Description of a new flat gecko (Squamata: Gekkonidae: Afroedura) from Mount Gorongosa, Mozambique

Abstract

A new species of flat gecko, Afroedura gorongosa sp. nov., is described from Gorongosa National Park, Sofala Province, central Mozambique. The new species is morphologically similar to A. transvaalica and A. loveridgei, from both of which it is genetically distinct (15–17% divergence; 400 bp of 16S rRNA). Morphologically it can be distinguished from both species by having fewer midbody scale rows (97–101) and a higher number of precloacal pores in males (8–13). The type series was collected on the western flanks of Mount Gorongosa (900 and 1100 m a.s.l.) in comparatively cool and moist microclimates, where it is threatened by illegal deforestation. Additional material was collected as low as 212 m a.s.l. on an inselberg near Mount Gorongosa. The new discovery adds to the growing number of endemic montane reptiles discovered in Mozambique in recent years, and highlights the need for a national conservation assessment of the country’s herpetofauna and continued protection of the Mount Gorongosa region.

Full text

142
Accepted by A. Bauer: 14 Aug. 2017; published: 26 Sept. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4324 (1): 142
–
160
http://www.mapress.com/j/zt/
Article
https://doi.org/10.11646/zootaxa.4324.1.8
http://zoobank.org/urn:lsid:zoobank.org:pub:B4FF9A5F-94A7-4E75-9EC8-B3C382A9128C
Description of a new flat gecko (Squamata: Gekkonidae: Afroedura)
from Mount Gorongosa, Mozambique
WILLIAM R. BRANCH
1,2,13
, JENNIFER A. GUYTON
3
, ANDREAS SCHMITZ
4
, MICHAEL F. BAREJ
5
,
PIOTR NASKRECKI
6,7
, HARITH FAROOQ
8,9,10,11
, LUKE VERBURGT
12
& MARK-OLIVER RÖDEL
5
1
Port Elizabeth Museum, P.O. Box 13147, Humewood 6013, South Africa
2
Research Associate, Department of Zoology, Nelson Mandela University, P.O. Box 77000, Port Elizabeth 6031, South Africa
3
Department of Ecology & Evolutionary Biology, Princeton University, Princeton, NJ 08544, U.S.A
4
Natural History Museum of Geneva, Department of Herpetology and Ichthyology, C.P. 6434, 1211 Geneva 6, Switzerland
5
Museum für Naturkunde Berlin, Leibniz Institute for Evolution and Biodiversity Science, Invalidenstrasse 43, 10115 Berlin, Germany
6
Edward O. Wilson Laboratory, Gorongosa National Park, Mozambique
7
Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A
8
Faculty of Natural Sciences, Lúrio University, Caixa Postal 958, Pemba, Mozambique
9
Department of Biology & CESAM, University of Aveiro, Aveiro, Portugal
10
Gothenburg Global Biodiversity Centre, Box 461, 405 30 Gothenburg, Sweden
11
Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, 405 30 Gothenburg, Sweden
12
Department of Zoology and Entomology, University of Pretoria, Pretoria, 0001, South Africa
13
Corresponding author: E-mail: williamroybranch@gmail.com
Abstract
A new species of flat gecko, Afroedura gorongosa sp. nov., is described from Gorongosa National Park, Sofala Province,
central Mozambique. The new species is morphologically similar to A. transvaalica and A. loveridgei, from both of which
it is genetically distinct (15–17% divergence; 400 bp of 16S rRNA). Morphologically it can be distinguished from both
species by having fewer midbody scale rows (97–101) and a higher number of precloacal pores in males (8–13). The type
series was collected on the western flanks of Mount Gorongosa (900 and 1100 m a.s.l.) in comparatively cool and moist
microclimates, where it is threatened by illegal deforestation. Additional material was collected as low as 212 m a.s.l. on
an inselberg near Mount Gorongosa. The new discovery adds to the growing number of endemic montane reptiles discov-
ered in Mozambique in recent years, and highlights the need for a national conservation assessment of the country’s her-
petofauna and continued protection of the Mount Gorongosa region.
Key words: Afroedura gorongosa sp. nov.; Afroedura loveridgei; Afroedura transvaalica; biodiversity; endemism; lizards
Resumo
Aqui descreve-se uma nova espécie de osga, Afroedura gorongosa sp. nov., do Parque Nacional da Gorongosa, na Provín-
cia de Sofala, centro de Moçambique. Do ponto de vista morfológico, a nova espécie assemelha-se às osgas A. transvaa-
lica e A. loveridgei, sendo geneticamente distinta de ambas (divergência de 15–17%; 400 bp de 16S rRNA). Distingue-se
morfologicamente de ambas as espécies por ter um número inferior de fileiras de escamas na secção mediana (97–101) e
um número superior de poros pré-cloacal nos machos (8–13). A série-tipo foi recolhida nos flancos ocidentais da Serra da
Gorongosa (900 e 1100 m a.s.l.), em microclimas relativamente mais frios e húmidos, onde se encontra ameaçada pela
desflorestação ilegal. Estas osgas foram também recolhidas à altitude de 212 m a.s.l., num inselberg e próximo da Serra
da Gorongosa. A nova descoberta junta-se ao crescente número de répteis endémicos de montanha descobertos em
Moçambique nos últimos anos, e realça a necessidade de uma avaliação da conservação da herpetofauna do país a nível
nacional, bem como a proteção da região do Monte Gorongosa.
Palavras-chave: Afroedura gorongosa sp. nov.; Afroedura loveridgei; Afroedura transvaalica; biodiversidade; endemis-
mo; lagartos

Zootaxa 4324 (1) © 2017 Magnolia Press
·
143
NEW AFROEDURA FROM MOZAMBIQUE
Introduction
Geckos have undergone an explosive radiation in southern Africa and are the dominant lizard family in the
subcontinent, with over 135 species currently known (Branch 2014 and updates). Despite their well-known digital
specializations they are otherwise often morphologically conservative, and the head and body are usually covered
with small granular scales, sometimes with scattered tubercles, that offer few opportunities for morphological
differentiation. Recent genetic studies, however, have uncovered unsuspected phylogenetic complexity (Bauer et
al. 1997; Bauer & Lamb 2005; Heinicke et al. 2014) and high levels of cryptic diversity within numerous
morphologically conservative gecko genera; e.g. Pachydactylus (Bauer & Branch 1995; Branch et al. 2011; Bauer
et al. 2006); Lygodactylus (Jacobsen 1992a, 1994a; Travers et al. 2014); or Goggia (Branch et al. 1995; Good et al.
1996; Bauer et al. 1996). This is particularly true of localized, allopatric populations of rupicolous species,
exemplified by crevice-dwelling flat geckos of the genus Afroedura Loveridge, 1944.
Loveridge (1944) erected Afroedura for a radiation of small to medium-sized rupicolous, leaf-toed geckos
restricted mainly to southern Africa. He separated them from Australian Oedura (Loveridge 1944), with which
they had originally been grouped, due to their lower scansor counts (1–3 in Afroedura, four or more in Oedura) and
in usually having a verticillate tail (unsegmented in Oedura), although some members of the Afroedura pondolia,
A. multiporis and A. langi-groups have been subsequently shown to possess unsegmented tails (Jacobsen 1992b).
In the last formal revision of African geckos, Loveridge (1947) recognized seven species and 12 taxa of Afroedura.
Onderstall (1984) named new taxa from South Africa, and noted further possible undescribed ones. Jacobsen
(1992b) presented a preliminary review of Transvaal populations, noting numerous new taxa but not formally
naming them. Aware of these reports, and of other problematic populations in the Cape region, the genus was
prioritized for further systematic study in the subcontinent (Branch 2006). Subsequently, known diversity within
the genus has greatly increased, with the description of nine new species, many with ultra-restricted (< 500 km
2
)
ranges (Jacobsen et al. 2014). Phylogenetic studies on the genus (Jacobsen et al. 2014; Makhubo et al. 2015)
placed the 27 currently recognized species in seven monophyletic clades, whilst further undescribed cryptic
diversity has also been signalled (Makhubo et al. 2015; Branch et al. 2017).
All Afroedura are considered endemic to southern Africa with the exception of A. loveridgei and A. bogerti,
both of the A. transvaalica-group (Jacobsen et al. 2014). Hewitt (1925) described Oedura transvaalica based on
material from “N’jelele River, Zoutpansberg District, Transvaal” (= Nzhelele River, a southern tributary of the
Limpopo River, that arises in the Soutpansberg east of Nzhelele (= Thohoyandou, 22°53'0.91"S, 30°29'2.79"E),
Limpopo Province, South Africa; Raper 1989). In the same paper Hewitt (1925) treated material from Umtali (=
Mutare) and Matopos, both north of the Limpopo River, as a northern race, O. t. platyceps, with Umtali as the type
locality. This race was distinguished (in part) by “…the form and size of the mental shield, which even at the base
is narrower than the first labial.” Loveridge (1947), after earlier transferring African Oedura to the new genus
Afroedura and describing A. bogerti from Angola (Loveridge 1944), felt that some features considered diagnostic
by Hewitt (1925) for A. t. platyceps, e.g. smaller size of mental, more depressed head, and less well-developed
distal scansors, were very variable. However, despite these observations, he still continued to recognize the race.
Subsequently, Broadley (1962) confirmed Loveridge’s observations and also considered an increase in precloacal
pore number in Zimbabwe populations of A. transvaalica to be clinal; e.g. 5–6 Beit Bridge (30 specimens), 6–8
Umatali (= Mutare, 48 specimens), 8–10 Mtoko (16 specimens). He, therefore, relegated Hewitt’s race H. t.
platyceps to the synonymy of A. transvaalica. Later, Bauer (2014) noted that a phylogenetic assessment of the
isolated populations of A. transvaalica was indicated, possibly based on a then unpublished molecular phylogeny
of the genus (Jacobsen et al. 2014). This paper revealed relatively deep sequence divergence among the widely
scattered A. transvaalica populations, indicating further potential cryptic diversity.
When describing A. transvaalica loveridgei from “five miles west of Tete, Mozambique”, Broadley (1963)
differentiated it from the typical race, which was widespread in Zimbabwe and entered South Africa along the
Limpopo River valley, by the rostral being excluded from the nostril and its smaller size. He noted that A. t.
loveridgei was “confined to the Zambezi valley below 1,500 feet” (500 m). Afroedura t. loveridgei was not the only
record of Afroedura from Mozambique, as Broadley (1966) recorded specimens of A. transvaalica from Magasso
and Mount Gorongosa, but noted that they occurred at higher altitudes than A. loveridgei. Branch (1998),
subsequently raised A. t. loveridgei to species status, and this was later supported by genetic analysis (Jacobsen et
al. 2014).

BRANCH ET AL.
144
·
Zootaxa 4324 (1) © 2017 Magnolia Press
Nocturnal and rupicolous, flat geckos usually shelter under exfoliating flakes on hard rock outcrops such as
granite, gneiss and some sandstones (Fig. 1). Their distributions are therefore often disjunct, with limited gene flow
between isolates. As a consequence the taxonomy of many populations has remained confused, and only recently
has the application of molecular assessments, coupled with increased surveys of previously inaccessible regions,
brought some resolution to species boundaries and phylogenetic relationships within the genus (Jacobsen et al.
2014; Makhubo et al. 2015). However, these taxonomic hypotheses require reassessment when new populations
are discovered or new genetic material becomes available. Broadley (1966) previously referred flat geckos from
Gorongosa National Park in Mozambique to A. transvaalica. Additional specimens collected in the park, including
genetic material, allowed us to reassess the taxonomic and phylogenetic relationships of this population.
FIGURE 1. Afroedura loveridgei (EI 0189) and its rocky habitat along the Zambezi River, just downstream of the Cahora
Bassa Dam, Tete Province, Mozambique (Photos: L. Verburgt).

Zootaxa 4324 (1) © 2017 Magnolia Press
·
145
NEW AFROEDURA FROM MOZAMBIQUE
Materials and methods
Species concept. We apply a general lineage-based species concept, treating as species those entities that represent
independent historical lineages (Frost & Hillis 1990; de Queiroz 1998). Our operational criteria to infer species
boundaries are both tree- and character-based.
Morphology. This study was based mainly on Afroedura material in the collections of the Port Elizabeth
Museum (PEM) and the Museum für Naturkunde Berlin (ZMB) (see Material Examined). Fresh material was
collected during two biodiversity surveys (April to December 2015) in Gorongosa National Park, Mozambique.
Specimens were euthanized with benzocaine. After death the geckos were fixed and stored in either 75% ethanol
(ZMB) or 55% isopropanol (PEM). Type material of Oedura transvaalica Hewitt, 1925 and Oedura transvaalica
platyceps Hewitt, 1925 was examined.
As with many other gecko genera Afroedura is morphologically conservative. Due to their granular scales
there are few discrete differences that are diagnostic. The number of paired and enlarged subdigital scansors has
been used to differentiate groups within the genus (FitzSimons 1943; Loveridge 1947; Onderstall 1984; Jacobsen
1997), but is invariant among the A. transvaalica-group, the only Afroedura occurring north on the Limpopo River
in south-eastern Africa. The following characters (detailed in Jacobsen 1992b) were assessed: 1: presence or
absence of internasal granules between the nasorostral scales; 2: number of postmental scales; 3: number of scales
between the eyes over crown of head; 4: number of scales between upper edge of earhole and rear margin of orbit;
5: number of scales between nostril and front edge of orbit, including postnasal; 6: number of supralabials; 7:
number of infralabials; 8: number of midbody scale rows; 9: number of scale rows on dorsal surface per tail whorl
(counted 3–6 verticils posterior to the cloaca); 10: number of scales rows on ventral surface per tail whorl; and 11:
number of precloacal pores in males.
Measurements have been taken by one person (WRB) and included: snout-vent length (SVL), tail length (TL);
head length (HL—tip of snout to back of jaw); head width (HW—widest point of head at level of eyes); snout
length (SL—tip of snout to front of orbit); ear-eye length (EE—top edge of earhole to back of orbit); and
internostril distance (IN—horizontal line between both nostrils).
Genetic analyses. Two recent, large-scale phylogenetic analyses of the genus Afroedura (Jacobsen et al. 2014;
Makhubo et al. 2015) have identified robust clades within the known species, and their known distributions are
well-documented. We therefore concentrated our genetic analyses on the species known to occur in Mozambique,
neighbouring Zimbabwe and adjacent South Africa, as well as the Angolan taxa referred to as A. bogerti. These all
form part of the previously defined tropical A. transvaalica-loveridgei-bogerti clade (Jacobsen et al. 2014), to
which the Gorongosa population is morphologically allied due to possessing two pairs of terminal scansors and a
verticillate tail. Our genetic assessment was therefore restricted to a species-level determination using the 16S
rRNA gene of the mitochondrial genome.
We rooted our phylogeny with Afroedura hawequensis Mouton & Mostert, 1985 from the southwestern Cape,
which is the sole member of the A. hawequensis-group sensu Jacobsen et al. (2014), and seemingly the most basal
member of the genus Afroedura (Jacobsen et al. 2014).
DNA was extracted from tissues using the E.Z.N.A. Tissue DNA Kit (VWR/Omega bio-tek) following the
manufacturers protocol. A portion of the mitochondrial genome (16S rRNA gene) was PCR amplified and
sequenced following the methods described in Schmitz et al. (2005). All sequences have been deposited in
GenBank (Table 1).
DNA sequences were aligned using the original chromatograph data in the program BioEdit (Hall 1999), using
ClustalX (Thompson et al. 1997) and the resulting alignment was manually edited. We managed to unambiguously
align 400 bp of the 16S rRNA gene including some fast-evolving, indel-rich loop regions. Bayesian Inference
(MrBayes, version 3.12; Huelsenbeck & Ronquist 2001) and Maximum Likelihood (RAxML version 7.0.4;
Stamatakis 2006) using the rapid hill climbing algorithm and the GTR+Γ model of nucleotide substitution
following Stamatakis et al. (2006) were applied to assess phylogenetic relationships. The best-fit model of
sequence evolution for the Bayesian analysis was selected using jModeltest 2.1.7 (Darriba et al. 2012), using the
Bayesian information criterion (BIC). Bootstrap analyses (BS) with 1000 pseudoreplicates in the ML analysis were
used to evaluate the relative branch support in the phylogenetic analysis. Clades with bootstrap values (ML) ≥ 70%
were considered strongly supported. Bayesian analyses were run for five million generations using four chains
sampling every 1000 generations, with a burn-in of 1000 trees. Clades with posterior probabilities (PP) ≥ 95% were
considered strongly supported.

BRANCH ET AL.
146
·
Zootaxa 4324 (1) © 2017 Magnolia Press
In total, sequences from 38 Afroedura specimens were included in the phylogenetic analyses (Table 1) to
corroborate our morphological analyses. The distribution of voucher and genetic material examined for species of
the A. transvaalica complex are shown in Fig. 2. Additional localities for A. transvaalica can be found in Broadley
(1966), Bauer (2014), and Jacobsen et al. (2014).
TABLE 1. Afroedura specimens (field and collection numbers), localities and GenBank accession numbers of vouchers
used in this study.
Species Locality Sample number GenBank
A. hawaquensis Limietberg, Western Cape Province, South Africa KTH10.09 LM993779
A. transvaalica Near border to Zimbabwe, Limpopo Province, South
Africa
MBUR 01714 LM993792
A. loveridgei Near Moatize, Tete Province, Mozambique EI 123 MF565446
A. loveridgei Near Moatize, Tete Province, Mozambique EI 189 MF565447
A. loveridgei Near Moatize, Tete Province, Mozambique EI 191/PEM R21992 MF565448
A. gorongosa sp. nov. (paratype) Gorongosa National Park, Sofala Province,
Mozambique
ZMB 83290 MF565449
A. gorongosa sp. nov. (holotype) Gorongosa National Park, Sofala Province,
Mozambique
ZMB 83293 MF565450
A. gorongosa sp. nov. (paratype) Gorongosa National Park, Sofala Province,
Mozambique
PEM R22220 MF565451
A. bogerti 1 Omauha Lodge, Iona National Park, Namibé
Province, Angola
KT 196 MF565452
A. bogerti 1 Omauha Lodge, Iona National Park, Namibé
Province, Angola
KT 197 MF565453
A. bogerti 1 Omauha Lodge, Iona National Park, Namibé
Province, Angola
KTH09-197 LM993777
A. bogerti 1 Omauha Lodge, Iona National Park, Namibé
Province, Angola
KTH09-196 LM993776
A. bogerti 2 50 km E Namibé on main tar road to Leba, Namibé
Province, SW Angola
ANG 289 MF565454
A. bogerti 2 Meva Beach, Benguella Province, Angola "30" MF565455
A. bogerti 2 20 km S Bentiaba, Namibe Province, Angola samp 62/E260.15 MF565456
A. bogerti 2 10.4 km S of Rio Makonga on tar road to Bentiaba,
Namibe Province, Angola
samp 58/E260.14 MF565457
A.bogerti 2 10.4 km S of Rio Makonga on tar road to Bentiaba,
Namibe Province,Angola
samp 57/E260.13 MF565458
A. bogerti 2 10.4 km S of Rio Makonga on tar road to Bentiaba,
Namibe Province,Angola
samp 39/E260.12 MF565459
A. bogerti 2 1 km E of Farm Mucongo, Namibe Province, SW
Angola
AG 1 37 MF565460
A. bogerti 2 52 km N on tar road on road to Lucira, Namibe
Province, Angola
ANG 311 MF565461
A. bogerti 2 1 km E of Farm Mucongo, Namibe Province, SW
Angola
AG 141 MF565462
A. bogerti 2 1 km E of Farm Mucongo, Namibe Province, SW
Angola
AG 1 38 MF565463
A. bogerti 3 400 m N of Mission de Namba grounds, Kwanza Sul
Province, Angola
samp 28/E260.5 MF565464
......continued on the next page

Zootaxa 4324 (1) © 2017 Magnolia Press
·
147
NEW AFROEDURA FROM MOZAMBIQUE
FIGURE 2. Distribution of the Afroedura transvaalica-complex in south-eastern Africa (excluding the Angolan A. bogerti-
complex); orange circles: A. loveridgei; green squares: A. transvaalica; red triangle: Mount Gorongosa, the type locality of A.
gorongosa sp. nov.; blue triangle: Bunga Inselberg, the location of additional A. gorongosa sp. nov. specimens. Localities
plotted only depict voucher and genetic material examined; additional localities for A. transvaalica can be found in: Broadley
(1966), Bauer (2014), and Jacobsen et al. (2014). The inset map shows in brown the African countries included in the
distribution map.
TABLE 1. (Continued)
Species Locality Sample number GenBank
A. bogerti 3 400 m N of Mission de Namba grounds, Kwanza Sul
Province, Angola
samp 27/E260.4 MF565465
A. bogerti 3 Farm Namba, Kwanza Sul Province, Angola samp 25/E260.3 MF565466
A. bogerti 3 Farm Namba, Kwanza Sul Province, Angola samp 23/E260.1 MF565467
A. bogerti 3 Farm Namba, Kwanza Sul Province, Angola samp 24/E260.2 MF565468
A. bogerti 4 Fm Victoria-Verdun, 2 km S of Mt Sandula, Kwanza
Sul Province, Angola
samp 34/E260.9 MF565469
A. bogerti 4 Fm Victoria-Verdun, 2 km S of Mt Sandula, Kwanza
Sul Province, Angola
samp 31/E260.6 MF565470
A. bogerti 4 Fm Victoria-Verdun, 2 km S of Mt Sandula, Kwanza
Sul Province, Angola
samp 33/E260.8 MF565471
A. bogerti 4 5 km SW Lepi, Huambo Province, Angola samp 37/E260.11 MF565472
A. bogerti 4 Candumbo Roacks Memorial, 20 km E Humana,
Huambo Province, Angola
samp 35/E260.10 MF565473
A. bogerti 4 Candumbo Rocks, Huambo Province, Angola WC 4037 MF565474
A. bogerti 4 Candumbo Rocks, Huambo Province, Angola WC 4038 MF565475
A. bogerti 4 Maka-Mombolo, NE of Balomba, Benguela Province,
Angola
samp 70/E260.16 MF565476
A. bogerti 4 5 km west of Maka-Mombolo, Benguela Province,
Angola
samp 71/E260.17 MF565477
A. bogerti 4 5 km west of Maka-Mombolo, Benguela Province,
Angola
samp 72/E260.18 MF565478
A. bogerti 4 5 km west of Maka-Mombolo, Benguela Province,
Angola
samp 73/E260.19 MF565479

BRANCH ET AL.
148
·
Zootaxa 4324 (1) © 2017 Magnolia Press
Results
Morphology. Results for the morphological analysis are summarised in Table 2 and discussed in the species
description below.
Molecular data. For the genetic dataset both the Bayesian and Maximum Likelihood analyses agreed fully in
the topology of the recovered phylogram (Fig. 3). The methods differed only in the degree of support recovered for
the individual nodes. These results fully corroborated those derived from the morphological analysis, and
confirmed that the Gorongosa population was not conspecific with any other members of the A. transvaalica-
group, including A. loveridgei and A. transvaalica (Table 3).
Our molecular assessment showed that the Gorongosa population forms part of the A. transvaalica-loveridgei
clade (sensu Jacobsen et al. 2014), which currently includes the species A. transvaalica (Hewitt 1925), A.
loveridgei Broadley, 1963, and A. bogerti Loveridge, 1944. Within this clade A. transvaalica is basal,
corresponding well to its proposed most basal phylogenetic position in previously published data on the genus
Afroedura (Makhubo et al. 2015). Afroedura bogerti is restricted to Angola, with a single Namibian record of
problematic status (Branch 1998; Griffin 2003). Although Angolan A. bogerti is now known to contain cryptic
diversity (Branch et al. 2017; this study), all known populations (1–4) form a subclade that is sister to A. loveridgei
and the Gorongosa population (Fig. 3).
All three Gorongosa Afroedura sequences were identical, and showed large uncorrected p-distances of ca.
15.7–17.7% to A. loveridgei, in which intraspecific variation was low at ca. 0.3–3.5% (Table 3). Similarly large
genetic distances of ca. 15% was revealed between the Gorongosa geckos and A. transvaalica, as well as between
A. transvaalica towards the different A. bogerti populations (13.1–17.3%; but see below) and A. loveridgei (16.9–
18.0%). The latter species was genetically even less similar to the different A. bogerti populations 1–4 (21.4%; see
below). These genetic distances correspond very well to the level of genetic differences found between other well
established species within the genus Afroedura for the same gene (Makhubo et al. 2015; this study, Table 3).
FIGURE 3. Phylogenetic tree topology for the 16S gene (identical for Bayes and ML) using Afroedura hawequensis as
outgroup. Support values for posterior probabilities and bootstraps are indicated by circles (PP: >0.95 / ML: >70%) : full black
circles indicated support by both methods and full open circles indicate no strong support by either method.
Similar to previous findings of clear and well supported phylogenetic substructure in A. transvaalica (Jacobsen
et al. 2014), our analyses revealed that Angolan Afroedura bogerti populations comprise four species-level
subclades, indicating the existence of three undescribed Angolan Afroedura species (Fig. 3). While the genetic
distances found between those taxa are lower than between the other analysed Afroedura species in this study, all

Zootaxa 4324 (1) © 2017 Magnolia Press
·
149
NEW AFROEDURA FROM MOZAMBIQUE
are mostly well supported. They form fully allopatric sub-clades which show consistently high inter- and lower
intrapopulational genetic distances, which roughly correspond to the distances within the A. transvaalica sub-
clades (see Jacobsen et al. 2014; Table 3), indicating prolonged reproductive isolation. A more detailed analysis of
the A. bogerti candidate species, including identification of the nominate population, will be presented elsewhere.
TABLE 2. Summary of morphological comparison of Afroedura transvaalica (Eastern Zimbabwe populations), A. loveridgei
and A. gorongosa sp. nov.; given is sample size (number), morphological characters (compare material & methods); values
are provided as range, mean () ± standard deviation (in parenthesis); M= male, F= female, N= sample size.
TABLE 3 . Uncorrected pairwise genetic distances of Afroedura gorongosa sp. nov. with other members of the
Afroedura transvaalica-loveridgei-bogerti clade and A. hawaquensis for 16S rRNA (compare text, Table 1 and Fig. 3).
Given are minimum (MIN), maximum (MAX) and values of uncorrected pairwise sequence divergence, presented in
% of 400 bp of the 16S rRNA gene; SD= standard deviation; N= number of vouchers per clade; Nc= number of direct
sequence comparisons.
In summary, both genetics and morphology (particularly diagnostic features such as the rostral and nasal
condition, and the number of midbody scale rows and precloacal pores) confirmed that the Gorongosa population
should be considered as a new, undescribed species within the genus Afroedura which is described below.
Character A. transvaalica (N= 22) A. loveridgei (N= 12) A. gorongosa sp. nov. (N= 6)
Snout vent (max) M 71 mm, F 68 mm M 59 mm, F 57 mm M 57 mm, F 61 mm
HL / HW (1.1 ± 0.04) (1.2 ± 0.06) (1.32± 0.05)
Snout / Eye (1.8 ± 0.15) (1.5 ± 0.22) (1.48 ± 0.10)
Snout / Ear-eye (1.1 ± 0.14) (1.1 ± 0.12) (1.34 ± 0.05)
Precloacal pores (males) 6–10 (7.9) 8–10 (8.8) 8–13 (10)
Dorsal rows tail whorl 6–8 (7.2 ± 0.34) 7–8 (7.8 ± 0.32) 7–8 (7.2 ± 0.45)
Ventral rows tail whorl 5–5 (5.15 ± 0.34) 5–6 (5.8 ± 0.41) 5 (5)
Scales below 4
th
toe 8–13 (9.86 ±1.17) 8–10 (9 ± 0.78) 10–12 (11 ± 0.84)
Midbody scale rows 108–119 (114.3 ± 3.5) 113–120 (116.6 ± 2.07) 97–101 (99.6 ± 1.67)
Scales between eyes 15–19 (17.9 ± 1.27) 15–19 (17.2 ± 1.12) 19-21 (20.2 ± 0.75)
Scales—nostril to eye 10-13 (11.45 ± 0.74) 10-12 (11.1 ± 0.67) 12-13 (12.5 ± 0.55)
Scales—ear to ey e 16–21 (19.2 ± 1.27) 17–21 (18.9 ± 1.38) 21–24 (22.7 ± 1.03)
Scales around nostril 5 (5) 3–5 (4.2 ± 0.58) 5 (5)
Contact anterior nasals 82% (4 with single granule) 0% 100% (1+2 to 2+3)
Upper labials 10–13 (11.7 ± 0.73) 9–10 (9.6 ± 0.52) 9–11 (10.0 ± 0.63)
Lower labials 9–12 (10.5 ± 0.89) 9–11 (9.4 ± 0.67) 9–10 (9.5 ± 0.55)
Species MIN (with
bogerti s.l. intra-
clade distances)
MIN (without
bogerti s.l. intra-
clade distances)
max  (with/without
bogerti s.l. intra-
clade distances)
SD (with/without
bogerti s.l. intra-
clade distances)
intra-clade
distances
NNc
Afroedura
hawaquensis
18.70 18.70 22.00 20.41 0.95 - 1 37
A. transvaalica 13.12 13.20 19.37 15.75 1.56 - 1 37
A. loveridgei 15.59 15.59 22.02 17.97 1.50 0.27–3.49 3 105
A. gorongosa
sp. nov.
14.47 14.47 19.53 16.72 1.33 0 3 105
A. bogerti 1 7.68 17.16 21.38 12.31/19.03 3.88/1.51 0 4 136
A. bogerti 2 7.68 16.44 20.96 13.56/18.26 3.15/1.19 0–4.60 10 296
A. bogerti 3 6.64 16.10 21.93 12.07/17.91 4.11/1.67 0 5 165
A. bogerti 4 6.64 13.12 20.63 12.25/16.10 3.24/1.77 0–4.61 11 297

BRANCH ET AL.
150
·
Zootaxa 4324 (1) © 2017 Magnolia Press
Afroedura gorongosa sp. nov.
(Figs. 4–5)
Afroedura transvaalica (part) Broadley 1966, p. 111.
Holotype. ZMB 83293 (GNP 484), adult male, western flank of Mount Gorongosa, Gorongosa National Park,
Sofala Province, Mozambique (1038 m a.s.l., 18˚28’04.3”S, 34˚02’53.2”E), collected by M.-O. Rödel and M.F.
Barej, 11 December 2015.
Paratypes (three specimens). ZMB 83290 (GNP 438), adult female, with incision in left thigh; PEM R22220
(GNP 439, previously ZMB 83291), adult female, with incision in left thigh; ZMB 83292 (GNP 440; will later be
deposited in the zoological collection of the E.O. Wilson Laboratory in Gorongosa National Park), adult male, with
everted hemipenes and incision in left thigh; collected on 22 May and 21 July 2015 by J.A. Guyton and P.
Naskrecki from gallery forest near Murombodzi Waterfall, Mount Gorongosa, Gorongosa National Park, Sofala
Province, Mozambique (842 m a.s.l; 18°29'0.1''S, 34°2'34.6''E).
Additional material (two specimens). ZMB 83288 (GNP 433), adult male, ZMB 83289 (GNP 434), adult
female; both collected in April 2015 by H. Farooq, P. Naskrecki and J.A. Guyton from the top of Bunga Inselberg,
Gorongosa National Park, Sofala Province, Mozambique (approx. 200 m a.s.l., 18˚35’58.1”S, 34˚20’34.8”E).
Etymology. The specific name refers to Mount Gorongosa and Gorongosa National Park in Central
Mozambique, to which the species is endemic. We suggest Gorongosa Flat Gecko is a suitable common name.
Afroedura, based on Oedura, is feminine and the specific epithet is treated as a noun in apposition.
Diagnosis. A member of the A. transvaalica-complex in possessing two pairs of enlarged scansors per digit
and a strongly verticillate and flattened tail (Jacobsen et al. 2014). Differs from other members of the complex by
having 97–102 midbody scale rows (less than 95 in A. bogerti-complex, 113–120 (= 117) in A. loveridgei and
102–119 in A. transvaalica (102–118, = 109 in South Africa—Jacobsen et al. 2014; 108–119, = 114 in
Zimbabwe adjacent to Mount Gorongosa); by the rostral bordering the nostril (nostril excluded from rostral in A.
loveridgei); by the anterior nasals being separated by 1–3 granules (always in broad contact in A. loveridgei;
usually in broad contact in A. transvaalica - <3% in South Africa populations, 18% separated by a single granule in
Zimbabwe adjacent to Mount Gorongosa); in having 19–22 scales between the anterior borders of the eyes (15–19
in A. loveridgei; 17–20 in South African A. transvaalica, and 15–19 in Zimbabwe populations adjacent to
Mozambique); and in having a higher average number of precloacal pores in males (8–13, = 10; 6–10, = 8 in A.
transvaalica; 8–10, = 9 in A. loveridgei).
Holotype description.
Morphology. Adult male; SVL 50.8 mm; tail 52.2 mm (partly regenerated), with a small incision in the left
thigh for the removal of muscle tissue. Head and body dorsoventrally depressed; HL 12.3 mm, HW 9.0 mm,
broadest at mid-eye and 1.37x longer than wide. Eye large (3.7 mm wide), pupil vertical with indented margins;
circumorbital scales small and granular, becoming elongate, almost ciliate, at upper posterior margin. Snout
rounded, 4.9 mm long, longer than distance between eye and ear openings (3.6 mm). Scales on top of snout
granular, rounded and convex, the largest being 2/3rds the width of the scales on the back, which are granular and
juxtaposed, with no intervening minute granules. Scales on snout almost twice as large as those on crown and
throat. Scales on belly flattened, imbricate, more-or-less ovate at mid-ventrum, and twice the size of lateral
granules and 1.5x those along backbone. Ear opening deep, oblique and roughly oval, only half as high as wide.
Limbs well developed, hindlimbs slightly longer than forelimbs, both without obvious mite pockets at
posterior margin of limb insertions. All digits with a large pair of distal scansors, separated by a large, curved claw,
and followed after a large gap (twice length of terminal scansor) by a smaller pair of scansors; infero-median row
of digital scales enlarged transversely, particularly towards the scansors, where the terminal scale adjoining the first
pair of scansors may be medially constricted, swollen and scansor-like; 12 scale rows under 4
th
toe.
Nostril pierced between rostral, 1
st
supralabial and three nasal scales; the supranasal being much larger than the
subequal postnasals and separated by a granule bordering the rostral, followed behind by two smaller granules.
Rostral roughly rectangular but with its upper edges elongated due to extensions into the nostril. Nine supralabials
on each side, the labial margin flexing dorsad at the rictus, with 1–2 minute scales proximal to the flexure. Nine
infralabials on either side, with a small scale proximad to the flexure. At the lip, mental slightly narrower than
adjacent infralabial, only 2/3rds the width of rostral, and in contact with two distinctly elongate postmental scales.

Zootaxa 4324 (1) © 2017 Magnolia Press
·
151
NEW AFROEDURA FROM MOZAMBIQUE
Approximately 20 scales across crown at level of front of eye; 13 scales between nostril and front of eye; 23 scales
from front of ear to back of eye; 99 scales around midbody.
FIGURE 4. Afroedura gorongosa sp. nov. a) male holotype (ZMB 83293, Mount Gorongosa, Mozambique; photo: M.-O.
Rödel); b) female paratype (ZMB 83292, Mount Gorongosa, Mozambique; photo: P. Naskrecki) showing dorsal pattern and
regenerated tail; c) Afroedura gorongosa (ZMB 83289, Bunga Inselberg, Gorongosa National Park, Mozambique; photo: C.
Dorse).

BRANCH ET AL.
152
·
Zootaxa 4324 (1) © 2017 Magnolia Press
FIGURE 5. Afroedura gorongosa sp. nov. female paratype (PEM R22229, previously ZMB 83291, Mount Gorongosa,
Mozambique; photos: P. Naskrecki); a) general habitus and dorsal coloration; b) close up of nasal region showing diagnostic
features of: presence of internasal granules, and posterior projection of rostral to border the nostril; c) close up of lower side of
left forelimb; and d) close up of right hind foot showing two paired scansors.
A roughly V-shaped row of 13 precloacal pores occurs eight scale rows anterior to the cloacal lip, with these
scales more imbricate and triangular adjacent to the precloacal pores. Original portion of tail slightly dorsoventrally
flattened and indistinctly verticillate, without obvious lateral constrictions; each verticil comprising 6–7 imbricate
rows of scales dorsally and 5 imbricate scale rows ventrally, and with ventral scales approximately twice the size of
those on dorsal surface. Hemipenal pouches well-developed, with an angular, ventro-lateral series of 3 enlarged
tubercular scales on each side (largest nearest cloaca), and with two openings of the cloacal sacs on the posterior lip
of the cloaca.
Colour in life (from photographs, Fig. 4a and unpublished). Dark black-brown above with a series of five
white dorsal spots (4–5 scales across) between the fore- and hind-limb insertions, that are surrounded by a
chocolate brown zone that encompasses a series of 2–4 laterally arranged smaller pale spots (2–3 scales wide), and
posteriorly by a diffuse pale region that forms an irregular band across the mid dorsum. Top of head, flanks and
upper surfaces of limbs with small, scattered pale spots. Head with a dark brown band across the posterior edge of
crown encompassing a small central pale spot; a vague, thin pale canthal stripe, extends on both sides from the
nasal region to front of eye; upper and lower labials grey-brown with diffuse yellowish bars; iris reddish brown
with a heavy dark brown reticulation, and pupil with crenulated edge. Original segment of tail mottled in dark
brown and pale blotches; regenerated tail grey with vague darker blotches. Ventrum mottled grey-cream.
Colour in preservative. Dark black-brown above with irregular markings on back, comprising four equally
spaced dark bars, each with a central white spot 4–5 scales wide, and an irregular, paler brown bar behind; top and
sides of head, flanks and upper surfaces of limbs with small, irregular pale spots; ventral surface of throat, neck,
belly, limbs and tail uniform dark grey. Side of head and labials dark brown, with a few scattered pale spots on
upper labials, and vague pale edges to lower labials.
Vari a t i o n in paratypes and additional material (Figs. 4b, 4c, 5; Tables 3–4).
Morphology. SVL from 54.6 mm (ZMB 83289) to 61.2 mm (PEM 22220); head length 1.24–1.38 times head
width (max. in paratypes 1.38); snout only 1.5 times diameter of eye in one specimen. The supranasals always
separated by granules, usually with a single large granule in contact with the rostral between the supranasals,
followed by 2–3 smaller granules in lateral contact; the first upper labial and rostral always enters the nostril, and
the width of the rostral at the lip margin is always wider than that of the mental (except in ZMB 83290 were they
are subequal); always two postmental scales; supralabials more than in holotype (10–11), infralabials (9–10);
scales between anterior edge of eye 19–21; scales between nostril and anterior edge of orbit 12–13; scales between

Zootaxa 4324 (1) © 2017 Magnolia Press
·
153
NEW AFROEDURA FROM MOZAMBIQUE
anterior edge of ear and rear margin of orbit 21–24; scales around mid-body 97–101; subdigital lamella beneath 4
th
toe 10–11; dorsal scales per tail verticel 7–8; ventral scales per tail verticel 5.
In one female (ZMB 83290) the terminal, medially-constricted subdigital lamella of the 3
rd
toe of left foot is
swollen and scansor-like, bearing small setae. Precloacal pores 8–13 in males, females without dimples in
precloacal scales (as in A. transvaalica, but unlike A. loveridgei where five females had 5–9 dimpled precloacal
scales, that lacked secretory activity, and only one had none).
Colour in life (based on female paratype PEM R22229). Boldly patterned, with six dorsal dark transverse
bands, posteriorly edged with cream, the first on the neck, the second and sixth at the fore- and hindlimb insertions,
respectively, and with three thin bands equally spaced between them on the body; background colour of body, head
and upper surfaces of limbs flecked on light brown and cream with occasional pale spots 2–4 scales wide; a dark
canthal stripe runs through the rostral to the front of the eye, with a paler zone above; the scales of the anterior-
dorsal region of the orbital rim are bright yellow, with a faint yellowish infusion of the crown scales between the
eyes; the original tail is boldly barred dorsally with seven, subequally-spaced, dark brown-black bars with a yellow
posterior edge; belly uniform cream with scattered yellowish scales, greyer on ventral surfaces of limbs and tail.
Colour in preservative. Similar to holotype, but dorsum pale brown to grey; larger dorsal markings usually
reticulated, occasionally forming irregular transverse bands (6–7 from occiput to between hindlimbs); smaller
dorsal markings vary from vague to distinct. A few specimens have irregular pale blotches along the posterior
edges of the darker dorsal patches/bands, and also scattered on the dorsal surfaces of the limbs and back. Dorsal
surface of regenerated tails usually with irregular pale and dark blotches that may form wavy bands, and which
become flecks or grey blotches below.
Size. Largest male (ZMB 83288)—SVL 56.5 mm, tail (partially regenerated) 55.7 mm; largest female (PEM
R22220)—SVL 61.2 mm, tail (original) 64.0 mm. It is thus intermediate in size between A. transvaalica (male—
max. SVL 72 mm, female—max. SVL 64 mm) and A. loveridgei (male—max. SVL 59 mm, female—max. SVL 58
mm). Only one of six specimens had an original tail (105% SVL).
Hemipenis (based on ZMB 83292). Approximately 5 mm long, with a simple, smooth, possibly longitudinally
flounced pedicel; capped with two large cups covered with fine calyculate ornamentation; sulcus simple and
draining into the conjoined base of the distal cups.
Distribution. Known from only three localities in the Mount Gorongosa region (Fig. 2). The holotype and
paratypes, plus various non-collected individuals, have been found on the western flanks of Mount Gorongosa
between about 900 and 1100 m a.s.l. (Figs. 6a–c). Access to further potential areas on the mountain, i.e. rock faces
at the eastern slopes, was not possible due to security concerns. The additional material and various non-collected
specimens have been observed at a lower altitude on the Bunga Inselberg (about 212 m a.s.l.) in the north-western
corner of the National Park, approximately 40 km from the type locality (Fig. 6d). No tissue samples from the
Bunga individuals were available, and the assignment of these individuals to the new species was thus based on
morphological similarity.
Natural history. The Gorongosa National Park is situated in Central Mozambique approx. 100 km south-west
of the Zambezi River, and is in the southern part of the great African Rift Valley. Mount Gorongosa is an isolated
granite mountain massif rising to 1863 m a.s.l., north-west of the National Park’s plains. At its highest elevations
the massif’s plateau is covered with montane grasslands and forest patches dominated by cypresses (Widdringtonia
nodiflora); the upper slopes of the mountain (900–1600 m) support a belt of moist evergreen forest. The type series
was collected in an area between 900 and 1100 m on the western flanks of the mountain (Figs. 6a–c, 7). The
paratypes were collected in a small alcove beneath a boulder pile in the closed-canopy riverine forest beside the
Murombodzi Waterfall, and thus in comparatively cool and moist microclimates (Figs. 6c, 7). Some rocks with
lizards present were located within the splash zone of the waterfall. Other specimens were observed but not
collected at this site. The holotype was collected from a deep narrow crack in a large granite rock (i.e. 8 m³) in
highly degraded grassland below the edge of the rainforest and almost fully exposed to the sun (Fig. 6b). It was the
only specimen seen at that site. At both sites the new species lived in syntopy with the cordylid Smaug
mossambicus (FitzSimons, 1958).
The additional non-type material comes from the Bunga Inselberg (Fig. 6d), a partly eroded inselberg
composed mostly of trachyte in the north-western corner of the national park, covered and surrounded by dense
Miombo woodland at a relatively low altitude (approx. 212 m a.s.l.). Here the geckos were observed during the day
and at night in several places in deep cracks between very large boulders. No geckos were observed outside of

BRANCH ET AL.
154
·
Zootaxa 4324 (1) © 2017 Magnolia Press
these cracks. Syntopic lizards were Smaug mossambicus, the skink Trachylepis margaritifera (Peters, 1854) and
the gecko Hemidactylus platycephalus Peters, 1854. Further sympatric, but not syntopic geckos were:
Lygodactylus cf. capensis (Smith, 1849), Pachydactylus punctatus Peters, 1854 and Chondrodactylus turneri
(Gray, 1864).
FIGURE 6. Habitats of Afroedura gorongosa sp. nov. on Mount Gorongosa (a–c); a) edge of rainforest on Mount Gorongosa,
the holotype was collected in a rock-crack of an isolated boulder in an open area close to the rainforest at 1038 m a.s.l. (b) and
in rocky areas of the Murombodzi river (c; compare Fig. 7); d) rocky habitat on the top of Bunga Inselberg, Gorongosa National
Park, Mozambique (photos: M.-O. Rödel).

Zootaxa 4324 (1) © 2017 Magnolia Press
·
155
NEW AFROEDURA FROM MOZAMBIQUE
FIGURE 7. Habitat of Afroedura gorongosa sp. nov. paratypes around Murombodzi waterfall, Mount Gorongosa; inset figure:
J.A. Guyton collecting one of the paratypes (photos: J. Poole).
TABLE 4. Museum accession numbers and measurements (mm) for the type series and additional material (AM) of
Afroedura gorongosa sp. nov.
The new species was observed regularly between April and December 2015. Like A. transvaalica, it is tolerant
of conspecifics and frequently several specimens, and sometimes as many as over 20, were observed sheltering in
deep rock crevices. One adult female (ZMB 83290) contained two eggs of approximately 11.5 x 7.5 mm.
Conservation Status. The population of the new gecko present on the lower slopes of Mount Gorongosa is
threatened by illegal deforestation taking place within the riverine forest adjacent to Murombodzi Waterfall. Slash-
and-burn removal of tall, old growth trees around the rocks may cause changes to the humidity and water
availability in these currently very moist, shaded microhabitats that seem to be preferred by the lizards.
Reforestation efforts led by Gorongosa National Park and introduction of shade-grown coffee as an alternative to
slash-and-burn agriculture still practiced around the mountain will hopefully slow or halt the loss of the native
riverine and evergreen vegetation in that area. On Bunga Inselberg the population of A. gorongosa is currently
Mus. No Status Sex SVL Tail Head
length
Head
width
Snout Eye Ear-eye Internostril
distance
ZMB 83293 Holotype M 50.8 52.2 12.3 9.0 4.9 3.7 3.6
PEM R22220
(ZMB 83291)
Paratype F 61.2 64.0 14.8 11.2 6.1 3.8 4.4 2.2
ZMB 83290 Paratype F 61.1 49.3 15.0 10.9 6.1 3.9 4.7 2.1
ZMB 83292 Paratype M 56.4 45.6 13.0 10.3 5.2 3.5 4.1 1.9
ZMB 83288 AM M 56.5 55.7 14.2 11.1 6.0 4.2 4.3 1.6
ZMB 83289 AM F 54.6 na 13.8 10.7 5.5 3.8 4.4 1.8

BRANCH ET AL.
156
·
Zootaxa 4324 (1) © 2017 Magnolia Press
protected within the core area of the National Park and no immediate threat to its survival exists, though
deforestation for agriculture is slowly encroaching on the area. However, based on the existence of only a few
known sites and small area of occupancy, i.e. the presumed restriction to a fragmented range on Mount Gorongosa
and potentially some rocky areas surrounding the mountain, as well as the ongoing high rate of habitat degradation
and conversion on the mountain, the species may be considered highly threatened, potentially justifying an
assessment as Endangered or even Critically Endangered. Further surveys and formal IUCN Red List assessment
are required to confirm this.
Key to the Afroedura transvaalica-group
1. Midbody scale rows more than 95 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Midbody scale rows less than 95; Angola. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bogerti (sensu lato)
2. Rostral bordering nostril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Rostral excluded from nostril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. loveridgei
3. Anterior nasals in contact (very rarely separated); scales around midbody-South Africa 102-118 (= 109), northern Zimbabwe
108-119 (= 114). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. transvaalica
- Anterior nasals separated by 1-3 granules; scales around midbody 99-101 (= 100) . . . . . . . . . . . . . . . . A. gorongosa sp. nov.
Discussion
The molecular phylogeny of Afroedura (Jacobsen et al. 2014) revealed cryptic diversity in the diverse populations
of A. transvaalica studied. They noted “There is clear and well supported genetic substructure within … A.
transvaalica”, and this presumably led Bauer (2014) to note that cryptic diversity in A. transvaalica deserved
further study. Broadley (1962) noted a clinal increase in precloacal pore number within populations of A.
transvaalica, with the lowest counts occurring on both sides of the Limpopo River. This was confirmed by
Jacobsen et al. (2014), and is also reflected in midbody scale row counts, which are also higher in northern
Zimbabwe (see Table 3) compared with South Africa (Jacobsen et al. 2014). This morphological variation
correlates with genetic substructure and supports Bauer’s (2014) call for further study on the species. We do not,
however, consider Hewitt’s name Oedura transvaalica platyceps, based on material from Umtali (= Mutare),
eastern Zimbabwe, to be available for the Gorongosa population due to their high midbody scale counts. It may,
however, be available for northern populations of A. transvaalica, should future studies on the species confirm
species-level divergence.
Mount Gorongosa and its surroundings exhibit high levels of biodiversity and represent an important centre of
regional endemism, of which A. gorongosa sp. nov. is the most recent addition. As noted by Jacobsen et al. (2014),
the major clades of Afroedura are generally restricted geographically, with the largest disjunction occurring within
the A. transvaalica clade, where A. bogerti occurs in western Angola. The inclusion of the latter within the A.
transvaalica clade is well-supported morphologically, as well as genetically, in both this study and that of Jacobsen
et al. (2014). Similar east-west disjunctions are known in other rupicolous groups, including the cordylid genera
Cordylus (Stanley et al. 2011, 2016) and Platysaurus (Branch & Whiting 1997; Whiting et al. 2015), and where
previously associated with movement of Aeolian Kalahari sands in the Plio-Pleistocene (Broadley 1978; Jacobsen
1994b). However, recent phylogenetic studies of these and related genera (Bauer & Lamb 2002; Broadley et al.
2014) indicate that initial ancestral isolation was associated with earlier cladogenic events (Scott et al. 2004),
although later climatic oscillations may have driven subsequent speciation in these genera (Whiting et al. 2015;
Stanley et al. 2016; Branch et al. 2017). Moore et al. (2017) discuss the effect on plant diversity and endemism of
widespread erosion along the eastern margin of southern and eastern Africa from the Late Palaeogene to the
present. This accentuated the relief of the eastern escarpment bordering the Manica Highlands, Zimbabwe, and
gave rise to the characteristic regional bornhardt or inselberg topography, of which Mount Gorongosa is the most
prominent survivor.
The close relationship of A. gorongosa sp. nov. to adjacent congeners in the eastern highlands of Zimbabwe is
reflected in other endemic taxa from Mount Gorongosa, e.g. the freshwater crab Potomonautes gorongosa
(Cumberlidge et al. 2016), the forest shrew Myosorex meesteri (Taylor et al. 2013), and the dwarf leaf chameleon
Rhampholeon gorongosae (Broadley 1971). All of these examples involve genera with species diversity associated

Zootaxa 4324 (1) © 2017 Magnolia Press
·
157
NEW AFROEDURA FROM MOZAMBIQUE
with the archipelago of montane isolates in adjacent Zimbabwe, Malawi and central Mozambique (freshwater
crabs: Daniels & Bayliss 2012; Phiri & Daniels 2012; shrews: Taylor et al. 2013; chameleons: Branch & Tolley
2010; Branch et al. 2014).
Acknowledgements
We thank the following curators/collections managers for support and/or access to material in their care: Werner
Conradie and Gill Watson, Port Elizabeth Museum; Frank Tillack, Museum für Naturkunde Berlin; Denise
Hamerton, IZIKO Museum (= South African Museum) for forwarding images of the type of A. t. platyceps; Chris
Dorse for images of the new species used in Figs. 4c and 5c, Joyce Poole for inset picture in Fig. 7 and Ninda
Baptista for the Portuguese Resumé. Travel costs to Berlin by WRB were supported by Incentive funding from the
National Research Foundation, South Africa. Travel and maintenance costs for JAG were supported by a U.S.
National Science Foundation Graduate Research Fellowship, a National Geographic Young Explorers Grant, and
Princeton University. Support for biodiversity surveys that resulted in the discovery of the new species was
provided by Greg Carr, Gorongosa Restoration Project, and the Prager Family. We also thank Mateus Mutemba,
Marc Stalmans, and Pedro Muagura for research and collecting permits, and Lucilia Chuquela from the Natural
History Museum, Maputo (MHN) for specimen export permits.
References
Bauer, A.M. (2014) Afroedura transvaalica: species account. In: Bates, M.F., Branch, W.R., Bauer, A.M., Burger, M., Marais,
J., Alexander, G.J. & de Villiers, M.S. (Eds.), Atlas and Red Data Book of the Reptiles of South Africa, Lesotho and
Swaziland. Suricata 1. South African National Biodiversity Institute, Pretoria, pp. 99.
Bauer, A.M. & Branch, W.R. (1995) Geographic variation in western populations of the Pachydactylus punctatus complex
(Reptilia Gekkonidae). Tropical Zoology, 8, 69–84.
https://doi.org/10.1080/03946975.1995.10539273
Bauer, A.M., Branch, W.R. & Good, D.A. (1996). A new species of rock-dwelling Phyllodactylus (Squamata: Gekkonidae)
from the Richtersveld, South Africa. Occasional Papers of the Museum of Natural Sciences, Louisiana State University,
71, 1–13.
Bauer, A.M., Good, D.A. & Branch, W.R. (1997) The taxonomy of the southern African leaf-toed geckos, with a review of Old
World “Phyllodactylus” (Squamata: Gekkonidae) and the description of five new genera. Proceedings of the California
Academy of Sciences, 49, 447–497.
Bauer, A.M. & Lamb, T. (2002) Phylogenetic relationships among members of the Pachydactylus capensis group of southern
African geckos. African Zoology, 37, 209–220.
https://doi.org/10.1080/15627020.2002.11657176
Bauer, A.M. & Lamb, T. (2005) Phylogenetic relationships of southern African geckos in the Pachydactylus group (Squamata:
Gekkonidae). African Journal of Herpetology, 54, 105–129.
Bauer, A.M., Lamb, T. & Branch, W.R. (2006) A revision of the Pachydactylus serval and P. weberi groups (Reptilia: Gekkota:
Gekkonidae) of Southern Africa, and with the description of eight new species. Proceedings of the California Academy of
Sciences, 57, 595–709.
Branch, W.R. (1998) Field Guide to Snakes and other Reptiles of Southern Africa. Struik, Pretoria, 236 pp.
Branch, W.R. (2006) Priorities for systematic studies on southern African reptiles. In: Branch. W.R., Tolley, K.A.,
Cunningham, M., Bauer, A.M., Alexander, G., Harrison, J.A., Turner, A.A. & Bates, M.F. (Eds.), A Plan for Phylogenetic
Studies of Southern African Reptiles: Proceedings of a Workshop Held at Kirstenbosch, February 2006. Biodiversity Series
No. 5. South African National Biodiversity Institute, Pretoria, pp. 2–20.
Branch, W.R. (2014) Reptiles of South Africa, Lesotho and Swaziland: Conservation status, diversity, endemism, hotspots and
threats. In: Bates, M.F., Branch, W.R., Bauer, A.M., Burger, M., Marais, J., Alexander, G.J. & de Villiers, M.S. (Eds.),
Atlas and Red Data Book of the Reptiles of South Africa, Lesotho and Swaziland. Suricata 1. South African National
Biodiversity Institute, Pretoria, pp. 22–50.
Branch, W.R., Bauer, A.M., Jackman, T.R. & Heinicke, M. (2011) A new species of the Pachydactylus weberi complex
(Reptilia: Squamata: Gekkonidae) from the Namib-Rand Reserve, southern Namibia. Brevoria, 524, 1–15.
https://doi.org/10.3099/0006-9698-524.1.1
Branch, W.R., Bayliss, J. & Tolley, K.A. (2014) Pygmy chameleons of the Rhampholeon platyceps complex (Squamata:
Chamaeleonidae): Description of four new species from isolated ‘sky islands’ of northern Mozambique. Zootaxa, 3814
(1), 1–36.
https://doi.org/10.11646/zootaxa.3814.1.1

BRANCH ET AL.
158
·
Zootaxa 4324 (1) © 2017 Magnolia Press
Branch, W.R., Haacke, W., Vaz Pinto, P., Conradie, W., Baptista, N. & Verburgt, L. (2017) Loveridge’s Angolan geckos,
Afroedura karroica bogerti and Pachydactylus scutatus angolensis (Sauria, Gekkonidae): new distribution records,
comments on type localities and taxonomic status. Zoosystematics & Evolution, 93, 157–166.
https://doi.org/10.3897/zse.93.10915
Branch, W.R. & Tolley, K.A. (2010) A new chameleon (Sauria: Chamaeleonidae: Nadzikambia) from Mount Mabu, Northern
Mozambique. African Journal of Herpetology, 59, 157–172.
https://doi.org/10.1080/21564574.2010.516275
Branch, W.R. & Whiting, M.J. (1997) A new Platysaurus (Squamata: Cordylidae) from the Northern Cape Province, South
Africa. African Journal of Herpetology, 46, 124–136.
https://doi.org/10.1080/21564574.1997.9649987
Broadley, D.G. (1962) On some reptile collections from the north-western and north-eastern districts of Southern Rhodesia,
1958-61, with descriptions of four new lizards. Occasional Papers of the National Museums of Southern Rhodesia, 3
(26B), 787–843.
Broadley, D.G. (1963) Three new lizards from south Nyasaland and Tete. Annals of the Magazine of Natural History, 6, 285–
288.
https://doi.org/10.1080/00222936308651355
Broadley, D.G. (1966) The herpetology of south-east Africa. Unpublished Ph.D. thesis, University of Natal, Pietermaritzburg,
680 pp.
Broadley, D.G. (1971) A review of Rhampholeon marshalli Boulenger, with the description of a new subspecies from
Mozambique (Sauria: Chamaeleonidae). Arnoldia (Rhodesia), 5, 1–6.
Broadley, D.G. (1978) A revision of the genus Platysaurus A. Smith (Sauria: Cordylidae). Occasional Papers of the National
Museums and Monuments of Rhodesia, Series B, Natural Sciences, 6, 129–185.
Broadley, D.G., Jackman, T.R. & Bauer, A.M. (2014) A review of the genus Homopholis Boulenger (Reptilia: Squamata:
Gekkonidae) in southern Africa. African Journal Herpetology, 63, 109–126.
https://doi.org/10.1080/21564574.2014.930071
Cumberlidge, N., Naskrecki, P. & Daniels, S.R. (2016) Potamonautes gorongosa, a new species of potamonautid freshwater
crab (Decapoda, Brachyura) from Mozambique, southeastern Africa. Nauplius, 24, 1–8.
https://doi.org/10.1590/2358-2936e2016029
Daniels, S.R. & Bayliss, J. (2012) Neglected refugia of biodiversity: Mountainous regions in Mozambique and Malawi yield
three novel freshwater crab species (Potamonautidae: Potamonautes). Zoological Journal of the Linnean Society, 164,
498–509.
https://doi.org/10.1111/j.1096-3642.2011.00773.x
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing.
Nature Methods, 9, 772.
https://doi.org/10.1038/nmeth.2109
De Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation. In: Howard, D.J.
& Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, Oxford, pp. 57–75.
FitzSimons, V.F. (1943) The lizards of South Africa: Systematic account of the lizards (Sauria) of Southern Africa. Transvaal
Museum Memoirs, 1, 1–6.
Frost, D.R. & Hillis, D.M. (1990) Species in concept and practice: herpetological applications. Herpetologica, 46, 87–104.
Griffin, M. (2003) Annotated checklist and provisional national conservation status of Namibian reptiles. Namibia Scientific
Society, Windhoek, 169 pp.
Good, D.A., Bauer, A.M. & Branch, W.R. (1996) A new species of Phyllodactylus (Squamata: Gekkonidae) from the Karoo
National Park, South Africa. African Journal of Herpetology, 45, 49–58.
https://doi.org/10.1080/21564574.1996.9649965
Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT.
Nucleic Acids Symposium, 41, 95–98.
Heinicke, M.P., Daza, J.D., Greenbaum, E., Jackman, T.R. & Bauer, A.M. (2014) Phylogeny, taxonomy and biogeography of a
circum-Indian Ocean clade of leaf-toed geckos (Reptilia: Gekkota), with a description of two new genera. Systematics and
Biodiversity, 12, 23–42.
https://doi.org/10.1080/14772000.2013.877999
Hewitt, J. (1925) On some new species of reptiles and amphibians from South Africa. Records of the Albany Museum
(Grahamstown), 3, 343–370, pls. 1–5.
Huelsenbeck, J.P. & Ronquist, F. (2001) MrBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
https://doi.org/10.1093/bioinformatics/17.8.754
Jacobsen, N.H.G. (1992a) New Lygodactylus taxa (Reptilia: Gekkonidae) from the Transvaal. Bonner zoologische Beiträge, 43,
527–542.
Jacobsen, N.H.G. (1992b) Flat geckos (genus Afroedura) in the Transvaal. Journal of the Herpetological Association of Africa,
40, 22–25.
https://doi.org/10.1080/04416651.1992.9650313
Jacobsen, N.H.G. (1994a) A new subspecies of Lygodactylus ocellatus (Roux)(Lacertilia: Gekkonidae) from the Soutpansberg,
South Africa. Journal of African Zoology, 108, 231–236.

Zootaxa 4324 (1) © 2017 Magnolia Press
·
159
NEW AFROEDURA FROM MOZAMBIQUE
Jacobsen, N.H.G. (1994b) The Platysaurus intermedius complex (Sauria: Cordylidae) in the Transvaal, South Africa, with
descriptions of three new taxa. South African Journal of Zoology, 29, 132–143.
https://doi.org/10.1080/02541858.1994.11448338
Jacobsen, N.H.G. (1997) Family Gekkonidae, Genus Afroedura. In: Van Wyk, J.H. (Ed.), Proceedings of the FitzSimons
Commemorative Symposium, South African Lizards: 50 years of Progress and Third H.A.A. Symposium on African
Herpetology, Transvaal Museum, Pretoria, South Africa, 11–15 October 1993. Herpetological Association of Africa,
Stellenbosch, pp. 34–39.
Jacobsen, N.H.G., Kuhn, A.L., Jackman, T.R. & Bauer, A.M. (2014) A phylogenetic analysis of the southern African gecko
genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and
Mpumalanga provinces of South Africa. Zootaxa, 3846 (4), 451–501.
https://doi.org/10.11646/zootaxa.3846.4.1
Loveridge, A. (1944) New geckos of the genera Afroedura, new genus, and Pachydactylus from Angola. American Museum
Novitates, 1254, 1–4.
Loveridge, A. (1947) Revision of the African lizards of the family Gekkonidae. Bulletin of the Museum of Comparative
Zoology, 98, 1–469, pls. 1–7.
Loveridge, A. (1953) Zoological results of a fifth expedition to East Africa. III Reptiles from Nyasaland and Tete. Bulletin of
the Museum of Comparative Zoology, 110, 141–322, pls. i–v.
Makhubo, B.G., Tolley, K.A. & Bates, M.F. (2015) Molecular phylogeny of the Afroedura nivaria (Reptilia: Gekkonidae)
species complex in South Africa provides insight on cryptic speciation. Molecular Phylogenetics and Evolution, 82, 31–
42.
https://doi.org/10.1016/j.ympev.2014.09.025
Moore, A., Cotterill, F.P.D. & Key, R.M. (2017) A geomorphic and geological framework for the interpretation of species
diversity and endemism in the Manica Highlands. Kirkia, 19 (1), 54–69.
Mouton, P. Le F.N. & Mostert, D.P. (1985) Description of a new species of Afroedura (Loveridge) (Reptilia: Gekkonidae) from
the south-western Cape. South African Journal of Zoology, 20, 246–249.
https://doi.org/10.1080/02541858.1985.11447943
Onderstall, D. (1984) Descriptions of two new subspecies of Afroedura pondolia (Hewitt) and a discussion of species groups
within the genus (Reptilia: Gekkonidae). Annals of the Transvaal Museum, 33, 497–509.
Phiri, E.E. & Daniels, S.R. (2013) Hidden in the highlands: the description and phylogenetic position of a novel endemic
freshwater crab from Zimbabwe. Invertebrate Systematics, 27, 530–539.
https://doi.org/10.1071/IS13012
Raper, P.E. (1989) Dictionary of Southern African Place Names. 2
nd
Edition. Jonathan Ball Publishers, Johannesburg, 608 pp.
Schmitz, A., Brandley, M.C., Mausfeld, P., Vences, M., Glaw, F., Nussbaum, R. A. & Reeder, T.W. (2005) Opening the black
box: phylogenetics and morphological evolution of the Malagasy fossorial lizards of the subfamily “Scincinae”. Molecular
Phylogenetics and Evolution, 34, 118–133.
https://doi.org/10.1016/j.ympev.2004.08.016
Scott, I.A.W., Keogh, J.S. & Whiting, M.J. (2004) Shifting sands and shifty lizards: molecular phylogeny and biogeography of
African flat lizards (Platysaurus). Molecular Phylogenetics and Evolution, 31, 618–629.
https://doi.org/10.1016/j.ympev.2003.08.010
Stamatakis, A. (2006) RAxML-VI-HPC: Maximum Likelihood-based phylogenetic analyses with thousands of taxa and Mixed
Models. Bioinformatics, 22, 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446
Stanley, E.L., Bauer, A.M., Jackman, T.R., Branch, W.R. & Mouton, P. Le F.N, (2011) Between a rock and a hard polytomy:
Rapid radiation in the rupicolous Girdled Lizards (Squamata: Cordylidae). Molecular Phylogenetics Evolution, 58, 53–70.
https://doi.org/10.1016/j.ympev.2010.08.024
Stanley, E., Ceríaco, L.M.P., Bandeira, S., Valerio, H., Bates, M.F. & Branch, W.R. (2016) A new species of Cordylus
(Squamata: Cordylidae) from the ProNamib of south-western Angola. Zootaxa, 4061 (3), 201–226.
https://doi.org/10.11646/zootaxa.4061.3.1
Taylor, P.J., Kearney, T.C., Kerbis Peterhans, J.C., Baxter, R.M. & Willows-Munro, S. (2013) Cryptic diversity in forest shrews
of the genus Myosorex from southern Africa, with the description of a new species and comments on Myosorex tenuis.
Zoological Journal of the Linnean Society, 169, 881–902.
https://doi.org/10.1111/zoj.12083
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: flexible
strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 24, 4876–4882.
https://doi.org/10.1093/nar/25.24.4876
Travers, S.L., Jackman, T.R. & Bauer, A.M. (2014) A molecular phylogeny of Afromontane dwarf geckos (Lygodactylus)
reveals a single radiation and increased species diversity in a South African montane center of endemism. Molecular
Phylogenetics and Evolution, 80, 31–42.
https://doi.org/10.1016/j.ympev.2014.07.017
Whiting, M.J., Branch, W.R., Pepper, M. & Keogh, J.S. (2015) A new species of spectacularly coloured flat lizard Platysaurus
(Squamata: Cordylidae: Platysaurinae) from southern Africa. Zootaxa, 3986 (2), 173–192.
https://doi.org/10.11646/zootaxa.3986.2.2

BRANCH ET AL.
160
·
Zootaxa 4324 (1) © 2017 Magnolia Press
MATERIAL EXAMINED
(not listed in type description)
Afroedura transvaalica: ZIMBABWE: PEM R10069, R10071, R10081-82 (AM 7317, 4 specimens), R10070 (AM 7406),
R10072-79 (AM 6695, 8 specimens), ‘Musami, near Salisbury’ (= 57 km E Harare, 1731DB, 17°44’S, 31°34’E); PEM
R10082-97 (AM 7512, 15 specimens), R12952, Bikita, Masvingo District (2031DB, 20°07’S, 31°43’E); PEM R10098-111
(AM 7250, 13 specimens), Mtoko (= Mutoko, 1732AC ,17°24’S, 32°13’E); PEM R12953-55 (AMR 0155, 3 specimens),
Mtoko (17°02'S, 32°01'E), PEM R12956-57 (AMR 0155, 2 specimens), Empandeni (20°03'S, 27°04'E); PEM R10112 (AM
7250), Driefontein (Mission), Gweru District (19°25’S, 30°42’E, 1930BD). SOUTH AFRICA: PEM R16072-73 (AM 5114, 2
specimens, co-types Oedura transvaalica Hewitt, 1925), ‘Nijelele River, Soutpansberg’, Limpopo Province (2229CC); PEM
R20786, 10 km south of Messina, Limpopo Province (22°28’27’’S, 29°59’25”E).
Afroedura loveridgei: ZMB 74437, 74384-85, “Mozambique”; PEM R8208-09, 8334-39 (eight specimens), 7 km west Songa
turn-off, Tete Province, Mozambique (16°20'49"S, 33°27'89"E, 271 m asl); PEM R21992
(EI 0191), 15 km east Moatize,
Mozambique (16
°
06'01" S, 33
°
53'04.52" E).