1907), who published sev-
eral papers on this material (e.g., Bocage 1867, 1873, 1896). Based on material from Anchieta, as
well as others, Bocage described several herpetological taxa from Namibe Province, including
Anchieta’s Tree Frog, Leptopelis anchietae (Bocage, 1873), the Double-scaled Chameleon,
Chamaeleo anchietae Bocage, 1872, Anchieta’s Ground Agama, Agama anchietae Bocage, 1896,
Anchieta’s Shovel-snout Lizard, Meroles anchietae (Bocage, 1867), the Reticulate Sand Lizard,
Meroles reticulatus (Bocage, 1867), the Speckled Sand Skink, Trachylepis punctulata (Bocage,
1872), the Speckled Western Burrowing Skink, Typhlacontias punctatissimus (Bocage, 1873),
Anchieta’s Worm-Lizard, Monopeltis anchietae (Bocage, 1873), a Skaapsteker, Psammophylax
occelatus Bocage, 1873 (currently a synonym of the Spotted Skaapsteker, Psammophylax
rhombeatus (Linnaeus, 1758)), a new variety of Striped Sand Snake, Psammophis sibilans var.
stenocephalus (Bocage, 1877), and a new species of Giant Blind Snake, Onychocephalus petersii
Bocage, 1873 that is currently considered a synonym of Afrotyphlops schlegelii (Bianconi, 1847).
At the dawn of the twentieth century, the Portuguese explorer Francisco Newton (1864–1909)
collected herpetological specimens in the province for the Natural History Museum of the Poly-
technic University of Porto, during a three year mission from 1903 to 1905, exploring the provinces
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 17
of Kwanza-Norte, Kwanza-Sul and Namibe. The initial collections made by Newton in the two first
provinces were studied and published upon by the Portuguese zoologist José Júlio Bettencourt Fer-
reira (1866–1948) on two different occasions (Ferreira 1906, 1906), but the Namibe Province mate-
rial had remained unstudied until today (Ceríaco et al., in prep.). In 1925, the Vernay Angola Expe-
dition explored central and southwestern Angola and collected specimens destined for the Ameri-
can Museum of Natural History. The herpetological results of this expedition were partly published
upon by the American herpetologist Charles M. Bogert (1908–1992), in a paper dealing with the
snakes (Bogert 1940). A second paper detailing the rest of the herpetological material collected on
the expedition was never published. Some years later, two Swiss scientific expeditions to Angola,
1928–1929 and 1932–1933, led by the Swiss naturalist Albert Monard (1886–1952) also explored
several locations in Namibe Province. The herpetological results of these expeditions were pub-
lished in four different papers (Monard 1931, 1937a, b, 1938). In between the two Swiss expedi-
tions, the Pulitzer-Carnegie Museum Expedition to Angola in 1930, led by influential American
publisher Ralph Pulitzer (1879–1939) and conducted by Wilfrid Rudyerd Boulton (1901–1983)
and his wife Laura Crayton Boulton (1899–1980), became one of the most important expeditions
in terms of herpetological results. The material was studied and published upon by Karl Patterson
Schmidt (1890–1957) in two papers — one dedicated to the reptiles (Schmidt 1933) and other to
the amphibians (Schmidt 1936). These works resulted in the description of Pulitzer’s Thick-toed
Gecko, Chondrodactylus pulitzerae (Schmidt, 1933), Boulton’s Namib Day Gecko, Rhoptropus
boultoni Schmidt, 1933, and the Angolan endemic subspecies of White-Throated Monitor, Varanus
albigularis angolensis Schmidt, 1933. More recently the Belgian herpetologist Raymond Laurent
(1917–2005) published on a collection of amphibians and reptiles from Namibe (Laurent 1964)
sent to him by the Portuguese entomologist and director of the former Museu do Dundo (northeast
Angola), António Barros de Machado (1912–2002). This contribution was of uttmost importance
for the knowledge of the southwestern Angolan herpetofauna. In addition to the several new taxa
added to the list of the provincial herpetofauna, he described four new taxa endemic to the south-
west of the country — Bogert’s Speckled Western Burrowing Skink, Typhlacontias bogerti Lau-
rent, 1964, Hellmich’s Wolf Snake, Lycophidion hellmichi Laurent, 1964, and two Namib Day
Geckos, Rhoptropus taeniostictus Laurent, 1964 and R. boultoni montanus Laurent, 1964. Wulf
Haacke conducted the last systematic field surveys in the Portuguese colonial period in 1971 and
1974 and deposited his collections in the Ditsong National Museum of Natural History (TM) in
Pretoria, South Africa, though this material has not been fully published upon. After independence
in 1975, Angola entered a long period of civil war, which ended only in 2002. This prevented fur-
ther field surveys and studies. In the past decade, several field surveys have been conducted,
including in Namibe. Teams from the Porth Elizabeth Museum — Bayworld (PEM) prospected the
province in three different expeditions so far, and a team from CAS, INBAC and VU conducted
the survey reported in this paper. These expeditions have increased our knowledge of the southern
Angolan herpetofauna, and since 2008 five new taxa from southwestern Angola (2008–2013) have
been described — the Chela Mountain Reed Frog, Hyperolius chelaensis Conradie, Branch,
Measey and Tolley, 2012, the Namib Spiny Tailed Gecko, Afrogecko plumicaudus Haacke, 2008,
subsequently made the type species of the monotypic genus Kolekanos Heinicke, Daza, Green-
baum, Jackman and Bauer, 2014, Haacke’s Sand Lizard, Pedioplanis haackei Conradie, Measey,
Branch and Tolley, 2012, Huntley’s Sand Lizard, Pedioplanis huntleyi Conradie, Measey, Branch
and Tolley, 2012, and the Kaokoveld Girdled Lizard, Cordylus namakuiyus Stanley, Ceríaco, Ban-
deira, Valério, Bates and Branch, 2016 — all but the first endemic to Namibe Province.
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MATERIAL AND METHODS
We conducted herpetological surveys in Namibe Province from 28 November to 11 December
2013, including both Iona National Park and Namibe Regional Natural Park. A total of 13 areas
were surveyed (Fig. 1). In each area, we attempted to sample a combination of habitat types. Over-
all conditions during this fieldwork were hot and dry as this was an unusually dry year. We cap-
tured specimens using long-nooses, rubber bands, or by hand during both diurnal and nocturnal
visual surveys. All specimens were euthanized following an approved IACUC protocol (#2014-2),
preserved in 10% buffered formalin in the field, and then transferred to 70% ethanol for storage.
Liver tissues were preserved in 95% ethanol and RNALateR. Voucher specimens and tissue sam-
ples are deposited in the herpetological collection of the California Academy of Sciences, with a
subset of specimens deposited in the reference collection of INBAC. In some cases, we further con-
firmed species identifications by sequencing the mitochondrial 16S ribosomal RNA gene. As noted
above, a complete list of all amphibians and reptile species reported from Namibe Province was
assembled (Table 1). This list, including localities and associated bibliographic references was
based on the ongoing project for the first atlas of the Angolan amphibians and reptiles (Marques
2015; Marques et al., in prep.). We do not include in the list unpublished museum records such as
the large series in the Ditsong National Museum of Natural History collected by Wulf Haacke in
the 1970s or the recent collections made by William R. Branch and Werner Conradie from the Port
Elizabeth Museum (Bayworld). These specimens will be a part of a forthcoming publication (W.R.
Branch, pers. comm.). However, museum material representing taxa vouchered on our expedition
are noted when relevant in the species accounts.
RESULTS
A total of 411 specimens were collected during the expedition, representing four amphibian
genera and 15 reptile genera. In the following species accounts, we provide information on CAS
voucher specimens, localities, and natural history. Latitude, longitude and elevation (in meters) of
the collection site are provided in each species account. In addition, when appropriate, we provide
brief taxonomic or geographic notes.
SPECIES ACCOUNTS
Amphibia
Anura
Bufonidae
HALLOWELLSTOAD
Sclerophrys maculata (Hallowell, 1854)
MATERIAL.— Leba Pass, between river and highway, 5 December 2013, 15º04ʹ13.2ʺS,
13º14ʹ37.7ʺE, 1676 m (CAS 254877–254878).
COMMENTS.— In Angola this species mostly occurs in the southwestern provinces of Namibe,
Benguela, Bié, and Huíla (Marques 2015). The nearest records are in “Cainde” and “16 km W of
Vila Nova” (Poynton and Haacke 1993; Ruas 1996, 2002). It is widespread in arreas to the south,
including much of northern Namibia (du Preez and Carruthers 2009). Ohler and Dubois (2016)
recently presented evidence identifying the type species of Sclerophrys Tschudi, 1838 as referable
to Bufo rangeri Hewitt, 1935, thus making Sclerophrys the oldest available name for the clade of
African toads recently referred to as Amietophrynus Frost et al., 2006.
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 19
Pyxicephalidae
ANGOLA RIVER FROG
Amietia angolensis (Bocage, 1866)
MATERIAL.— Leba Pass, between river and highway, 5 December 2013, 15º4ʹ12.2ʺS,
13º14ʹ38.9ʺE, 1676 m (CAS 254876).
COMMENTS.— This specimen represents the first record for the species for the province,
although there are several records from the province of Huíla, in Boca de Humpata (Laurent 1964a;
Ruas 1996; Channing and Baptista 2013), and Huila (Bocage 1895; Themido 1941; Perret 1976;
Ruas 1996, 2002) less than 20 km east of Leba Pass. The species is widespread across the rest of
the country (Marques 2015) as well as in much of the more mesic areas of southern Africa. In
Namibia it occurs only where there are permanent rivers (du Preez and Carruthers 2009).
DAMARALAND SAND FROG
Tomopterna damarensis Dawood and Channing, 2002
MATERIAL.— Pediva Hot Springs, 2 December 2013, 16º17ʹ4.62ʺS, 12º33ʹ47.86ʺE, 241 m
(CAS 254855).
COMMENTS.— The specimen was collected by locals on the border of the largest pond at Pedi-
va Springs. The lower jaw is broken, but the specimen is otherwise in good condition. A dark pig-
mentation is visible along the jaw-line, which identifies the specimen as a male, as is common in
the genus. Comparing our specimen to the recently described Tomopterna damarensis Dawood and
Channing, 2002, from Damaraland, northwestern Namibia, it agrees with the smooth dorsum and
most important morphological characters. Comparison of 16S mtDNA sequence to the type speci-
men confirms the identification of this specimen as T. damarensis (GenBank KU662310; p-dis-
tance from GenBank AY255091.1, the holotype of T. damarensis, is 0.7 %). Additional details on
the distribution of the species in Angola and Namibia are being prepared for publication
(M. Heinicke et al., in prep.). This is the first record of the species for the country, extending the
range of the species considerably northwards from the type locality at Khorixas, Namibia (Dawood
and Channing 2002).
Microhylidae
MARBLED RUBBER FROG – Fig. 2
Phrynomantis annectens Werner, 1910
MATERIAL.— Omauha Lodge, 3 December 2013, 16º11ʹ55.4ʺS, 12º24ʹ0.3ʺE, 338 m (CAS
255056).
COMMENTS.— Both this specimen and another now in the INBAC collections were collected
inside a toilet water tank, one of few available sources of standing water in the area. These speci-
mens represent the fourth record for the species in the country. The species reaches its northern dis-
tribution in Angola, in Novo Redondo/Gabela, Kwanza Sul Province (Poynton and Haacke 1993).
Phrynomantis annectens has previously been recorded from the Mutiambo River and Caraculo in
Namibe Province (Poynton and Haacke 1993). The current records represent the southernmost
known distribution of the species in Angola, however it is common in Namibia and South Africa
(Channing 2001; du Preez and Carruthers 2009) and is likely to occur throughout the province
wherever water is available.
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Reptilia
Squamata
Agamidae
ANCHIETASGROUND AGAMA – Fig. 3
Agama anchietae Bocage, 1896
MATERIAL.— INP, 29 November 2013, 16º39ʹ27.12ʺS, 12º26ʹ17.04ʺE, 459 m (CAS 254778);
Pico Azevedo, 7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE, 359 m (CAS 254942); NNP, 28
November 2013, 15º46ʹ27.4ʺS. 12º19ʹ59.2ʺE, 264 m (CAS 254956).
COMMENTS.— These specimens have distinctive black-tipped spines on the palmar scales,
which distinguish A. anchietae from the morphologically similar A. aculeata Merrem, 1820 that
occurs sympatrically in Southern Angola and Namibia (Branch 1993). Agama anchietae was
described from Angola by Bocage based on specimens from Catumbela, Benguela and Dombe (all
in Benguela Province), and Moçamedes (currently Namibe, Namibe Province). In Angola, the
species is known to occur in Namibe Province (Bocage 1896, 1897; Laurent 1964a), Benguela
Province (Bocage 1863, 1896, 1897), and Bié Province (Schmidt 1933). Our specimens represent
the southernmost localities of the species in Angola, although it is certainly distributed continu-
ously across the entire province.
NAMIB ROCK AGAMA – Fig. 4
Agama planiceps Peters, 1862
MATERIAL.— ‘Lion Cave’ at 3.4 km SW of Espinheira camp, 30 November 2013,
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 21
FIGURE2. Marbled Rubber Frog, Phrynomantis annectensWerner, 1910, from Palmwag, Kunene Region, Namibia.
Photo courtesy of Randall Babb.
16º48ʹ45.0ʺS, 12º20ʹ22.9ʺE, 463
m (CAS 254753); Omauha
Lodge camp, 2 December 2013,
16º11ʹ55.4ʺS, 12º24ʹ0.3ʺE, 335
m (CAS 254832); INP, north of
Tambor, 4 December 2013,
15º59ʹ46.0ʺS, 12º24ʹ24ʺE, 307 m
(CAS 254839); INP, south side
of Curoca River crossing, 29
November 2013, 16º18ʹ15.6ʺS,
12º25ʹ1.56ʺE, 209 m (CAS
254845), 1 December 2013,
16º18ʹ14.8ʺS, 12º24ʹ2159.8ʺE,
210 m (CAS 254848); Pediva
Hot Springs, 2 December 2013,
16º7ʹ19.7ʺS, 12º33ʹ40.0ʺE, 244
m (CAS 254859); Namibe-
Lubango road, road marker 59,
1.8 km west by road from Carac-
ulo, north side of the road,
6 December 2013, 15º0ʹ57.6ʺS, 12º38ʹ36.8ʺE, 500 m (CAS 254900, CAS 254910); Pico Azevedo,
7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE, 359 m (CAS 254941).
22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 3. Anchieta’s Ground Agama, Agama anchietae Bocage, 1866, from Sesfontein, Kunene Region, Namibia.
Photo courtesy of Johan Marais.
FIGURE 4. Adult male Namib Rock Agama, Agama planiceps Peters, 1862,
from Pico Azevedo. Photo by Luis Ceríaco.
COMMENTS.— The species has been cited throughout Angola, although many previous records
remain doubtful. Agama planiceps appears to be restricted to arid savannas and Namibe Province,
or perhaps Beguela, is most probably its northernmost limit. Mertens (1938) described the sub-
species Agama planiceps shackii from Cubal (Benguela Province), and it is likely that the central
and northern Angolan records of planiceps are in fact shackii. The status of this form remains in
question, but preliminary examination of topotypical material suggests that it is specifically dis-
tinct. Our specimens fit the morphological description and current known distribution of nomino-
typical planiceps. The species is cited for several localities in Namibe Province, including Biballa
(Bocage 1895), Fazenda Bumbo (Laurent 1964a), and Pico Azevedo (Schmidt 1933). Some of our
records represent a southern range extension of the species in the Province, although a record from
the Kwito region, in Cunene Province (Angel 1923) is the southernmost Angolan record. The
species likely also occurs in rocky areas extending south to the Namibian border. The majority of
these specimens were seen basking on the top of rocks. Males and females present a striking sex-
ual dimorphism, with the males being considerably larger than the females, and having an intense-
ly red head and a dark blue body (the posterior half of the tail is usually also red). Females are typ-
ically dark-grey on the dorsum with yellow marking on the head and dorsum.
Gekkonidae
FITZSIMONS’ THICK-TOED GECKO – Fig. 5
Chondrodactylus fitzsimonsi (Loveridge, 1947)
MATERIAL.— Espinheira, 30 November 2013, 16º47ʹ14.3ʺS, 12º21ʹ29.4ʺE, 457 m (CAS
254814); INP, north of Tambor, 4 December 2013, 15º59ʹ46.9ʺS, 12º24ʹ24ʺE, 300 m (CAS
254841).
COMMENTS.— Only four records of this species are known for the country: Ongueria, 55 km
from Sá da Bandeira (presently Lubango), in Huíla Province (Laurent 1964a), Praia das Conchas
(Laurent 1964a), “around Moçâmedes in the road to da Bandeira” (Laurent 1964a) and Pico
Azevedo (Schmidt 1933), both in Namibe Province. Our records represent southern range exten-
sions for the species in the country, although the species extends into west-central Namibia (Bauer
et al. 1993). In both Namibia and Angola it can occur sympatrically with other, similarly sized con-
geners, but is generally more restricted to rocky habitats than either C. pulitzerae or C. turneri.
PULITZERSTHICK-TOED GECKO – Fig. 6
Chondrodactylus pulitzerae (Schmidt, 1933)
MATERIAL.— INP, 9.65 km (by air) west-south-west of Espinheira, 30 November 2013,
16º48ʹ43.19ʺS, 12º16ʹ16.55ʺE, 488 m (CAS 254790–CAS 254792); Espinheira, 29 November
2013, 16º47ʹ11.01ʺS, 12º21ʹ28.77ʺE, 457 m (CAS 254796–254798), 16º47ʹ8.1ʺS, 12º21ʹ16.44ʺE,
457 m (CAS 254804, 30 November, 16º47ʹ14.3ʺS, 12º21ʹ29.4ʺE, 457 m (CAS 254814–254815),
16º47ʹ15.1ʺS, 12º21ʹ23.8ʺE, 457 m (CAS 254816), 16º47ʹ14.0ʺS, 12º21ʹ29.6ʺE, 457 m (CAS
254817), 16º47ʹ17.3ʺS, 12º21ʹ25.1ʺE, 457 m (CAS 254818), 16º47ʹ11.1ʺS, 12º21ʹ30.2ʺE, 457 m
(CAS 254819); Omauha Lodge, 28 November 2013, 16º11ʹ55.01ʺS, 12º24ʹ3.12ʺE, 338 m (CAS
254830), 2 December 2013, 16º11ʹ55.4ʺS, 12º24ʹ0.3ʺE, 338 m (CAS 254833), 28 November 2013,
16º11ʹ54.19ʺS, 12º24ʹ2.45ʺE, 338 m (CAS 254843); INP, Rio Curoca in Pediva Hot Springs area,
3 December 2013, 16º17ʹ0.93ʺS, 12º33ʹ39.81ʺE, 247 m (CAS 254854); Namibe-Lubango road,
road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road, 6 December 2013,
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 23
24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 5. Adult male FitzSimons’Thick-toed Gecko, Chondrodactylus fitzsimonsi (Loveridge, 1947), from northern
Kaokoveld, Kunene Region, Namibia. Photo courtesy of Johan Marais.
FIGURE 6. Subdult male Pulitzer’sThick-toed Gecko, Chondrodactylus pulitzerae. (Schmidt, 1933), from Chimalavera,
Benguela Province, Angola. Photo by Luis Ceríaco.
15º0ʹ57.3ʺS, 12º38ʹ32.6ʺE, 497 m (CAS 254915); Pico Azevedo, 7 December 2013, 15º32ʹ2.4ʺS,
12º29ʹ31.1ʺE, 359 m (CAS 254920, CAS 254943).
COMMENTS.— This species occurs from the southern regions of Angola in Namibe (Bocage
1867, 1887, 1895; Laurent 1964a; Schmidt 1933) and Cunene (Monard 1937) provinces to
Malange, where it reaches its northern distribution in Capanda (Ceríaco et al. 2014). There are
records from the far northwest of Namibia as well, although these have been consistently treated
as either C. turneri or C. laevigatus. This species was originally described as a subspecies of
C. bibronii (now regarded as limited to South Africa, southern Namibia and adjacent areas; Benyr
1995; Bauer and Lamb 2005) based on material from Pico Azevedo (Schmidt 1933). Heinz (2011)
provided evidence for the specific distinctness of C. pulitzerae; see Ceríaco et al. (2014) for brief
discussion ofthe nomenclatural and taxonomic history of this taxon. One specimen (CAS 254920)
represents topotypical material. The species was common and found hiding on shaded areas of
rocky crevices, houses, and other structures.
LARGE-SCALED THICK-TOED GECKO
Pachydactylus scutatus Hewitt, 1927
MATERIAL.— Espinheira, 30 November 2013, 16º47ʹ51.8ʺS, 12º21ʹ15.2ʺE, 457 m (CAS
254826).
COMMENTS.— This specimen is the first published record of Pachydactylus scutatus for
Angola. Pachydatylus scutatus angolensis is now recognized as a distinct species (see below;
Bauer et al. 2002). Five additional specimens, all from Iona, are present in the Ditsong National
Museum of Natural History (TM 40615–18 from Espinheira, TM 40751 from 16°54ʹS, 12°35ʹE).
ANGOLAN THICK-TOED GECKO – Fig. 7
Pachydactylus angolensis (Loveridge, 1944)
MATERIAL.— Namibe-Lubango road, 2 km E (by road) of Mangueiras, south side of the road,
5 December 2013, 15º2ʹ37ʺS, 13º9ʹ36ʺE, 625 m (CAS 254887).
COMMENTS.— This poorly known taxon was described from Benguela Province (Loveridge
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 25
FIGURE 7. Juvenile Angolan Thick-toed Gecko, Pachydactylus angolensis (Loveridge, 1944), from Chimalavera,
Benguela Province, Angola. Photo by Luis Ceríaco.
1944a). Laurent (1964a) subsequently reported additional specimens from the “environs de
Moçâmedes” (now Namibe, Namibe Province). More recently, Wulf Haacke collected 16 speci-
mens from both Bengela and Namibe Provinces. The Namibe localities include Lungo, Lucira, San
Nicolau, and Saco de Giraul. There is a single record from extreme northern Namibia (J. Boone,
pers. comm.), suggesting that the species actually has a fairly broad range from south of the
Kunene to Hanha in Benguela (TM 46558).
SPECKLED THICK-TOED GECKO
Pachydactylus punctatus Peters, 1854
MATERIAL.— INP, 29 November 2013, 16º39ʹ24.1ʺS, 12º26ʹ12.2ʺE, 460 m (CAS 254781–
254784); Espinheira, 29 November 2013, 16º47ʹ11.01ʺS, 12º21ʹ28.77ʺE, 457 m (CAS 254799–
254800), 16º47ʹ7.02ʺS, 12º21ʹ16.86ʺE, 457 m (CAS 254806); 16º47ʹ4.2ʺS, 12º21ʹ16.62ʺE, 457 m
(CAS 254809–254810), 16º47ʹ12.71ʺS, 12º21ʹ28.67ʺE, 457 m (CAS 254812), 16º47ʹ20.3ʺS,
12º21ʹ27.6ʺE, 457 m (CAS 254960).
COMMENTS.— The species is known from southwestern Angola in the provinces of Benguela
(Bocage 1867, 1895; Boulenger 1885; Hellmich 1957b; Laurent 1954), Huila (Monard 1931, 1937,
Laurent 1964a), Cunene (Laurent 1964a), and Namibe (Schmidt 1933, Laurent 1964a). Members
of the P. punctatus complex have been confused with other southern African Pachydactylus,
including P. occellatus and P. geitje (Bocage 1867, 1885, 1895; Boulenger 1905; Frade 1963). The
previous report of P. serval in Angola (Monard 1931) likely corresponds to P. punctatus. A phylo-
geographic study of P. punctatus is being currently undertaken (Heinz 2011) and it appears that this
taxon comprises multiple unnamed cryptic species. At least two, possibly three, taxa in this com-
plex occur in southwestern Angola. Pachydactylus punctatus sensu lato is the most common ter-
restrial gecko in most of northern Namibia as well as southern Angola.
KAOKOLAND ROCK GECKO
Pachydactylus cf. oreophilus McLachlan and Spence, 1967
MATERIAL.— Omauha Lodge, 28 November 2013, 16º11ʹ55.01ʺS, 12º24ʹ3.12ʺE, 338 m (CAS
254829).
COMMENTS.— Pachydactylus oreophilus was described from near Sesfontein in northwestern
Namibia. Specimens assigned to this species extend northwards at least as far as the southern low-
land portions of Benguela Province, Angola. Preliminary molecular phylogenetic data suggest that
northern populations, including all of those in Angola and possibly those along the Kunene River
in Namibia, are not conspecific with the nominotypic form. The specimen was collected at night,
basking near a lamp 2.5 m off the ground, in Omauha Lodge.
BARNARDSNAMIB DAY GECKO – Fig. 8
Rhoptropus barnardi Hewitt, 1926
MATERIAL.— Approximately 7.35 km north-west (by road) of Pico Azevedo, 7 December
2013, 15º28ʹ30.7ʺS, 12º27ʹ47.5ʺE, 420 m (CAS 254759, CAS 254761); Omauha Lodge, 4 Decem-
ber 2013, 16º12ʹ1.2ʺS, 12º24ʹ0.1ʺE, 343 m (CAS 254837); INP, Rio Curoca crossing, North side of
the river, 1 December 2013, 16º18ʹ6.8ʺS, 12º25ʹ13.0ʺE, 206 m (CAS 254844); INP, Rio Curoca
crossing, south side of the river, 1 December 2013, 16º18ʹ14.7ʺS, 12º25ʹ0.0ʺE, 210 m (CAS
254846–254847); INP, Rio Curoca in the Pediva Hot Springs area, 2 December 2013,
26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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16º17ʹ0.93ʺS, 12º33ʹ39.81ʺE, 247 m (CAS 254852), 16º17ʹ14.3ʺS, 12º33ʹ35.9ʺE, 238 m (CAS
254856), 16º17ʹ24.01ʺS, 12º33ʹ43.9ʺE, 270 m (CAS 254863), Namibe-Lubango road, 2 km east
(by road) of Mangueiras, south side of the road, 5 December 2013, 15º2ʹ40.8ʺS, 13º9ʹ32.6ʺE, 664
m (CAS 254890); NNP, 28 November 2013, 15º46ʹ23.4ʺS, 12º19ʹ58.9ʺE, 264 m (CAS 254954).
COMMENTS.— Until now, this species has been known only from one published locality in
Angola. Laurent (1964a) cites the specimen from a locality “60 km on the road from Moçâmedes
[presently Namibe] to Sá da Bandeira [presently Lubango]”, the same locality from which he
described R. taeniostictus, which we also collected (see account below). This species is widely dis-
tributed in northwestern Namibia, occurring as far inland as the Otavi-Grootfontein region, due
south of western Cuando Cubango Province. It is rupicolous and can be found on small rocky piles
and ridges, as well as on larger boulders. The extent of its distribution in Angola is poorly known,
in part becase many records from Namibe and Huila are assignable to a morphologically similar,
but undescribed congener (see Rhoptropus sp. below).
TWO-PORED NAMIB DAY GECKO – Fig. 9
Rhoptropus biporosus Fitzsimons, 1957
MATERIAL.— INP, 29 November 2013, 16º32ʹ0.48ʺS, 12º26ʹ44.16ʺE, 378 m (CAS 254779,
16º39ʹ26.04ʺS, 12º26ʹ13.5ʺE, 460 m (CAS 254780); INP, 20 km south-south-west (by air) of Espi-
nheira, 30 November 2013, 16º55ʹ54.1ʺS, 12º14ʹ42.0ʺE, 631 m (CAS 254786–254788),
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 27
FIGURE 8. Adult Barnard’s Namib Day Gecko, Rhoptropus barnardi Hewitt, 1926, from Kamanjab, Kunene Region,
Namibia. Photo courtesy of Johan Marais.
16º48ʹ43.19ʺS, 12º16ʹ16.55ʺE, 485 m (CAS 254959), Espinheira, 29 November 2013,
16º47ʹ19.9ʺS, 12º21ʹ27.4ʺE, 457 m (CAS 254794), 16º47ʹ7.08ʺS, 12º21ʹ16.02ʺE, 457 m (CAS
254802–254803), 16º47ʹ7.02ʺS, 12º21ʹ16.86ʺE, 457 m (CAS 254805), 16º47ʹ4.26ʺS,
12º21ʹ16.62ʺE, 457 m (CAS 254811), 16º47ʹ12.71ʺS, 12º21ʹ28.67ʺE, 457 m (CAS 254813),
16º47ʹ20.2ʺS, 12º21ʹ27.9ʺE, 457 m (CAS 254958), 30 November 2013, 16º47ʹ14.3ʺS,
12º21´29.4ʺE (CAS 254820), 16º47ʹ8.7ʺS, 12º21ʹ30.3ʺE, 457 m (CAS 254821), 16º47ʹ18.1ʺS,
12º21ʹ26.2ʺE, 457 m (CAS 254822), 16º47ʹ33.6ʹS, 12º21ʹ19.0ʺE, 457 m (CAS 254823),
16º47ʹ41.5ʺS, 12º21ʹ17.3ʺE, 457 m (CAS 254824), 16º47ʹ45.3ʺS, 12º21ʹ15.9ʺE, 457 m (CAS
254825); NNP, 28 November 2013, 15º46ʹ27.4ʺS, 12º19ʹ59.2ʺE, 264 m (CAS 254957–254958).
COMMENTS.— The species occurs in the rocky outcrops in arid habitats inland of the northern
Namib dune fields in the vicinity of Orupembe, in the Kaokoveld and across the Cunene River to
Angola (Bauer and Good 1996). The only published record of this species for Angola is from the
Pico Azevedo region (Bauer and Good 1996), although Wulf Haacke collected numerous speci-
mens from localities across southern Namibe, as well as from near Otchinjau, Cunene Province
(specimens in Ditsong National Museum of Natural History).
BOULTONSNAMIB DAY GECKO – Fig. 10
Rhoptropus boultoni boultoni Schmidt, 1933
MATERIAL.— INP, 3.4 km southwest (by air) of Espinheira, vicinity of “Lion Cave”, 30
November 2013, 16º48ʹ73.5ʺS, 12º20ʹ23.2ʺE, 463 m (CAS 254752); Approximately 7.35 km
north-west (by road) of Pico Azevedo, 7 December 2013, 15º28ʹ33.2ʺS, 12º27ʹ45.7ʺE, 421 m (CAS
254757–254758); Espinheira, 16º47ʹ29.4ʺS, 12º21ʹ6.06ʺE, 457 m (CAS 254795); Omauha Lodge,
28 November 2013, 16º11ʹ52.5ʺS, 12º23ʹ59.3ʺE, 335 m (CAS 254828, 2 December 2013,
28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 9. Close-up of head of adult specimen of Two-Pored Namib Day Gecko, Rhoptropus biporosus FitzSimons,
1957 from northwest of Palmwag, Kunene Region, Namibia. Photo courtesy of Johan Marais.
16º12ʹ1.2ʺS, 12º24ʹ0.1ʺE, 343 m (CAS 254834); INP, Rio Curoca crossing, south side of river,
1 December 2013, 16º18ʹ14.7ʺS, 12º25ʹ0.0ʺE, 210 m (CAS 254849–254850), 2nd December 2013,
16º17ʹ19.7ʺS, 12º33ʹ40.0ʺE, 244 m (CAS 254857–254858, CAS 254861–254862), 29 November
2013, 16º18ʹ15.6ʺS, 12º25ʹ1.56ʺE, 209 m (CAS 254865); Leba Pass, between river and highway, 5
December 2013, 15º4ʹ13.2ʺS, 13º14ʹ37.7ʺE, 1676 m (CAS 254880); Namibe-Lubango road, 2.0
km east (by road) of Mangueiras, south side of the road, 5 December 2013, 15º2ʹ40.8ʺS,
13º9ʹ32.6ʺE, 664 m (CAS 254892), 15º2ʹ40.7ʺS, 13º9ʹ31ʺE, 640 m (CAS 254894, 15º0ʹ55.1ʺS,
12º38ʹ32.8ʺE, 497 m (CAS 254902); Namibe-Lubango road, road marker 59, 1.8 km west by road
of Caraculo, 6 December 2013, 15º00ʹ55.1ʺS, 12º38ʹ32.8ʺE, 497 m (CAS 254903); Pico Azevedo,
7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE, 359 m (CAS 254921–254926), 15º32ʹ5.8ʺS.
12º29ʹ29.5ʺE, 366 m (CAS 254946–254947, CAS 254949–254950); Espinheira, 29 November
2013, 16º47ʹ20.2ʺS, 12º21ʹ27.9ʺE, 457 m (CAS 254958).
COMMENTS.— This taxon is widespread from northwestern Namibia north at least to northern
Namibe Province.
MONTANE NAMIB DAY GECKO
Rhoptropus boultoni montanus Laurent, 1964
MATERIAL.— Leba Pass overlook, 5 December 2013, 15º4ʹ37.2ʺS, 13º13ʹ58.5ʺE, 1682 m
(CAS 254866, CAS 254867; 15º4ʹ38.3ʺS, 13º13´57.0ʺE, 1682 m (CAS 254868); 15º4ʹ36.0ʺS,
13º14ʹ1.6ʺE, 1682 m (CAS 254869–254872); Leba Pass, between river and highway, 15º04ʹ13.2ʺ
S, 13º14ʹ37.3ʺE, 1676 m (CAS 254882).
COMMENTS.— The subspecies was described from the Leba Escarpment (“60 km on the road
to Moçâmedes [now Namibe, Namibe Province] from Sá da Bandeira [now Lubango, Huila
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 29
FIGURE 10. Adult Boulton’s Namibe Day Gecko, Rhoptropus boultoni boultoni Schmidt, 1933, from east of Kaman-
jab, Kunene Region, Namibia. Photo courtesy of Johan Marais.
Province]”, Laurent 1964a). A large series of specimens in the Ditsong National Museum of Nat-
ural History are derived from localities near Lubango, in Huila. Our specimens are topotypical and
were collected on the Namibe side of the provincial boundary. Specimens were found basking on
high elevation granite rocks covered with bryophytes. Molecular phylogenetic studies (A. Kuhn,
unpublished) reveal that this taxon is specifically distinct from R. boultoni. Its formal elevation to
specific status will be justified in detail elsewhere.
ANGOLAN NAMIB DAY GECKO
Rhoptropus taeniostictus Laurent, 1964
MATERIAL.— Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of the
road, 5 December 2013, 15º2ʹ40.8ʺS, 13º9ʹ32.6ʺE, 664 m (CAS 254889); Namibe-Lubango road,
road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road, 6 December 2013,
15º0ʹ58.0ʺS, 12º38ʹ37.3ʺE, 490 m (CAS 254895);15º0ʹ57.9ʺS, 12º38ʹ42.3ʺE, 472 m (CAS 254897–
254898), 15º0ʹ57.6ʺS, 12º38ʹ36.8ʺE, 500 m (CAS 254901), 15º1ʹ0.7ʺS, 12º38ʹ31.9ʺE, 492 m (CAS
254904–254905), 15º0ʹ58.8ʺS, 12º38ʹ33.8ʺE, 491 m (CAS 254908), 15º0ʹ58.9ʺS, 12º38ʹ32.4ʺE,
497 m (CAS 254911), 15º0ʹ57.3ʺS, 12º38ʹ32.6ʺE, 497 m (CAS 254916), 15º1ʹ0.1ʺS, 12º38ʹ31.9ʺE,
497 m (CAS 254917–254918), 15º1ʹ0.9ʺS, 12º38ʹ30.4ʺE, 503 m (CAS 254919).
COMMENTS.— The Angolan endemic R. taeniostictus was described from a single specimen
from “60 km on the road from Moçâmedes [presently Namibe] to Sá da Bandeira [presently Luban-
go]”. Laurent (1964a) also considered the populations of R. barnardi from Mucungo cited by
Schmidt (1933) as referable to R. taeniostictus. The species appears restricted to Namibe Province
and is represented by many specimens in our collection as well as more widespread Namibe local-
ities in the Ditsong National Museum of Natural History.
Rhoptropus sp.
MATERIAL.— Espinheira, 29 November 2013, 16º47ʹ32.7ʺS, 12º21ʹ14.4ʺE, 562 m (CAS
254801); Omauha Lodge, 4 December 2013, 16º12ʹ1.2ʺS, 12º24ʹ0.1ʺE, 343 m (CAS 254836, CAS
254955); INP, north of Tambor, 4 December 2013, 15º59ʹ46.9ʺS, 12º24ʹ24.0ʺE, 300 m (CAS
254842, CAS 254762, CAS 254766), 15º28ʹ31.7ʺS, 12º27ʹ43.9ʺE, 408 m (CAS 254765, CAS
254760); Leba Pass, between river and highway, 5 December 2013, 15º4ʹ12.1ʺS, 13º14ʹ36.5ʺE,
1680 m (CAS 254873), 15º4ʹ13.2ʺS, 13º14ʹ37.7ʺE, 1676 m (CAS 254879, CAS 254881, CAS
254883); Namibe-Lubango road, 2 m east (by road) of Mangueiras, south side of the road,
5 December 2013, 15º2ʹ40.8ʺS, 13º9ʹ32.6ʺE, 664 m (CAS 254890–254891), 15º2ʹ40.7ʺS,
13º9ʹ31.0ʺE, 640 m (CAS 254893, CAS 254894).
COMMENTS.— This undescribed species is morphologically similar to both R. barnardi and
R. biporosus, but appears to be endemic to southern Angola. Populations from the Escarpment are
both morphologically and genetically different from those below the Escarpment.This taxon is cur-
rently under study as part of a phylogenetic analysis and revision of the genus as a whole (A. Kuhn
and A. Bauer, in prep.).
Scincidae
BOGERTSSPECKLED WESTERN BURROWING SKINK – Fig. 11
Typhlacontias punctatissimus bogerti Laurent, 1964
MATERIAL.— Espinheira, 29 November 2013, 16º47ʹ7.02ʺS, 12º21ʹ16.86ʺE, 457 m (CAS
30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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254807); Pico Azevedo, 7 De-
cember 2013, 15º32ʹ2.4ʺS,
12º29ʹ31.1ʺE, 359 m (CAS
254932–254938), 15º32ʹ5.8ʺS,
12º29ʹ29.5ʺE, 366 m (CAS
254944–254945).
CO M M E N T S . — Ha a ck e
(1997) reviewed the taxonomic
and nomenclatural history of
Typhlacontias punctatissimus
and its subspecies and recog-
nized two sympatric subspecies
in southern Angola — T. puncta-
tissimus punctatissimus Bocage,
1873, and the Angolan endemic
T. punctatissimus bogerti Lau-
rent, 1964. In all of our speci-
mens, the second and third upper labials are in contact with the eye and there is a second supraoc-
ular. Both characters fit the description presented by Haacke (1997) as diagnostic for
T. punctatissimus bogerti. The species is known to be viviparous and one female specimen (CAS
254945) contains an almost fully developed neonate.
VARIABLE SKINK
Trachylepis varia (Peters, 1867)
MATERIAL.— Leba Pass, 5 December 2013, 15º4ʹ12.1ʺS, 13º14ʹ36.2ʺE, 1680 m (CAS
254874), 15º4ʹ13.2ʺS, 13º14ʹ37.7ʺE, 1676 m (CAS 254884).
COMMENTS.— The species occurs throughout Angola, with many records in the provinces of
Benguela (Parker 1936; Bocage 1895, 1896; Monard 1937; Hellmich 1957a; Mertens 1938;
Boulenger 1905) and Huila (Bocage 1895; Monard 1937). Although other records from Namibe are
12 km W of Humbia (TM 40128–29) and Chapeau Armado turnoff (TM 41131). This species is
typically associated with relatively mesic microclimates and is, therefore, excluded from the hyper-
arid areas of Namibe. This skink has a broad distribution across much of sub-Saharan Africa and
includes several cryptic species, two of which are present in Angola. There are records from across
Angola and from all bordering countries as well. The phylogeography of the T. varia complex is
presently under study (J. Weinell, pers. comm.).
SPECKLED SAND SKINK – Fig. 12
Trachylepis punctulata (Bocage, 1872)
MATERIAL.— Espinheira, 29 November 2013, 16º47ʹ20.6ʺS, 12º21ʹ27.2ʺE, 457 m (CAS
254793); Namibe-Lubango road, road marker 59, 1.8 km west (by road) of Caraculo, on the north
side of the road, 6 December 2013, 15º0ʹ55.1ʺS, 12º38ʹ32.8ʺE, 497 m (CAS 254903); Praia do
Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013, 16º16ʹ20.4ʺS, 11º49ʹ53.9ʺE,
8 m (CAS 254769–254771), 16º16ʹ39.3ʺS, 11º49ʹ20.5ʺE, 8 m (CAS 254775).
COMMENTS.— The species was originally described by Bocage based on material from “Rio
Coroca, sur le littoral de Mossamedes, Angola” (Bocage 1872). The type locality is presumably the
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 31
FIGURE 11. Adult Bogert’s Speckled Western Burrowing Skink Typhla-
contias punctatissimus bogerti Laurent, 1964, from Pico Azevedo. Photo by
Edward Stanley.
region near the mouth of Curoca River, from the vicinity of Tombwa (formerly Porto Alexandre).
Our specimens were collected among plants between dunes along the coast south of Tombwa, and
these agree morphologically with the original description for the species. A comparison with the
type material was impossible due to its destruction in the fire that destroyed the Lisbon Museum
in 1978. Several uncatalogued specimens in the Museu Nacional de História Natural do Porto col-
lected in 1905 by the Portuguese explorer Francisco Newton are congruent with the specimens col-
lected by us. Newton’s specimens are still in their original jar and are labeled “Mossamedes” (pre-
sumably refering to the province as a whole, not the city of Mossamedes = Namibe). They are part
of a collection of vertebrates that the explorer made in the region. The herpetological specimens
were only partly studied and published upon (Ferreira 1904, 1906; Ceríaco et al. 2014), in contrast
to the bird and mammals collections (Seabra 1906a, 1906b, 1906c, 1906d, 1907). It is probable that
these specimens are from Tombwa, as this was the main place where Newton collected while in the
province (see bird records – Seabra 1906a). This is a common species in much of Namibia,
Botswana, and central South Africa, as well as portions of Zambia, Zimbabwe, and Mozambique
(Portik and Bauer 2012).
HOESCHSSKINK – Fig. 13
Trachylepis hoeschi (Mertens, 1954)
MATERIAL.— Rio Curoca in the Pediva Hot Springs area, 2 December 2013, 16º17ʹ0.93ʺS,
12º33ʹ39.81ʺE, 247 m (CAS 254851); NNP, 15º46ʹ25.9ʺS, 12º19ʹ59.0ʺE, 247 m (CAS 254952).
COMMENTS.— The only published Angolan record is from Laurent (1964a), from “Plage das
Conchas,” Namibe Province. Our specimens are, respectively, 75 and 135 km SE of Laurent’s site.
the Ditsong National Natural History Museum holds a small series of this species (TM 40733–37)
32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 12. Adult Speckled Sand Skink, Trachylepis punctulata (Bocage, 1872), from Kamanjab, Kunene Region,
Namibia. Photo courtesy of Johan Marais.
from Iona National Park. The Angolan records extend the core distribution of its range in north-
western Namibia (Branch 1998).
ANGOLAN BLUE-TAILED SKINK
Trachylepis laevis (Boulenger, 1907)
MATERIAL.— INP, north of Tambor, 4 December 2013, 15º59ʹ47.1ʺS. 12º24ʹ25.6ʺE, 314 m
(CAS 254838).
COMMENTS.— The species was described by Boulenger from Maconjo, in northern Namibe
Province (Boulenger 1907b). Laurent (1964a) recorded this species in Namibe Province from
“Munhino 50 km west of da Bandeira.” Hellmich (1957a) cited the species for Piri-Dembos,
Kwanza Norte Province, but this record is dubious. The Ditsong National Natural History Muse-
um houses numerous specimens from localities in Namibe and southern Benguela below the
Escarpment. The species occurs also in the Kamanjab area and Damaraland in northwestern
Namibia (Bauer et al. 1993). This lizard is extremely dorsoventrally depressed in association with
its crevice dwelling habits and was, for a time, placed in a monotypic genus, Oelofisa, in recogni-
tion of its highly autapomorphic morphology (Steyn and Mitchell 1965).
WESTERN THREE-STRIPED SKINK
Trachylepis occidentalis (Peters, 1867)
MATERIAL.— Pico Azevedo, 7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE, 359 m (CAS
254931).
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 33
FIGURE 13. Adult Hoesch’s Skink, Trachylepis hoeschi (Mertens, 1954), from Kamanjab, Kunene Region, Namibia.
Photo courtesy of Johan Marais
COMMENTS.— The species has been cited from 35 km south of the city of Namibe (Laurent
1964a) and from Curoca River (Bocage 1895). Three specimens in the Ditsong Natural National
History Museum originate from the Rio Curoca mouth and from Namibe. In addition to our spec-
imen from Pico Azevedo several individuals of the species were observed near Espinheira camp
(specimens not collected). In Angola this skink takes refuge in holes it digs in the sand at the base
of spiny shrubs of the genus Blepharis. It is widely distributed in western South Africa and much
of central and western Namibia (Branch 1988).
WEDGED-SNOUTED SKINK – Fig. 14
Trachylepis acutilabris (Peters, 1862)
MATERIAL.— INP, 3.4 km south-west (by air) of Espinheira, vicinities of “Lion Cave”, 30
November 2013, 16º48ʹ54.4ʺS, 12º20ʹ13.7ʺE, 450 m (CAS 254751); INP, car wreck 20 km south-
south-west (by air) of Espinheira, 30 November 2013, 16º55ʹ53.81ʺS, 12º14ʹ45.42ʺE, 616 m (CAS
254789); Namibe-Lubango road, road marker 59, 1.8 km (by road) of Caraculo, north side of the
road, 6 December 2013, 15º0ʹ59.3ʺS, 12º38ʹ33.6ʺE, 488 m (CAS 254899), 15º0ʹ58.8ʺS,
12º38ʹ33.8ʺE, 491 m (CAS 254907); Pico Azevedo, 7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE,
359 m (CAS 254927–254931).
COMMENTS.— This species is similar to lacertid lizards in morphology and diet (Castanzo and
Bauer 1992). Its elongate toes and countersunk lower jaw are consistent with its burrowing habits.
It typically occupies burrows at the base of vegetation in sandy soils from Namibia through west-
ern Angola to the Democratic Republic of Congo and Cabinda (Branch 1998).
WESTERN ROCK SKINK
Trachylepis sulcata (Peters, 1867)
MATERIAL.— INP, Rio Curoca in Pediva Hot Springs area, 3 December 2013, 16º17´0.93ʺS,
12º33ʹ39.81ʺE, 247 m (CAS 254853); Leba Pass, 5 December 2013, 15º4ʹ12.1ʺS, 13º14ʹ36.2ʺE,
1680 m (CAS 254875); Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of
the road, 5 December 2013, 15º2ʹ40.7ʺS, 13º9ʹ31ʺE, 625 m (CAS 254886), 15º2ʹ40.8ʺS,
13º9ʹ32.6ʺE, 664 m (CAS 254888).
COMMENTS.— Trachylepis sulcata is a rupicolous skink ranging from the Western Cape
Province of South Africa north to southern Angola. Trachylepis sulcata ansorgii (Boulenger
1907b) was described from southern Angola to accommodate specimens with bright throat and
infralabial coloration. Laurent (1964a) and Mertens (1971) considered it valid and the latter iden-
tified some Namibian specimens as intergrades between T. s. ansorgii and T. s. sulcata, whereas
Haacke (1972) considered specimens on the Namibian side of the Kunene river to be referable to
T. s. ansorgii. Some specimens from west of the Great Escarpment in northwestern Namibia exhib-
it the diagnostic coloration of ansorgii, but the two subspecies seem to have no fixed differences
in scalation (Bauer et al. 1993). Although Portik et al. (2011) did not include typical T. s. ansorgii
in their molecular sampling, preliminary integration of samples from our collection into their data
set reveals no significant difference from putative T. s. ansorgii from Namibe and the nominotyp-
ical form. We therefore treat T. sulcata as a monotypic species.
34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 35
FIGURE 14. Adult Wedged-Snouted Skink, Trachylepis acutilabris (Peters, 1862), from Kamanjab, Kunene Region,
Namibia. Photo courtesy of Johan Marais.
Lacertidae
ANCHIETASSHOVEL-SNOUT LIZARD
Meroles anchietae (Bocage, 1867)
MATERIAL.— Praia do Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013,
16º16ʹ29.1ʺS, 11º49ʹ05.0ʺE, 8 m (CAS 254773).
COMMENTS.— Bocage (1867) described this species from “Mossamedes” (Bocage 1867). Sur-
prisingly, this remains the only published locality for this species in Angola (Bocage, 1867, 1895),
despite it being common and widely distributed in barchan dunes from the Klinghardt Mountains
north through the Namib of western Namibia. In the Newton collections in Porto, there are sever-
al uncatalogued specimens corresponding to this species from “Mossamedes” and the Ditsong
National Natural History Museum has material collected by Wulf Haacke from Porto Alexandre
and Foz de Cunene.
RETICULATE SAND LIZARD – Fig. 15.
Meroles reticulatus (Bocage, 1867)
MATERIAL.— Praia do Navio coastal dunes, ca 124 km SSW of Namibe, 8 December 2013,
16º16ʹ39.3ʺS, 11º49ʹ20.5ʺE, 8 m (CAS 254776).
COMMENTS.— Bocage (1867) described this species from “Mossamedes.” Bocage (1895) sub-
sequently clarified that the types had come from the littoral zone at Rio Coroca [= Rio Curoca,
southern Namibe Province, Angola]. The range of this species extends towards Namibia to the area
of Conception Bay on the central coast. Although it is well documented within its Namibian range,
this specimen is only the third published locality for Angola. The species is, however, well repre-
sented from numerous localities in Namibe by specimens in the Ditsong National Museum of Nat-
ural History.
36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 15. Adult specimen of Anchieta’s Shovel-snout Lizard, Meroles anchietae, (Bocage, 1867) from gravel plains
north of Henties Bay, Erongo Region, Namibia. Photo courtesy of Johan Marais.
HAACKESSAND LIZARD
Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012
MATERIAL.— 5 km NW (by road) of Pico Azevedo, 7 December 2013, 15º28ʹ33.6ʺS,
12º27ʹ41.4ʺE, 399 m (CAS 254767), 15º28ʹ31.7ʺS, 12º27ʹ43.9ʺE, 408 m (CAS 254763–253764);
RNN, 28 November 2013, 15º46ʹ22.5ʺS, 12º19ʹ57.7ʺE, 262 m (CAS 254951), 15º46ʹ25ʺS,
12º19ʹ54.9ʺE, 262 m (CAS 254953); Pico Azevedo, 7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE,
359 m (CAS 254939); Pediva Hot Springs, south side of the river, 2 December 2013, 16º17ʹ37.7ʺS,
13º33ʹ37.2ʺE, 235 m (CAS 254860), 16º17ʹ24.01ʺS, 12º33ʹ43.9ʺE, 270 m (CAS 254864); INP,
north of Tambor, 4 December 2013, 15º59ʹ43.4ʺS, 12º24ʹ23.3ʺE, 306 m (CAS 254840); Omauha
Lodge, 4 December 2013, 16º12ʹ1.2ʺS, 12º24ʹ00.1ʺE, 343 m (CAS 254835).
COMMENTS.— Pedioplanis haackei is one of the latest additions to the herpetofauna of Ango-
la, and is endemic to southern Angola. Each of our specimens has 10 to 12 longitudinal rows of
ventral scales, a semi-transparent lower eyelid with a brille formed of two large scales, five to six
supralabials anterior to the subocular, two rows of granules separating supraoculars from supracil-
iaries, and the typical coloration with dots on the flanks, concordant with the diagnostic characters
presented by Conradie et al. (2012) in the species description. Genetically, our specimens (Gen-
Bank accession numbers KU662311–KU662318) have an average 16S p-distance of 1% from
those of Conradie et al. (2012) (GenBank accession numbers HE794000, HE793999.1;
HE793998.1; HE793997.1; HE793996.1; HE793995.1; HE793994.1; HE793994.3).
BENGUELA SAND LIZARD
Pedioplanis benguellensis (Bocage, 1867)
MATERIAL.— Namibe-Lubango road, road marker 59, 1.8 km W (by road) from Caraculo,
north side of the road, 6 December 2013, 15º00ʹ57.8ʺS, 13º38ʹ41.4ʺE, 476 m (CAS 254909),
15º00ʹ58.8ʺS, 13º38ʹ33.8ʺE, 491 m (CAS 254906), 15º00ʹ57.5ʺS, 12º38ʹ38.3ʺE, 482 m (CAS
254896).
COMMENTS.— All of these specimens have ten to eleven longitudinal series of ventral plates
and a single transparent scale in the lower eyelid. However, they also have two rows of granules
separating supraoculars from supraciliaries, a character given by Conradie et al. (2012) as synapo-
morphic for P. haackei. The number of upper labials in front of the subocular is variable: CAS
254896 has three, CAS 254906 has five, and CAS 254909 has four. Molecular comparisons with
specimens of P. benguellensis from Conradie et al. (2012) show these specimens to be conspecif-
ic (average uncorrected 16S p-distance of 1% from our specimens [GenBank accession numbers
KU662319 to KU662321] from those of Conradie et al. [2012] [GenBank accession numbers
HE794014.1, HE794012.1, HE794011.1, HE794010.1 and HE794013.1]).
BUSHVELD LIZARD
Heliobolus lugubris (Smith, 1838)
MATERIAL.— Namibe-Lubango road, 2 km E of Mangueiras, south side of the road, 5 Decem-
ber 2013, 15º02ʹ37.0ʺS, 13º09ʹ36.0ʺE, 625 m (CAS 254885).
COMMENTS.— The species is widespread over much of southern Africa, particularly on sandy
substrates (Bauer et al. 1993). In Angola, the majority of both bibliographic (Bocage 1867, 1895;
Monard 1937; Mertens 1938; Boulenger 1921) and museum records (e.g., TM 46525) are from
Benguela. However, there are also records of the species for Namibe Province from “Maconjo”
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 37
(Bocage 1895), “Capangombe” (Bocage 1895), and “Konondoto” (Boulenger 1921). The species’
distribution in the country extends to the southeast in Huíla and Cunene Provinces. Our specimen
is a sub-adult (55.5 mm SVL).
Cordylidae
KAOKOVELD GIRDLED LIZARD
Cordylus namakuiyus Stanley, Ceríaco, Bandeira, Valério, Bates and Branch, 2016
MATERIAL.— 7.5 km NW (by road) of Pico Azevedo, 7 December 2013, 15º28ʹ33.2ʺS,
12º27ʹ45.7ʺE, 421 m (CAS 254754–254755, CAS 256530–256531); Namibe-Lubango road, road
marker 59, 1.8 km W (by road) from Caraculo, north side of the road, 6 December 2013,
15º00ʹ59.4ʺS, 12º38ʹ31.3ʺE, 503 m (CAS 254912–254914); 15º00ʹ57.3ʺS, 12º38ʹ32.6ʺE, 509 m
(CAS 256529).
COMMENTS.— Based on a combination of morphological and molercular data, Stanley et al.
(2016) described this new species endemic to the arid lowlands west of the southern Angolan
escarpment. The majority of the type material of C. namakuiyus was collected during this trip. The
new species is morphologically and genetically distinct from its sister taxon Cordylus machadoi
Laurent, 1964, which occurs in the highlands of Huila, not far from Namibe. In addition to an aver-
age 7.1% uncorrected p-distance for the mitochondrial marker ND2, the two species differ in the
extent of osteodermal armament. The more complete body armor of C. namakuiyus may be an
adaptation to the semi-arid and refuge-scarce habitat where the species occurs. Within the new
species there is also a degree of internal genetic structure, with specimens from Iona (PEM
R18005) and Pico Azevedo (CAS 254754, 254755, 256530, 256531) more closely related to one
another than to the two specimens from Caraculo. Specimens in the American Museum of Natural
Histiry identified as Cordylus cordylus and collected in 1925 during the Vernay Expedition in
Angola are assignable to C. namakuiyus. Although the Vernay specimens lack specific locality
information, the expedition field notes mention that significant numbers of unidentified lizards
were collected at “Pico Azevedo” and “100 km east of Moçâmedes” the same areas where we col-
lected eight specimens of C. namakuiyus.
DWARF PLATED LIZARD – Fig. 16
Cordylosaurus subtessellatus (Smith, 1844)
MATERIAL.— INP, 20 km
SSW of Espinheira, 30 Novem-
ber 2013, 16º55ʹ54.1ʺS,
12º14ʹ42.0ʺE, 631 m (CAS
254785).
COMMENTS.— The species
is known from the coastal areas
of Benguela Province (Bocage
1867, 1895; Boulenger 1887)
and from the Curoca River in
Namibe Province (Bocage 1895).
This specimen represents the
southernmost record for Angola.
The specimen was found hiding
38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 16. Adult Dwarf Plated Lizard, Cordylosaurus subtessellatus
(Smith, 1844), from near Espinheira. Photo by Edward Stanley.
in a granite boulder crevice. Extralimitally, the species is widely distributed from the Little Karoo
in the Western Cape of South Africa, north through the entire length of Namibia (Branch 1998).
Gerrhosauridae
DESERT PLATED LIZARD – Fig. 17
Gerrhosaurus skoogi (Andersson, 1916)
MATERIAL.— Praia do Navio coastal
dunes, ca 124 km SSW of Namibe, 8 December
2013, 16º16ʹ20.4ʺS, 11º49ʹ53.9ʺE, 8 m (CAS
254772), 16º16ʹ29.1ʺS, 11º49ʹ50.0ʺE, 8 m
(CAS 254774), 16º16ʹ42.1ʺS, 11º49ʹ21.7ʺE, 8
m (CAS 254777).
COMMENTS.— This species was encoun-
tered basking at the sun in the coastal dunes
SSW of Namibe, especially in dune valleys
areas. When approached, these lizards dive into
the sand, disappearing rapidly. The species is
easily identified by its unique morphology and
peculiar ecology. Sand trails resulting from the
specimens walking in the dunes were noted. A
total of five specimens were collected, three of
which are at CAS and two at INBAC. Several
additional animals were observed in the area
but not collected. Males have a distinct black
throat and venter, and are considerably larger
than the females. The only published Angolan
records are from the type locality, Porto Alexan-
dre, between Mossamedes and the mouth of the
Cunene River (Andersson 1916; Fitzsimons
1953), approximately in the same area as our
material. Although the species is the most dis-
tinctive of all Gerrhosaurus (Nance 2007), it is
unambiguously nested among more typical
taxa, implying its unique features are relatively recently derived autapomorphies (Lamb et al. 2003;
Lamb and Bauer 2013).
Varanidae
ANGOLAN ROCK MONITOR
Varanus albigularis angolensis Schmidt, 1933
MATERIAL.— 7.5 km NW (by road) of Pico Azevedo, adult, found in a rock crevice, 7 Decem-
ber 2013, 15º28ʹ33.6ʺS, 12º27ʹ41.4ʺE, 399 m (CAS 254768).
COMMENTS.— The subspecies was described from Gauca, Bihé (Bié) Province (Schmidt
1933). According to the original description, the subspecies differed from the nominotypic form by
having larger scales everywhere on the body so that the scales around the body are about 125
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 39
FIGURE 17. Adult female Desert Plated Lizard, Ger-
rhosaurus skoogi Andersson, 1916, from the coastal dunes,
near Praia do Navio. Photo by Arianna Kuhn
instead of 150, and the transverse rows of scales from the collar to thighs are 75 instead of 100
(Schmidt 1933). Laurent (1964b) noted that the morphological differences between the nomino-
typic form and angolensis are quite subtle and that angolensis may, in fact, be a synonym of albigu-
laris. However, Bayless (2002) considered the material from nearby localities, such as Bibala and
Caraculo as V. a. angolensis. Given the currently accepted distribution of angolensis and the lower
number of scales around the midbody and between the collar and the thighs, we tentatively identi-
fy our specimen as the Angolan subspecies. The subspecies appears to be the prevalent form in
Angola (and possibly extending to some neighboring regions of the DRC, even if the nominotyp-
ic form occurs sympatrically, especially in the southern regions of the country (Bayless 2002).
Pythonidae
SOUTHERN AFRICAN ROCK PYTHON – Fig. 18
Python natalensis Smith, 1840
MATERIAL.— Beginning of
the forested areas, at the start
of the climb to Leba Pass (by
road), near Bruco village, 5
December 2013, 15º07ʹ15.82ʺS,
13º11ʹ11.56ʺE. Individual
observed but not collected.
COMMENTS.— A local at a
site near Bruco village was sell-
ing a single live individual of
Python natalensis, presumably
collected nearby. In the province,
this species is known from
Maconjo (Bocage 1895;
Broadley 1984) and from Giraul
River (Bocage 1896; Broadley
1984). Python natalensis was for
many years considered as a sub-
species of Python sebae (Gmelin,
1789) (Broadley 1984), but was
elevated to specific status by
Broadley (1999) based on mor-
phological differences as well the
evidence of the overlapping dis-
tributions (Broadley and Cotterill
2004). Although the current taxo-
nomic arrangement appears
appropriate, molecular analyses are needed to conclusively resolve the relationship between
P. natalensis and P. sebae (Alexander 2007). Spawls and Branch (1995) and Bellosa et al. (2007)
provided maps with the distribution ranges for both species, with P. natalensis occurring in central
and southern Angola, as far north as the Kwanza River, overlaping in Luanda Province with
P. sebae, which occurs in northern regions of Angola, including Cabinda. According to the local
selling this individual, this species is sometimes collected for food or sold to tourists as pets.
40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 18. Adult Southern Rock Python, Python natalensis (Gmelin,
1788), being sold by a local near Bruco village. Photo by Luis Ceríaco.
Lamprophiidae
KAROO SAND SNAKE – Fig. 19
Psammophis notostictus Peters, 1867
MATERIAL.— Espinheira, 30 November 2013, 16º47ʹ13.8ʺS, 12º21ʹ27.5ʺE, 457 m (CAS
254827); Pico Azevedo, 7 December 2013, 15º32ʹ2.4ʺS, 12º29ʹ31.1ʺE, 359 m (CAS 254940).
COMMENTS.— Psammophis notostictus is easily recognizable from all other southern African
Psammophis by its single cloacal shield and the presence of two preoculars (Broadley 1975b, 1977,
2002). These two specimens have both of these diagnostic characters. The species is known for
Angola, but only from Namibe Province. The closest published records of the species are in Rio
São Nicolau (Loveridge 1940; Broadley 1975b, 2002), Moçamedes [Namibe city] (Bocage 1887;
Loveridge 1940), and Curoca River (Loveridge 1940; Broadley 2002). Our records expand the
known distribution of the species further south in the country, although it is continuous southwards
throughout much of western southern Africa (Branch 1998).
DISCUSSION
Namibe Province hosts a high diversity of reptile taxa, with approximately one-third of all the
reptile species known for Angola (see Table 1). Not surprisingly for an arid region, the diversity of
amphibians is considerably lower in this province. However, the anuran species Tomopterna
damarensis, adapted to drier climates, is reported here for the first time. The lizard families Scin-
cidae and Gekkonidae are the most species-rich groups for the province with 21 and 20
species/subspecies known, respectively, and for the snakes the family Lamprophiidae has the high-
est diversity of taxa, with 13 species known for the province. These numbers, however, are under-
estimates. We did not take into account unpublished voucher specimens; as noted above, a more
complete synopsis of the Namibe Province taxa will be provided elsewhere (W.R. Branch, pers.
comm.). There are several examples of taxa not previously recorded for Namibe, but which are
found both north and south of Namibe Province and can be expected to be found here in the
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 41
FIGURE 19 – Adult Karoo Sand Snake, Psammophis notostictus Peters, 1867, from Espinheira. Photo by Luis Ceríaco.
future.The presence of conspicuous species, such as Anchieta’s Dwarf Python, Python anchietae
Bocage, 1887, known from Benguela Province and from the northern regions of Namibia, is an
excellent example of this pattern. Biogeographically, the province is interesting because it repre-
sents the northern limit of several southern African habitats and species. The entry of the coastal
Namib Desert from Namibia into the southern coastal areas of the province as far north as the city
of Namibe, as well as the continuation of the Kaokoveld and the Namibian savanna woodlands pro-
vides a clear dispersal path for southern African taxa. Likewise, the so-called Pro-Namib extends
from northwestern Namibia, through Namibe, to the southern areas of Benguela Province. Despite
these similarities, the influence of the Cunene River as a barrier should not be underestimated.
There are a number of endemic Angolan species of widespread genera in southern Africa, such as
Pedioplanis haackei, P. huntleyi, and the undescribed species of Rhoptropus collected in our expe-
dition, as well as the endemic genus Kolekanos.
Given the concentration of reptile diversity in Namibe Province, conservation in this region is
of special concern. Of the 95 species of reptiles that occur in the province (Table 1), several have
small distributions within Namibe and ten are endemic, including the Slender Feather-tailed
Gecko, Kolekanos plumicaudus and the recently described Kaokoveld Girdled-Lizard, Cordylus
namakuiyus. As noted by Marques (2015), the majority of the amphibians and reptiles of Angola
(1) have not been accessed by the IUCN or are listed as Data Deficient, and (2) are known from
fewer than five published records for the country since the first studies published in early 1860s.
Roughly one third of Namibe Province, 19,600 km2out of 57,091 km2, is protected as either
national park or nature reserve. In contrast to more populated provinces, human activities in
Namibe that present significant threats to the herpetofauna are limited. The majority of the human
population is concentrated around Namibe and Tombwa, and the main economic activities are fish-
eries and traditional pastoralism. While there are no studies of the impact of livestock on the her-
petofauna within Namibe Province, negative impacts are known worldwide in other regions,
including South Africa (Bauer and Branch 2003; Fabricius et al. 2003; Smart et al. 2005). Yet the
low densities of livestock and the nomadic nature of the populations practicing pastoralism suggest
that this is not a major threat to the amphibians and reptiles of Namibe. In contrast to neighboring
Namibia (Herrmann and Branch 2013), there are no major mining activities in the province, even
if these activites show recent signs of increase, which may in the future threathen some species.
Other threats, such as climate change, are believed to have negative consequences on the distribu-
tion and abundance of southern African lizards (Erasmus et al. 2002; Bates et al. 2014). Extreme
climatic events can facilitate wildfires, which unambiguously affect the availability of important
habitats for reptiles (Meik et al. 2002). Lastly, for several taxa (e.g., pythons, chameleons,
varanids), the impact of human harvesting for food, traditional medicine and the pet trade should
not be overlooked. This is known from other African countries (Weldon et al. 2007; Alves et al.
2008; Segniagbeto et al. 2013), and we did encounter one instance of a python being sold during
our brief survey.
Additional faunal surveys are clearly still needed for Namibe Province. Even if this is one of
the most herpetofaunally well known provinces in Angola, the new species recently described sig-
nal that diversity likely remains underestimated, especially for groups of species that are similar in
external appearance. In addition to surveying new areas, it is also important to sample type locali-
ties of previously described species. Because many of the original topotypes were lost or destroyed,
especially material described by Bocage and originally housed in the Lisbon Museum (Ceríaco
2014), new topotypic material with associated genetic resources will help to address many taxo-
nomic issues, including for groups containing undescribed cryptic diversity. Angola sits at a cros-
sroads of southern and central Africa and is important to understanding phylogenetic and biogeo-
42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2
graphic patterns across sub-Saharan Africa. The specimens reported here were collected in the first
of an ongoing series of joint American-Angolan herpetofaunal expeditions that will build local
capacity within Angola and provide accessible data resources to the scientific and conservation
communities through georeferenced biodiversity informatics databases (e.g., GBIF.org,
Vertnet.org), molecular databases (GenBank), and digitized morphological resources (high-resolu-
tion x-ray CT-scans). Combining data from new field surveys with information from both museum
specimens and literature records (Marques et al., in prep.) will provide the first detailed picture of
Angola’s herpetofaunal diversity.
ACKNOWLEDGMENTS
We thank the Angolan Ministry of Environment and INBAC for providing the institutional
support and permits for this work. Soki Kuedikuenda, former director of INBAC provided the nec-
essary collecting and exporting permits and gave institutional support to the expedition. The Sec-
retary of State for Biodiversity, Paula Francisco, provided institutional support and guidance to the
expedition since its beginning. We also thank provincial and regional offices of Biodiversity Affairs
for all their support and cooperation. Alvaro Baptista and his family provided logistical help dur-
ing field work in Namibe. We also want to thank to the Methodist University of Angola, especial-
ly Teresa Silva Neto and Luis Sebastião for their support during our stay in Angola. Two anony-
mous reviewers provided thoughtful criticisms that gave us pause and that we believe led to an
improvement of the presentation. However, we accept full responsibility of all acts of commission
and/or omission that persist herein.
This work was partly funded by a US National Science Foundation grant to DCB and AMB
(DEB 1202609 and 1019443) and a grant to DCB and AMB from the JRS Biodiversity Founda-
tion.
LITERATURE CITED
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distribution géographique des oiseaux de l’Afrique occidentale. Bulletin de la Société Portugaise des
Sciences Naturelles 1:41–45.
SEGNIAGBETO, G.H., F. PETROZZI, A. AÏDAM, AND L. LUISELLI. 2013. Reptiles traded in the fetish market of
Lomé, Togo (West Africa). Herpetological Conservation and Biology 8(2):400–408.
SMART, R., M. J. WHITING, AND W. TWINE. 2005. Lizards and landscapes: integrating field surveys and inter-
views assess the impact of human disturbance on lizard assemblages and selected reptiles in a savanna
in South Africa. Biological Conservation 112(2005):23–31.
STANLEY, E.L., L.M.P. CERÍACO, S. BANDEIRA, H. VALERIO, M.F. BATES, AND W.R. BRANCH. 2016. A review of
Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the description of a new
species from the Pro-Namib desert. Zootaxa 4061:201–226.
THEMIDO, A.A. 1941. Répteis e batráquios das colónias Portuguesas (Catálogo das colecções do Museu Zoo-
lógico de Coimbra). Memórias e Estudos do Museu Zoológico da Universidade de Coimbra 119:1–28.
TILBURY, C. 2010. Chameleons of Africa, an Atlas including the Chameleons of Europe, the Middle East and
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WALLACH, C., AND J. BOUNDY. 2014. Snakes of the World: A Catalogue of Living and Extinct Species. CRC
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WELDON, V., K. A.L.D. VILLIERS, AND L.H.D. PREEZ. 2007. Quantification of the trade in Xenopus laevis from
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77–83.
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50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 51
Appendix
Table 1
52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2
Notes
1Because of the morphological similarity among many Tomopterna species, some older records might be referable to
this taxon.
2Records attributed in the literature to Tomopterna cryptotis, e.g., 25 km W of Virei (Poynton and Haacke 1993; Ruas
1996, 2001) and Miranda (Boulenger 1907a; Poynton and Haacke 1993; Ruas 1996, 2001) are likely referable to T. tandyi,
but this requires reexamination of the relevant material.
3Some records may be referable to L. bradfieldi Hewitt, 1932.
4Lygodactylus lawrencei Hewitt, 1926 is a rocky, dry savanna species from the Kaokoveld regions in northern Namib-
ia, extending into southern Angola (Pasteur 1964:70 [Fig. 18]; Branch 1998:247; Uetz and Hošek 2014). New surveys and
aquisition of new fresh material is absolutely required to clarify the extention range of L. lawrencei in Angola, given the
lack of literature records for this species in the country.
5Certainly in error, see Greer (1967).
6It is plausible that these records correspond to the subspecies angolensis. However, due to the destruction of the
specimens used by Bocage in the Lisbon Museum fire and the impossibility of confirming their identity, we opt to main-
tain the original identification.
7The taxonomy of this species is in flux and a revisionary work on the nigrolineatus complex is underway (D.G.
Broadley and M. Bates, pers. comm.).
8The species was described by Broadley and Schätti (1997: 172), from Namibia, near the Cunene River at Ruacana,
western Ovamboland. Bauer et al. (2001:75–76, 79) suggested that this species should be expected to occur in southern
Angola, due to the continuity of the mopaneveld habitat of the species on either side of the Cunene River.
9The species has an irregular distribution from eastern Zimbabwe and the Okavango Swampa, to Angola and Lake
Malawi, through Cameroon (Hughes 1985:519 [Fig. 11]; Branch 1998:94; Chirio and LeBreton 2007:518; Wallach et al.
2014:546).
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 53
Taxon Occurrences & References
AMPHIBIANS
ANURA
Pipidae
Genus Xenopus Wagler, 1827
Xenopus petersii Bocage, 1895 Mossamedes [= Namibe] (Bocage 1867; Ruas 1996, 2002).
Bufonidae
Genus Sclerophrys Tschudi, 1838
Sclerophrys gutturalis (Power, 1927)
Mossamedes [= Namibe] (Bocage 1867); Cima [= Giraul de Cima]
(Poynton and Haacke 1993; Ruas 1996, 2002); Saco do Giraul
(Poynton and Haacke 1993; Ruas 1996, 2002); Curoca (Poynton and
Haacke 1993; Ruas 1996, 2002).
Sclerophrys maculata (Hallowell,
1854)
Cainde (Poynton and Haacke 1993; Ruas 1996, 2002); 16 km W of
Vila Nova (Poynton and Haacke 1993, Ruas 1996, 2002); this study.
Sclerophrys garmani (Meek, 1897) Mossamedes [= Namibe] (Bocage 1895; Ruas 1996, 2001).
Genus Poyntonophrynus Frost, Grant, Falvovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson,
Donnellan, Raxworthy, Campbell, Biotto, Moler, Drewes, Nussbum, Lynch, Green and Wheeler, 2006
Poyntonophrynus dombensis (Bocage
1895) Assunção (Poynton and Haacke 1993; Ruas 1996).
Poyntonophrynus grandisonae (Poyn-
ton and Haacke 1993)
5 km E of Assunção (Poynton and Haacke 1993; Ruas 1996); Carac-
ulo (Poynton and Haacke 1993; Ruas 1996); Salona [= Saiona]
River, 2 km N of Cainde (Poynton and Haacke 1993; Ruas 1996); 20
km W of Virei (Poynton and Haacke 1993; Ruas 1996).
Microhylidae
Genus Phrynomantis Peters, 1867
Phrynomantis annectens Werner,
1910
Mutiambo River (Poynton and Haacke 1993; Ruas 1996); Caraculo
(Poynton and Haacke 1993; Ruas 1996); this study.
Brevicipitidae
Genus Breviceps Merrem, 1820
Breviceps adspersus Peters, 1882 Biballa [= Bibala] (Bocage 1895; Ruas 1996, 2002); Mossamedes
[= Namibe] (Bocage 1873b).
Hemisotidae
Genus Hemisus Wagler, 1827
Hemisus guineensis Cope, 1865 Mossamedes [= Namibe] (Bocage 1887b, 1895; Ruas 1996).
Arthroleptidae
Genus Leptopetlis Gunther, 1859
Leptopelis anchietae (Bocage, 1873) Mossamedes [= Namibe] (Bocage 1873b; Boulenger 1882;
Loveridge 1957).
Phrynobatrachidae
Genus Phrynobatrachus Gunther, 1862
Phrynobatrachus natalensis (Smith
1849) Cunene mouth (Poynton and Haacke 1993; Ruas 1996, 2002).
TABLE 1 – Only published records are included. Additional species known from the province based on
unpublished museum records are not included unless also supported by published records.
54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Pyxicephalidae
Genus Amietia Dubois, 1987
Amietia angolensis (Bocage, 1866) This study.
Genus Tomopterna Duméril and Bibron, 1841
Tomopterna damarensis Dawood and
Channing, 1999 This study1.
Tomopterna krugerensis Passmore
and Carruthers, 1975 Namibe (Channing 2001).
Tomopterna tandyi Channing and
Bogart, 1996 Namibe (Channing 2001)2.
Tomopterna tuberculosa (Boulenger,
1882)
Biballa [= Bibala] (Bocage 1895; Ruas 1996); bottom of Leba Pass
(Poynton and Haacke 1993; Ruas 1996).
REPTILES
TESTUDINES
Pelomedusidae
Genus Pelomedusa Wagler, 1830
Pelomedusa subrufa (Bonnaterre,
1789)
Mossamedes [= Namibe] (Bocage 1887, 1895; Loveridge 1941);
Mucungo (Shmidt 1933; Loveridge 1941); Maconjo (Bocage 1895;
Loveridge 1941); Capangombe (Bocage 1887, 1895; Loveridge
1941).
Testudinidae
Genus Kinixys Bell, 1827
Kinixys belliana Gray, 1830 Capangombe (Bocage 1895; Loveridge and Williams 1957).
Genus Stigmochelys Gray, 1873
Stigmochelys pardalis (Bell, 1828) Mossamedes [= Namibe] (Bocage 1895); Capangombe (Loveridge
and Williams, 1957).
SQUAMATA
Gekkonidae
Genus Afrogecko Bauer, Good and Branch, 1997
Afrogecko ansorgii (Boulenger,
1907). Maconjo (Boulenger 1907b; Bauer et al. 1997).
Genus Chondrodactylus Peters, 1870
Chondrodactylus fitzsimonsi
(Loveridge, 1947)
Praia das Conchas (Laurent 1964a); around Moçâmedes [= Namibe]
in the road to Sá da Bandeira [= Lubango] (Laurent 1964a); this
study.
Chondrodactylus pulitzerae (Schmidt,
1933)
Mossamedes [= Namibe] (Bocage 1867, 1895; Loveridge 1947;
Laurent 1964a); Pico Azevedo (Schmidt 1933; Barbour and
Loveridge 1946, 1947; Marx 1959); Curoca River (Bocage 1887;
Loveridge 1947); this study.
Genus Hemidactylus Oken, 1817
Hemidactylus longicephalus Bocage,
1873.
Capangombe (Bocage 1873b, 1895, 1897; Loveridge 1947); Curoca
Rriver (Bocage 1895, 1897; Loveridge 1947).
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 55
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Genus Kolekanos Heinicke, Daza, Greenbaum, Jackman and Bauer, 2014
Kolekanos plumicaudus (Haacke, 2008).
Tambor (Haacke 2008; Mashinini and Mahlangu 2013); Curoca
River (Haacke 2008); 11 km NE of Iona along track towards
Oncocua (Haacke 2008; Heinicke et al. 2014).
Genus Lygodactylus Gray, 1864
Lygodactylus capensis (Smith, 1849) Mucungu (Schmidt 1933; Loveridge 1947); Capangombe (Bocage
1895)3.
Lygodactylus lawrencei Hewitt, 1926 4
Genus Pachydactylus Wiegamnn, 1834
Pachydactylus angolensis Loveridge,
1944
Moçâmedes [= Namibe] to Sá da Bandeira [= Lubango] at Praia
das Conchas (Laurent 1964a); Lucira (Bauer 1999); San Nicolau
[= São Nicolau] (Bauer 1999); Lungo (Bauer 1999); Saco de
Giraul (Bauer 1999); this study.
Pachydactylus caraculicus FitzSimons,
1959
Lungo (FitzSimons 1959); Caraculo (FitzSimons 1959; Haacke
1970; Mashinini and Mahlangu 2013); Giraul de Cima, river
(FitzSimons 1959); 36 mi. northwest of Mocamedes [= Namibe]
(Bauer 1999).
Pachydactylus cf. oreophilus McLach-
land and Spence, 1967
Assuñcao (= Assunção) (Bauer 1999); Caraculo (Bauer 1999); 20
km W Virei (Bauer 1999); 6 km S of Coroca River towards Iona
(Bauer 1999); Saiona River, 25 km NW Cainde (Bauer 1999);
Mutiambo River on road to Lucira (Bauer 1999); Tambor (Bauer
1999); 7 km from Iona towards Oncocau, Iona Reserve (Bauer
1999); Furnas (Bauer 1999).
Pachydactylus punctatus Peters, 1854
60 km of the road of Moçâmedes [= Namibe] to Sá da Bandeira
(Laurent 1964a); Moçâmedes [= Namibe] (Laurent 1964a); 35 km
south of Moçâmedes [= Namibe] (Laurent 1964a); Pico Azevedo
(Schmidt 1933; Loveridge 1947); 11 mi NE of Mocamedes [=
Namibe] (Bauer and Branch 1995).
Pachydactylus rangei (Andersson,
1908)
Mossamedes [= Namibe] (Haacke 1976b); Curoca River (Haacke
1976b); Porto Alexandre (Haacke 1976b); Cunene mouth (Haacke
1976b); Lacrau (Haacke 1976b); Namib Desert (Mertens 1937).
Pachydactylus vanzyli (Steyn and
Haacke, 1966) Espinheira (Haacke 1976a); Kakolo windmill (Haacke 1976a).
Genus Rhoptropus Peters, 1869
Rhoptropus afer Peters, 1869
Maconjo (Bocage 1873b); Capangombe (Bocage 1873b, 1895,
1897b); Moçâmedes [= Namibe] (Boulenger 1885); Curoca River
(Bocage 1887b, 1895, 1897b).
Rhoptropus barnardi Laurent 1964a 60 km from Moçâmedes [= Namibe] to Sá da Bandeira [= Luban-
go] (Laurent 1964a); this study.
Rhoptropus biporosus Fitzsimons, 1957 Pico Azevedo (Bauer and Good 1996); this study.
Rhoptropus boultoni boultoni Schmidt,
1933
60 km from Moçâmedes [= Namibe] to Sá da Bandeira [= Luban-
go] (Laurent 1964a); Pico do Azevedo (Schmidt 1933; Mertens
1938; Barbour and Loveridge 1946; Marx 1959; McCoy and Rich-
mond 1966); this study.
56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & Referencess
Rhoptropus boultoni montanus Lau-
rent, 1964 This study.
Rhoptropus taeniostictus Laurent
1964a
Mucungu (Schmidt 1933); “60 km from Moçâmedes [= Namibe] to Sá
da Bandeira [= Lubango] (Laurent 1964a); this study.
Lacertidae
Genus Heliobolus Fitzinger, 1843
Heliobolus lugubris (A. Smith,
1838)
Maconjo (Bocage 1895); Capangombe (Bocage 1895); Konondoto
(Boulenger 1921); this study.
Genus Meroles Gray, 1838
Meroles anchietae (Bocage, 1867) Moçâmedes (= Namibe) (Bocage 1867, 1895, 1897; Boulenger 1887;
Loveridge 1936); this study.
Meroles reticulatus (Bocage, 1867)
Moçâmedes (= Namibe) (Bocage 1867; Boulenger 1887; Loveridge
1936); Coroca River (Bocage 1895, 1897; Boulenger 1887; Bauer and
Günther 1995; this study.
Genus Nucras Gray, 1838
Nucras tessellata (Smith 1838)
Maconjo (Bocage 1895; Broadley 1972); 34 km from Moçâmedes [=
Namibe] to Sá da Bandeira [= Lubango] (Laurent 1964; Broadley
1972).
Genus Pedioplanis Fitzinger, 1843
Pedioplanis benguellensis (Bocage,
1867)
Maconjo (Boulenger 1921; Bauer and Günther 1995; Conradie et al.
2012); Capangombe (Bocage 1895; Monard 1937); Mossamedes [=
Namibe] (Bocage 1887, 1895; Monard 1937); this study.
Pedioplanis haackei Conradie,
Measey, Branch and Tolley, 2012
Red Canyon at Lake Arco (Conradie et al. 2012); 10 km south of Lake
Arco (Conradie et al. 2012); Road to Tambor at giant Welwitchia (Con-
radie et al. 2012); Road from Lake Arco to Espinheira (Conradie et al.
2012); Omauha Lodge (Conradie et al. 2012); Road to Tambor (Con-
radie et al. 2012); 20 km north of Omauha Lodge (Conradie et al.
2012); this study.
Pedioplanis huntleyi Conradie,
Measey, Branch and Tolley, 2012
Omauha Lodge (Conradie et al. 2012). 14 km west of Moimba (Con-
radie et al. 2012); 23 km west of Moimba (Conradie et al. 2012); 26
km east of Iona (Conradie et al. 2012); 16 km east of Iona (Conradie et
al. 2012); 8 km northeast of Iona (Conradie et al. 2012); Road to
Onocua 7 km NE from Iona (Conradie et al. 2012); 26 km SE of
Onocua (Conradie et al. 2012).
Scincidae
Genus Eumecia Bocage, 1870
Eumecia anchietae Bocage, 1870 Moçâmedes [= Namibe] (Boulenger 1887)5.
Genus Mochlus Günther, 1864
Mochlus sundevallii (Smith, 1849)
Campangombe (Bocage 1895); Moçâmedes (= Namibe) (Bocage 1867;
1895); Curoca River (Bocage 1895); 10mls E of Caracul [= Caraculo]
(Haacke 1965; Broadley 1966).
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 57
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Genus Panaspis Cope, 1868
Panaspis cabindae (Bocage, 1866) Capangombe (Bocage 1895, 1897).
Genus Sepsina Bocage, 1866
Sepsina angolensis Bocage, 1866 Capangombe (Bocage 1895; Monard 1937).
Sepsina copei Bocage, 1873 Biballa [= Bibala] (Bocage 1895, 1897).
Genus Trachylepis Fitzinger, 1843
Trachylepis acutilabris (Peters, 1862)
Cahinde-Ongueira (Hellmich 1957a); Mossâmedes [= Namibe]
desert, 35 km south from the city (Laurent 1964a); Curoca River
(Bocage 1895); this study.
Trachylepis bayonii bayonii (Bocage,
1872) Moçâmedes [= Namibe] (Boulenger 1887).
Trachylepis binotata (Bocage, 1867)
Capangombe (Bocage 1895); 50 km Moçâmedes [= Namibe] to Sá
da Bandeira [= Lubango]" (Laurent 1964a); Maconjo (Bauer et al.
2003).
Trachylepis chimbana (Boulenger,
1887)
Chimba River (Boulenger 1887; Bocage 1872, 1895, 1897;
Broadley 1975a); Assunção (Broadley 1975a); Maconjo (Bocage
1895, 1897; Broadley 1975a), Moçâmedes [= Namibe] (Boulenger
1887); Lucira (Broadley 1975a); Chapeau Armando [= Chapéu
Armando] turnoff, Mossamedes [= Namibe] (Broadley 1975a);
Caraculo (Broadley 1975a); 14 km NE Caraculo (Broadley 1975a);
Capangombe (Bocage 1895, 1897; Broadley 1975a); Mossamedes
[= Namibe] (Boulenger 1887); Saiona River, NW of Cainde
(Broadley 1975a); Cainde (Broadley 1975a); Coroca River
(Broadley 1975a).
Trachylepis hoeschi (Mertens, 1954) Praia das Conchas, Moçâmedes [= Namibe] (Laurent 1964a); this
study.
Trachylepis lacertiformis (Peters, 1854) Cainde (Broadley 1975); 14 km NE of Caraculo (Broadley 1975a).
Trachylepis laevis (Boulenger, 1907) Maconjo (Boulenger 1907b); Munhino (Laurent 1964a); this study.
Trachylepis occidentalis (Peters, 1867) Curoca River (Bocage 1895), 35 km south of the city of
Moçâmedes [= Namibe] (Laurent 1964a); this study.
Trachylepis punctulata (Bocage, 1872)
Inamango River on Lucira road (Broadley 1975a); Mucungu
(Schmidt 1933; Broadley 1975a); Sao Nicolau [= São Nicolau]
(Broadley 1975a); 17 km N of Sao Nicolau [= São Nicolau]
(Broadley 1975a); Caraculo (Broadley 1975a); 15 km W of Carac-
ulo (Broadley 1975a); Cima [= Giraul de Cima] (Broadley 1975a);
Pico Azevedo (Broadley 1975a); 23 km W of Virei (Broadley
1975a); Coroca River (Bocage 1872, 1895, 1897; Boulenger 1887;
Broadley 1975a, 2000); 6 km S of Rio Coroca on Iona road
(Broadley 1975a); Porto Alexandre [= Tômbua] (Broadley 1975a);
30 km N of Tambor (Broadley 1975a); Octchinfengo River on
Onocua road, Iona Reserve (Broadley 1975a); Cunene mouth
(Broadley 1975a); this study.
Trachylepis sulcata (Peters, 1867) Capangombe (Bocage 1895); Munhino (Laurent 1964a); this study.
58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Trachylepis varia (Peters, 1867) Moçâmedes [= Namibe] (Bocage 1867); Biballa [= Bibala]
(Bocage 1872); this study.
Genus Typhlacontias Bocage, 1873
Typhlacontias johnsonii Anderson, 1916 Curoca River (Bocage 1895; Haacke 1997); Porto Alexandre (=
Tombwa) [= Tômbua] (Haacke 1997); Lacrau (Haacke 1997).
Typhlacontias punctatissimus punctatis-
simus Bocage, 1873
Coroca River (Bocage 1873, 1887 1895, 1897); Mossamedes [=
Namibe] (Boulenger 1887); Porto Alexandre [= Tômbua] (Haacke
1997).
Typhlacontias punctatissimus bogerti
Laurent, 1964
Mossamedes [= Namibe] desert, 35 km south from the city (Lau-
rent 1964a); Curoca River (Haacke 1997); Moçâmedes [= Namibe]
(Haacke 1997); 10 km S of Moçâmedes [= Namibe] (Haacke
1997); 34 km S of Moçâmedes [= Namibe] (Haacke 1997); 8 km
SE of Pico Azevedo (Haacke 1970); Kakolo windmill (Haacke
1997); this study.
Typhlacontias rudebecki Haacke, 1997 São Nicolau (Haacke 1997).
Varanidae
Genus Varanus Merrem, 1820
Varanus albigularis angolensis
Schmidt, 1933 This study.
Varanus albigularis albigularis
(Daudin, 1802) Chimba River (Bocage 1895); Biballa [= Bibala] (Bocage 1895)6.
Chamaeleonidae
Genus Chamaeleo Laurenti, 1768
Chamaeleo anchietae Bocage, 1872 Namibe (Tilbury 2010: 451).
Chamaeleo dilepis Leach, 1819 Moçâmedes [= Namibe] (Bocage 1867b; 1887; 1895); Chimba
(Hellmich 1957a).
Chamaeleo namaquensis Smith, 1831 Moçâmedes [= Namibe] (Günther 1865; Bocage 1867, 1872, 1895;
Boulenger 1887).
Agamidae
Genus Agama Daudin, 1802
Agama aculeata Merrem, 1820
Biballa [= Bibala] (Bocage 1895); Moçâmedes [= Namibe]
(Bocage 1887b, 1895); Virei-Cahinde (Hellmich 1957a); Molundo
(Monard 1937); Chimporo (Monard 1937).
Agama anchietae Bocage, 1896
Moçâmedes [= Namibe] (Bocage 1896, 1897); 100 km southeast of
Moçâmedes [= Namibe] (Laurent 1964); Maconjo (Boulenger and
Power 1921); this study.
Agama planiceps planiceps Peters, 1862 Biballa [= Bibala] (Bocage 1895); Pico Azevedo (Schimdt 1933);
this study.
Cordylidae
Genus Cordylus Laurenti, 1768
Cordylus namakuiyus Stanley, Ceríaco,
Bandeira, Valério, Bates and Branch,
2015
Caraculo, on the road from Lubango and Namibe (Stanley et al.
2016); Pico Azevedo (Stanley et al. 2016); road between Namibe
and Omauha Lodge (Stanley et al.2016); this study.
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 59
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Gerrhosauridae
Genus Cordylosaurus Gray, 1865 [1866]
Cordylosaurus subtessellatus (Smith,
1844) Curoca River (Bocage 1895); this study.
Genus Gerrhosaurus Wiegmann, 1828
Gerrhosaurus nigrolineatus Hallowell,
1857 Capangombe (Bocage 1895; Loveridge 1942)7.
Gerrhosaurus skoogi Andersson, 1916 Porto Alexandre [= Tômboa] (FitzSimons 1953); this study.
Genus Matobosaurus Bates and Tol-
ley, 2013
Matobosaurus maltzahni (De Grys,
1938)
Chimba River (Bocage 1895); Moçâmedes [= Namibe] (Bocage
1895); Tambor (Bates et al. 2013); Omauha Lodge (Bates et al.
2013).
SERPENTES
Typhlopidae
Genus Afrotyphlops Broadley and Wallach, 2009
Afrotyphlops anomalus (Bocage, 1873) Biballa [= Bibala] (Bocage 1895; Broadley and Wallach 2009);
Moçâmedes [= Namibe] (Bocage 1873).
Afrotyphlops schlegelii (Bianconi,
1847)
Biballa [= Bibala] (Bocage 1873, 1886, 1895, 1897a; Loveridge
1933).
Leptotyphlopidae
Genus Leptotyphlops Fitzinger, 1843
Leptotyphlops scutifrons (Peters, 1854) Biballa [= Bibala] (Bocage 1873, 1895), Capangombe (Bocage
1895).
Pythonidae
Genus Python Daudin, 1803
Python natalensis Smith, 1840 Maconjo (Bocage 1895; Broadley 1984); Giraul River (Bocage
1896; Broadley 1984); this study.
Viperidae
Genus Bitis Gray, 1842
Bitis arietans (Merrem, 1820) Moçâmedes (= Namibe) (Günther 1865; Manaças 1981).
Bitis caudalis (Smith, 1839)
Capangombe (Bocage 1895, Manaças 1981); Moçâmedes
[= Namibe] (Günther 1865; Monard 1937; Manaças 1981);
Mossamedes [= Namibe] Desert, 35 km south from the city (Lau-
rent 1964a); Curoca River (Bocage 1895; Monard 1937; Manaças
1981).
Genus Causus Wagler, 1830
Causus resimus (Peters, 1862)
Chimba River (Bocage 1895; Manaças 1981); Biballa [= Bibala]
(Bocage 1895; Manaças 1981); Maconjo (Bocage 1895); Macujo [=
Maconjo] (Manaças 1981).
Causus rhombeatus (Lichtenstein,
1823)
Cuce River (Ferreira 1897); Mossamedes [= Namibe] (Bocage
1887a, 1895).
60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Series 4, Volume 63, No. 2
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Lamprophiidae
Genus Aparallactus Smith, 1849
Aparallactus capensis capensis Smith,
1849
Biballa [= Bibala] (Bocage 1895, 1897a; Loveridge 1944b; de
Witte and Laurent 1947).
Genus Boaedon Duméril, Bibron and
Duméril, 1854
Boaedon fuliginosus complex Biballa [= Bibala] (Bocage 1895); Capangombe (Bocage 1895).
Genus Lycophidion Fitzinger, 1843
Lycophidion hellmichi Laurent, 1964 Capolopopo (Laurent 1964a; Broadley 1991, 1996).
Lycophidion multimaculatum Boettger,
1888
Moçâmedes [= Namibe] (Bocage 1895; Laurent 1968; Broadley
1996).
Genus Hemirhagerrhis Boettger, 1896
Hemirhagerrhis viperina (Bocage,
1873)
Cuce River (Ferreira 1897); Munhino (Bogert 1940; Broadley
1995b; Broadley and Hughes 2000); Maconjo (Bocage 1895;
Broadley 1995b; Broadley and Hughes 2000); Capangombe
(Bocage 1895; Broadley 1995b; Broadley and Hughes 2000); Huxe
(Broadley 1997b; Broadley and Hughes 2000); Lungo (Broadley
1997b; Broadley and Hughes 2000); Caraculo (Broadley 1997b;
Broadley and Hughes 2000).
Genus Psammophis Boie, 1825
Psammophis leopardinus Bocage, 1887
Capangombe (Bocage 1887; Broadley 2002); Moçâmedes
[= Namibe] (Bocage 1887, 1895; Loveridge 1957); Iona (Broadley
2002; Hughes and Wade 2002).
Psammophis mossambicus Peters, 1882
Mossamedes [= Namibe] (Loveridge 1940; Broadley 2002); Coroca
River (Bocage 1895); Porto Alexandre [= Tômboa] (Loveridge
1940).
Psammophis namibensis Broadley,
1975
Mossamedes [= Namibe] (Broadley 1975b, 2002b); Pico Azevedo
(Broadley 2002); Coroca River (Broadley 1975b, 2002); Cunene
mouth (Broadley 1975b, 2002b); Cunene Forde, 15 km NE, Iona
Res. (Broadley 2002).
Psammophis notostictus Peters, 1867
São Nicolao River (Boulenger 1896; Loveridge 1940; Broadley
1975b; 2002); Coroca River (Bocage 1887, 1895; Monard 1937;
Loveridge 1940; Broadley 1975b, 2002); this study.
Psammophis subtaeniatus Peters, 1882
Biballa [= Bibala] (Bocage 1895, 1897a; Loveridge 1940); Macon-
jo (1895, 1897a; Loveridge 1940; Broadley 2002); Chao de Chella
(Broadley 2002).
Psammophis trigrammus Günther,
1865
Sao Nicolao [= São Nicolau] River (Bocage 1887; Loveridge 1940;
Broadley 1977, 2002; Wallach et al. 2014); Catara River (Broadley
2002); Coroca River (Broadley 2002b); Iona Reserve, 7 km to
Oncócua (Broadley 2002).
Genus Prosymna Gray, 1849
Prosymna frontalis (Peters, 1867) Moçâmedes [= Namibe] (Boulenger 1893).
Prosymna visseri Fitzsimons, 1959
near Caracul [= Caraculo], S. Angola (Fitzsimons 1959; Bauer et al.
2001); Balabaia (Broadley 1980); 5 km S of Chibemba (Broadley
1980).
CERÍACO ET AL.: AMPHIBIANS AND REPTILES OF WESTERN ANGOLA 61
TABLE 1 (continued) – Only published records are included. Additional species known from the province
based on unpublished museum records are not included unless also supported by published records.
Taxon Occurrences & References
Elapidae
Genus Afronaja Wallach, Wüster and
Broadley 2009
Afronaja mossambica (Peters, 1854) Maconjo (Manaças 1981; Broadley 1974).
Afronaja nigricincta (Bogert, 1940)
Munhino (Bogert 1940; Manaças 1981); Maconjo (Manaças
1981); Capangombe (Bocage 1895); Cunene mouth (Manaças
1981).
Afronaja nigricollis (Reinhardt, 1843) Capangombe (Ferreira 1900b; Manaças 1981).
Genus Aspidelaps A. Smith, 1849
Aspidelaps lubricus cowlesi Bogert,
1940
Munhino (Bogert 1940; Manaças 1981; Broadley and Baldwin
2003).
Genus Boulengerina Dollo, 1886
Boulengerina melanoleuca Hallowell,
1857 Capangombe (Ferreira 1900b; Manaças 1981).
Genus Elapsoidea Bocage, 1866
Elapsoidea semiannulata semiannulata
Bocage, 1882
Maconjo (Bocage 1895, 1897c; Loveridge 1944b; Broadley 1971,
1998b; Manaças 1981).
Colubridae
Genus Coluber Linnaeus, 1758
Coluber zebrinus Broadley and Schätti,
1997 8
Genus Crotaphopeltis Fitzinger, 1843
Crotaphopeltis hotamboeia (Laurenti,
1768) Biballa [= Bibala] (Bocage 1895).
Genus Philothamnus Smith, 1840
Philothamnus angolensis Bocage 1882 Capangombe (Bocage 1882a, 1897a; Loveridge 1951, 1957;
Chirio and LeBreton 2007; Wallach et al. 2014).
Philothamnus irregularis (Leach, 1819) Capangombe (Bocage 1882a, 1887, 1895); Mossamedes [=
Namibe] (Bocage 1887, 1895).
Philothamnus ornatus Bocage, 1872 9
Philothamnus semivariegatus (Smith,
1840)
Maconjo (Bocage 1882a); Capangombe (Bocage 1895; Haacke
1985).
Genus Telescopus Wagler, 1830
Telescopus finkeldeyi Haacke, 2013 5 km north Namibé [= Namibe] (Haacke 2013).
Genus Thelotornis A. Smith, 1849
Thelotornis capensis oatesi (Günther,
1881) Biballa [= Bibala] (Bocage 1895; Loveridge 1944b).
... In order to address these shortcomings, national inventories have been published to help summarise all the historical materials from Angola (Branch 2018;Marques et al. 2018;Baptista et al. 2019b;Branch et al. 2019c) and have set the foundation for future herpetological work. Several formal conservation areas, including national parks and reserves, have been the focus of herpetological surveys in the last decade (Ceríaco et al. 2014(Ceríaco et al. , 2016a(Ceríaco et al. , 2018bConradie et al. 2016Butler et al. 2019;Baptista et al. 2019a;Ernst et al. 2020). The rapid improvement of our knowledge of Angolan herpetofauna, especially the number of new species descriptions, highlights the need to re-evaluate estimates of biodiversity richness across the country . ...
... Numerous herpetologists have visited the region in recent years, and this has led to the description of five new species from the park and immediate surroundings (Haacke 2008;Conradie et al. 2012b;Branch et al. 2021;Parrinha et al. 2021) and the documentation of several range restricted species, such as Pachydactylus scutatus Hewitt 1927, P. rangei (Andersson, 1908), P. vanzyli (Steyn & Haacke, 1966), and Lygodactylus lawrencei Hewitt 1927. To date, 37 herpetofaunal species (within four amphibian and 15 reptile genera) are documented to occur in extreme southwestern Angola including both Iona NP and surrounds, which includes the Namibe Partial Reserve to the north (Ceríaco et al. 2016a). More broadly, 14 species of amphibians and over 95 species of terrestrial reptiles have been recorded for the entirety of Namibe Province (Marques et al. 2018), which includes Iona NP. ...
... In addition to these new survey data, distributional information from existing herpetological records were retrieved from the literature (Ceríaco et al. 2016a;Branch et al. 2017;Marques et al. 2018;Baptista et al. 2019;Marques et al. 2020;. After consolidation of new and existing records, all new records were examined to assess whether they represented extensions to the known range. ...
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Angola has experienced an incredible increase of the knowledge of its herpetofauna over the past decade. However, accurate biodiversity inventories remain deficient for certain regions of particular conservation interest. We therefore provide an updated checklist of Iona National Park’s herpetofauna, with 75 recorded species, including five amphibians and 70 reptiles, 40 of these recorded for the first time in Iona National Park. These species comprise ~80% of the reptile diversity of Namibe Province. Therefore, this work recognises Iona National Park as the most reptile-diverse protected area in Angola and is potentially one of the richest in southern Africa. Consequently, this work enhances the importance of specific conservation plans in the area and the need for further investigation into the hidden biodiversity of this region.
... Chondrodactylus fitzsimonsi is restricted to the western regions of northern Namibia and adjacent southern Angola (Fig. 12, right). In Namibe Province, Angola it is found south of Moçâmedes (formerly Namibe) to the Namibian border in Iona National Park (Ceríaco et al., 2016;Marques et al., 2018) with a single record from Ongueria, just above the escarpment in Huíla Province (Laurent, 1964). The distribution commonly reported for the species (e.g., Branch, 1998) extends southward in the Kunene Region of Namibia at least as far as the Grootberg Pass and the type locality at Kamanjab. ...
... In fact, only C. laevigatus occurs in this region. Ceríaco et al. (2016) noted the presence of C. pulitzerae in far northwestern Namibia but did not provide details. It occurs from the border at the Kunene River south as far as Sesfontein (entire Namibian distribution in the Kunene Region) with a single locality further south at ''10 km N of the Hunkab River'' (but unknown where along the river, TM 52910-11). ...
... It was regarded as a valid subspecies of P. laevigatus by Benyr (1995) and as a full species of Chondrodactylus by Heinz (2011) in their respective unpublished theses. Ceríaco et al. (2014) first used C. pulitzerae as a specifically valid name on the basis of Heinz's (2011) molecular data, and it has since been used consistently (e.g., Ceríaco et al., 2016Ceríaco et al., , 2017Conradie et al., 2016;Heinicke et al., 2017;Marques et al., 2018;Branch et al., 2019). ...
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Despite being among the largest and most conspicuous geckos across southern and eastern Africa, the toe-padded species of Chondrodactylus have remained one of the most taxonomically difficult groups of African lizards, due chiefly to their overall morphological conservativeness accompanied by high intraspecific variation. Current recognition of taxa is based on recent molecular phylogenetic analyses, but the application of the currently recognized nomina to particular populations has not yet been presented. We present a much-expanded multigene analysis of 234 representatives of the genus Chondrodactylus that supports the recognition of 6 species-level taxa, one without toepads, C. angulifer, as sister to five with pads: C. bibronii, C. turneri, C. laevigatus, C. pulitzerae, and C. fitzsimonsi. In general, the species can be recognized on the basis of the relative size of chin and gular scales, dorsal scalation, and head shape. However, the most widespread species, C. laevigatus is only very subtly distinct from C. turneri, with which it is likely parapatric in East Africa (although western populations of C. laevigatus are unambiguously diagnosable from all other congeners). Intraspecific divergences are high in some of the species. In C. fitzsimonsi there is evidence of shared nuclear haplotypes with C. pulitzerae and potential morphological evidence for hybridization or introgression with C. laevigatus. Chondrodactylus turneri exhibits a mitochondrial gene rearrangement that is unique among all geckos followed by an insertion of roughly 200 base pairs that do not correspond to known sequences. Most Chondrodactylus species are primarily distributed in arid to semiarid southwestern Africa, where as many as 4 species occur in sympatry in northern Namibia. In contrast, C. turneri is limited to the lowlands of the southeast and C. laevigatus follows the arid-corridor traversing sub-Saharan Africa southwest to northeast.
... In this period, the country's biodiversity was neglected and over-exploited for its natural resources (Huntley and Matos 1992). Following the cessation of hostilities and the ongoing redevelopment of regional infrastructure, the modern biodiversity surveys have begun to focus on poorly surveyed regions of the country in the south-west (Huntley 2009;Ceríaco et al. 2016a;Baptista et al. 2018Baptista et al. , 2019aButler et al. 2019), northeast (Branch and Conradie 2015;Huntley and Francisco 2015), south-east (Brooks 2012(Brooks , 2013Conradie et al. 2016;NGOWP 2017), central (Ceríaco et al. 2014(Ceríaco et al. , 2016b(Ceríaco et al. , 2018a, and north-west regions. Some of these surveys have targeted areas that had never been scientifically surveyed until recently, leading to the discovery and description of several new species of amphibians (Conradie et al. 2012aCeríaco et al. 2018bCeríaco et al. , 2021Nielsen et al. 2020) and reptiles (Conradie et al. 2012bStanley et al. 2016;Branch et al. 2019Branch et al. , 2021Marques et al. 2019Marques et al. a,b, 2020Ceríaco et al. 2020a-c;Hallermann et al. 2020;Parrinha et al. 2021). ...
... Some of these surveys have targeted areas that had never been scientifically surveyed until recently, leading to the discovery and description of several new species of amphibians (Conradie et al. 2012aCeríaco et al. 2018bCeríaco et al. , 2021Nielsen et al. 2020) and reptiles (Conradie et al. 2012bStanley et al. 2016;Branch et al. 2019Branch et al. , 2021Marques et al. 2019Marques et al. a,b, 2020Ceríaco et al. 2020a-c;Hallermann et al. 2020;Parrinha et al. 2021). Other targeted surveys resulted in the addition of new country records Conradie and Bourquin 2013;Ernst et al. 2014Ernst et al. , 2015Ceríaco et al. 2014Ceríaco et al. , 2016aBranch 2018) or resolved previous taxonomic confusion (Channing et al. 2013;Channing and Baptista 2013;Ernst et al. 2015;Marques et al. 2018). Recent synopses of the Angolan herpetofauna include an atlas of all the herpetofauna of Angola (Marques et al. 2018), a checklist of snakes (Branch 2018), and overviews of the reptiles ) and amphibians (Baptista et al. 2019b) of Angola. ...
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The herpetofauna of Angola has been neglected for many years, but recent surveys have revealed previously unknown diversity and a consequent increase in the number of species recorded for the country. Most historical surveys focused on the north-eastern and south-western parts of the country, while mostly neglecting the central and south-eastern parts, comprising the provinces of Bié, Moxico, and Cuando Cubango. To address this sampling bias and investigate the conservation importance of the region, a series of rapid biodiversity surveys of the upper Cuito, Cubango, Cuando, Zambezi, and Cuanza river basins were conducted by the National Geographic Okavango Wilderness Project between 2015 and 2019. The first part of those survey results are presented here as an updated checklist of current and historical snake records from the south-eastern region of Angola. In summary, 160 new specimens were collected from the region, comprising 36 species, bringing the total number of recognized snake species for the region to 53. These surveys revealed three new country records (Amblyodipsas ventrimaculata, Crotaphopeltis barotseensis, and Dasypeltis confusa) and led to the description of two novel species in previous publications (Boaedon branchi and B. fradei), increasing the total number of snake species in Angola to 133. Finally, updated geographic distribution maps are provided for all species encountered in this study for the whole country. This contribution increases our knowledge of this poorly known region of Africa and highlights the need for and importance of similar studies in other undersampled areas.
... Recently, a fifth site at Omauha farm was added to the species' distribution (Agarwal et al., 2017). Various herpetological surveys conducted in Iona NP and across the Angolan province of Namibe failed to record the species elsewhere (Ceríaco et al., 2016;Marques et al., 2018;Branch et al., 2019), strongly suggesting a highly restricted distribution range. In all these cases the species was found associated with granite outcrops overlaying a sandy substrate at mid-elevations below 800 m elevation. ...
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This work provides an updated distribution for K. plumicaudus within Namibe Province, the known range now spanning ~240 km from north of the city of Moçamedes southwards to the margins of the Cunene River on the Namibian border. Furthermore, using the GeoCAT IUCN tool (following Bachman et al., 2011) we update the EOO to 11,333 km2, which almost triples the previous estimates (Ceríaco et al., 2020; Fig. 1). These results support the hypothesis of K. plumicaudus being a species characteristic of the Angolan Kaokoveld within the Namib Desert system, but also suggest that they may be marginally present in north-western Namibia near the Cunene River at the edge of the sand sea, a possibility that should be investigated further in future works. Our new findings highlight some key habitat requirements of this species: sparse bushes with thin branches for nocturnal foraging, in close proximity to rock formations with deep, narrow crevices (often formed by flaked weathering patterns) for diurnal shelter. While it was previously assumed to occur only on large granite boulders, smaller outcrops of quartzite or other formations with similar weathering patterns are also suitable for this species. We further demonstrate a wide elevational tolerance, ranging from below 100 m to above 2,000 m elevation. Finally, we provide the first evidence of fluorescence in a leaf-toed gecko, adding to at least three other distantly related species of geckos in the Namib Desert who are known to fluoresce, suggesting that this feature may have evolved independently in different lineages in this environment.
... In the last two decades, Angolan herpetological knowledge has greatly improved, thanks to enhanced social stability and accessibility to previously unexplored regions, which helped to unearth this incredible local biodiversity (Marques et al. 2018;). These efforts have led to the description of new species (Haacke 2008;Conradie et al. 2012Conradie et al. , 2020Stanley et al. 2016;Marques et al. 2019aMarques et al. ,b, 2020Branch et al. 2019bBranch et al. , 2021Ceríaco et al. 2020a,b,c;Hallerman et al. 2020), the rediscovery of several poorly known species, and documenting new range extensions and additions to the country's species list (Branch and Conradie 2013;Conradie and Bourquin 2013;Ceríaco et al. 2014;Ernst et al. 2014Ernst et al. , 2015Ceríaco et al. 2016;Branch 2018;Branch et al. 2019a;Vaz Pinto et al. 2019;Baptista et al. 2019Baptista et al. , 2020Ernst et al. 2020Ernst et al. , 2021. ...
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The taxonomy of Angolan Hemidactylus has recently been revised. However, the lack of fresh material for some groups and regions, has led to the misidentification of some taxa and an underestimation of actual diversity in others. To shed light on the evolutionary history and systematics of Angolan Hemidactylus, we generated a new phylogenetic hypothesis for the group, and updated the tax-onomy following an integrative approach. This resulted in the description of four new species (H. pfindaensis sp. nov., H. faustus sp. nov., H. carivoensis sp. nov. and H. cinganji sp. nov.), the reevaluation of two recently described species (H. vernayi and H. paivae) and the synonymization of a recently described species (H. hannahsabinnae). We estimate divergence times for these lineages, providing a preliminary interpretation of their speciation process. Moreover, we suggest and outline 13 Angolan Main Biogeographic Units (AMBUs) in the area, defining a new biogeographic context for future works on Angolan herpetofauna. We now recognize eleven Hemidactylus species in Angola, and we provide here a new morphological key for Angolan Hemidactylus to help with identifications and species assignments in this group.
... In the last two decades, Angolan herpetological knowledge has greatly improved, thanks to enhanced social stability and accessibility to previously unexplored regions, which helped to unearth this incredible local biodiversity (Marques et al. 2018;). These efforts have led to the description of new species (Haacke 2008;Conradie et al. 2012Conradie et al. , 2020Stanley et al. 2016;Marques et al. 2019aMarques et al. ,b, 2020Branch et al. 2019bBranch et al. , 2021Ceríaco et al. 2020a,b,c;Hallerman et al. 2020), the rediscovery of several poorly known species, and documenting new range extensions and additions to the country's species list (Branch and Conradie 2013;Conradie and Bourquin 2013;Ceríaco et al. 2014;Ernst et al. 2014Ernst et al. , 2015Ceríaco et al. 2016;Branch 2018;Branch et al. 2019a;Vaz Pinto et al. 2019;Baptista et al. 2019Baptista et al. , 2020Ernst et al. 2020Ernst et al. , 2021. ...
Article
Full-text available
The taxonomy of Angolan Hemidactylus has recently been revised. However, the lack of fresh material for some groups and regions, has led to the misidentification of some taxa and an underestimation of actual diversity in others. To shed light on the evolutionary history and systematics of Angolan Hemidactylus , we generated a new phylogenetic hypothesis for the group, and updated the taxonomy following an integrative approach. This resulted in the description of four new species ( H. pfindaensis sp. nov. , H. faustus sp. nov. , H. carivoensis sp. nov. and H. cinganji sp. nov. ), the reevaluation of two recently described species ( H. vernayi and H. paivae ) and the synonymization of a recently described species ( H. hannahsabinnae ). We estimate divergence times for these lineages, providing a preliminary interpretation of their speciation process. Moreover, we suggest and outline 13 Angolan Main Biogeographic Units (AMBUs) in the area, defining a new biogeographic context for future works on Angolan herpetofauna. We now recognize eleven Hemidactylus species in Angola, and we provide here a new morphological key for Angolan Hemidactylus to help with identifications and species assignments in this group.
... The central and eastern localities of B. ombelanonga, as well as the latter from either B. adspersus or B. poweri, may be separated by drainage basins; however, with no contemporary sampling across regions spanning hundreds of kilometers, it is difficult to test these broad biogeographic hypotheses. Many recent initiatives have improved the current state of knowledge of Angola's herpetofauna, as well as to identify priority areas for future field survey work (Ceríaco et al. 2014(Ceríaco et al. , 2016Conradie et al. 2016;Heinicke et al. 2017;Marques et al. 2018;Baptista et al. 2019;Butler et al. 2019;Ernst et al. 2020), yet these efforts have still only scratched the surface. Additional, comprehensive field surveys, particularly those with focused/specialized efforts to record hard-to-find, seasonal, and/or fossorial taxa (e.g., by deploying pitfall traps, drift fence arrays, artificial refuges, etc., for an extended period of time or repeatedly throughout the year), should be priorities in the near future. ...
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Recent molecular phylogenetic work has found that Breviceps Merrem, 1820 comprises two major clades, one of which, the B. mossambicus group, is widely distributed across southern sub-Saharan Africa. This group is notable for harboring abundant cryptic diversity. Of the four most recently described Breviceps species, three are members of this group, and at least five additional lineages await formal description. Although Breviceps has long been known to occur in Angola, no contemporary material has been collected until recently. The three most widespread taxa, B. adspersus, B. mossambicus, and B. poweri, may all occur in Angola, but accurate species assignment remains challenging given the rampant morphological similarity between these taxa, and, until recently, the lack of genetic resources. Phylogenetic, morphological, and acoustic analyses of recently collected samples from disparate localities within Angola provide evidence for an undescribed species that is sister to B. poweri. The new species can be diagnosed from its sister taxon by lacking pale spots along the flanks, a pale patch above the vent, and a short, dark band below the nares (all present in B. poweri). Additionally, the male advertisement call differs from the three other Breviceps that might occur in Angola in having both a longer interval between consecutive calls and a higher average dominant frequency. We here describe this lineage as a distinct species, currently only known from Angola, and discuss the presence of other Breviceps taxa within Angola.
... Lygodactylus angolensis Bocage, 1896 (Tables 4; Fig. 1-11) Lygodactylus angolensis (Bocage 1896(Bocage : 110, 1897Monard 1937: 52;Loveridge 1947: 207;1957: 187;Hellmich 1957: 35;Pasteur 1965Pasteur [1964Branch 1998: 245;Broadley & Cotterill 2004: 41;Ceríaco et al. 2016aCeríaco et al. : 57, 2016bCeríaco et al. : 56, 57, 2018bMarques et al. 2018: 188,189;Branch et al. 2019: 315) Lygodactylus laurae (Schmidt 1933: 4;Mertens 1937: 6;Barbour & Loveridge 1946: 147;Marx 1959 Despite being a widespread species in Angola and surrounding countries, the species was only described at the end of the nineteenth century based on two specimens collected by the Portuguese explorer José d'Anchieta "on the walls of a corral" in Hanha, Benguela Province (Bocage 1896). Bocage (1896) noted that some specimens from Cahata and Galanga, both in Benguela Province, that he had previously (Bocage 1895) identified as L. capensis were in fact the newly described species as well. ...
Article
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At present the genus Lygodactylus is represented by three species in Angola confirmed by voucher specimens-L. angolensis, L. bradfieldi, and L. capensis-and two others believed to be present, but without specimens with precise localities, L. chobiensis and L. lawrencei. We present a detailed taxonomic revision of the group in Angola and describe three new species, Lygodactylus baptistai sp. nov. L. nyaneka sp. nov. and Lygodactylus tchokwe sp. nov. Phylogenetic analysis using the mitochondrial marker ND2, as well as morphological data support the recognition of the new species. In addition, data suggest that specimens historically assigned to L. capensis in Angola represent misidentifications of L. nyaneka sp. nov. and L. tchokwe sp. nov. We revisit the identity of Lygodactylus laurae, a junior synonym of L. angolensis. We also present the first confirmed record of L. lawrencei in the country, using both morphological and molecular data. The description of the new species and the revision of the taxonomic identity of the Angolan populations of the genus, raises the number of species occurring in the country to five. A key to the Angolan species is presented.
... Angola has always been a country of interest to biologists because of its rich, but poorly documented biodiversity, but was largely inaccessible to researchers, as a result of almost three decades of violent civil war. Since the end of the war in 2002, several field surveys have been conducted in the country, which have led to new species inventories of poorly explored regions, and has led to the discovery of taxa previously unrecorded from the country (Ernst et al. 2014(Ernst et al. , 2015(Ernst et al. , 2020Ceríaco et al. 2014aCeríaco et al. , 2016aCeríaco et al. , 2016bCeríaco et al. , 2018aBranch and Conradie 2015;Conradie et al. 2016;Baptista et al. 2019a;Butler et al. 2019), the description of new species (Conradie et al. 2012a(Conradie et al. , 2012b(Conradie et al. , 2013Stanley et al. 2016;Ceríaco et al. 2018bCeríaco et al. , 2020aCeríaco et al. , 2020bBranch et al. 2019a;Marques et al. 2019aMarques et al. , 2019b, and the study of rare and data-deficient species (e.g. Oliveira et al. 2016;Branch et al. 2017aBranch et al. , 2019bHeinicke et al. 2017;Agarwal et al. 2017;Gonçalves et al. 2019;Vaz Pinto et al. 2019). ...
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An integrative taxonomic review of the genus Boaedon in Angola is provided. A molecular phylogeny, based on 99 genetic samples for which the mitochondrial markers 16S rRNA have been sequenced, reveals 23 monophyletic species-level groups in Africa and indicates the presence of nine species in Angola. Based on both phylogenetic and morphological data, we revalidate and designate a neotype for B. angolensis, describe three new species for Angola (e.g. B. bocagei sp. nov., B. branchi sp. nov., and B. fradei sp. nov.), revalidate B. variegatus from its synonymy with B. lineatus and designate a lectotype for this taxon, and identify B. lineatus var. lineolatus as a junior synonym of B. variegatus. The taxonomic status of the recently described B. paralineatus from Central Africa is discussed with respect to the more inclusive B. lineatus group. Moreover, we report on a new country record for Angola, namely B. mentalis, which we elevate here to full species and discuss the taxonomic status of this species in southern Africa. Finally, we provide an identification key and updated distribution maps for all Boaedon species occurring in Angola, including the Cabinda enclave.
... The new studies on Angolan biodiversity have analyzed a wide diversity of taxonomic groups (Huntley et al. 2019) with herpetofauna being one of the groups that has attracted significant attention . These recent studies include the description of new species (Conradie et al. 2012a(Conradie et al. ,b, 2013Stanley et al. 2016;Ceríaco et al. 2018b;Marques et al. 2019a,b;Branch et al. 2019a,b), surveys of poorly explored areas (Ceríaco et al. 2014(Ceríaco et al. , 2016a(Ceríaco et al. ,b, 2018cBranch & Conradie 2015;Conradie et al. 2016;Baptista et al. 2018;Butler et al. 2019) and the rediscovery of rare and elusive species (Heinicke et al. 2017;Gonçalves et al. 2019). ...
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