Content uploaded by Jessica Marie da Silva
Author content
All content in this area was uploaded by Jessica Marie da Silva on Dec 24, 2013
Content may be subject to copyright.
Accepted by S. Carranza: 19 Nov. 2013; published: 23 Dec. 2013
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3750 (5): 465–493
www.mapress.com
/
zootaxa
/
Article
465
http://dx.doi.org/10.11646/zootaxa.3750.5.3
http://zoobank.org/urn:lsid:zoobank.org:pub:DC8E9834-EBFE-41EC-91D1-69EE0ED2DDF5
A molecular phylogeny of the African plated lizards, genus Gerrhosaurus
Wiegmann, 1828 (Squamata: Gerrhosauridae), with the description of two new
genera
MICHAEL F. BATES1,7, KRYSTAL A. TOLLEY2,3, SHELLEY EDWARDS2,3, ZOË DAVIDS2,
JESSICA M. DA SILVA2,4 & WILLIAM R. BRANCH5,6
1 Department of Herpetology, National Museum, P.O. Box 266, Bloemfontein 9300, South Africa. E-mail: herp@nasmus.co.za
2 Applied Biodiversity Research Division, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South
Africa. E-mail: K.Tolley@sanbi.org.za, S.Edwards@sanbi.org.za, Z.Davids@sanbi.org.za, jessica.m.dasilva@gmail.com
3 Department of Botany and Zoology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa
4 Department of Conservation Ecology and Entomology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa
5 Department of Zoology, Nelson Mandela Metropolitan University, P.O. Box 77000, Port Elizabeth 6031, South Africa
6 Department of Herpetology, Port Elizabeth Museum, P.O. Box 13147, Humewood 6013, South Africa.
E-mail: wrbranch@bayworld.co.za
7 Corresponding author: E-mail: herp@nasmus.co.za
Abstract
We constructed a molecular phylogeny of the African plated lizard family Gerrhosauridae using two mitochondrial markers
(ND2, 732 bp; 16S, 576 bp) and one nuclear marker (PRLR, 538 bp). This analysis showed that the subfamily Gerrhosaurinae
consists of five major clades which we interpret as representing five genera. The genera Tetradactylus and Cordylosaurus were
each recovered as monophyletic, but Gerrhosaurus as currently conceived is paraphyletic, consisting of three distinct genus-
level assemblages. The two clades consisting of Gerrhosaurus major Duméril, 1851 and Gerrhosaurus validus Smith, 1849 are
both described here as new genera, namely Broadleysaurus Bates & Tolley gen. nov. and Matobosaurus Bates & Tolley gen.
nov., respectively. Two subspecies of ‘Gerrhosaurus major’ that were historically separated on the basis of differences in
colour pattern are not reciprocally monophyletic, so Gerrhosaurus bottegoi Del Prato, 1895 is relegated to the synonomy of
Broadleysaurus major (Duméril, 1851) comb. nov., which is rendered monotypic. Gerrhosaurus validus maltzahni De Grys,
1938 is genetically and morphologically well differentiated from G. v. validus and the two taxa also occur in allopatry. We
therefore re-instate the former as Matobosaurus maltzahni (De Grys, 1938) comb. nov., rendering Matobosaurus validus
(Smith, 1849) comb. nov. a monotypic species. Our analysis also showed that Gerrhosaurus sensu stricto comprises two major
subclades, one consisting of Gerrhosaurus typicus (Smith, 1837) + Gerrhosaurus skoogi Andersson, 1916, and the other
containing the remaining species. In this latter subclade we show that west-Central African Gerrhosaurus nigrolineatus
Hallowell, 1857 is most closely related to Gerrhosaurus auritus Boettger, 1887 rather than to G. nigrolineatus from East and
Southern Africa. The west-Central African clade of G. nigrolineatus differs from the East and Southern African clade by a p-
distance of 13.0% (ND2) and 6.9% (16S), and can be differentiated morphologically. We accordingly apply the name
Gerrhosaurus intermedius Lönnberg, 1907 comb. nov. to populations from Kenya, Uganda, Rwanda, Tanzania, Malawi,
Mozambique, Zimbabwe and South Africa previously identified under the name G. nigrolineatus. Our analysis also confirms
that Gerrhosaurus bulsi Laurent, 1954 is a distinct species and sister taxon to a clade containing G. nigrolineatus, G. auritus and
G. intermedius. The latter four taxa form a closely-related ‘G. nigrolineatus species complex’ with a widespread distribution in
Africa. Most closely related to this complex of species is Gerrhosaurus flavigularis Wiegmann, 1828 which has an extensive
range in East and Southern Africa, and displays genetic substructure which requires further investigation. The status of
Gerrhosaurus multilineatus Bocage, 1866, and Angolan populations referred to G. nigrolineatus, remains problematic.
Key words: Sauria, Cordyliformes, Gerrhosaurinae, systematics, taxonomy, phylogeography
Introduction
The lizard families Gerrhosauridae and Cordylidae together constitute the clade Cordyliformes (Lang 1991). There
exists a long history of disagreement among authors as to whether this clade comprises a single family, the
BATES ET AL.
466 · Zootaxa 3750 (5) © 2013 Magnolia Press
Cordylidae, without mention of subfamilies (e.g. Odierna et al. 2002―molecular and karyological data); two
families, namely Cordylidae and Gerrhosauridae (e.g. Loveridge 1942; FitzSimons 1943; Lang 1991―morphology);
or one family with two subfamilies, namely Cordylinae and Gerrhosaurinae (e.g. Wermuth 1968; Auerbach
1987―morphology). However, the molecular analyses of Frost et al. (2001), Lamb et al. (2003) and Lamb & Bauer
(2013) have confirmed the monophyly of the two families, and revealed sufficient genetic divergence for their
acceptance.
The Gerrhosauridae consists of two subfamilies: the Gerrhosaurinae from mainland sub-Saharan Africa, and
the Zonosaurinae from Madagascar (including the offshore islands of Comoros, Gloriosa and Cosmoledo) (Lang
1991). Gerrhosaurinae consists of three genera, namely Gerrhosaurus Wiegmann (nine species), Tetradactylus
Merrem (eight species; including T. fitzsimonsi Hewitt which differs from T. africanus [Gray] by lacking
hindlimbs, and the two taxa are separated by over 500 km), and Cordylosaurus Gray (one species), while the
Zonosaurinae currently contains two genera, namely Zonosaurus Boulenger (17 species) and Tracheloptychus
Peters (two species) (Branch 1998; Broadley 2007; Glaw & Vences 2007; Uetz 2013; Bates et al. in press.),
although the latter may be embedded within the former (Raselimanana et al. 2009). A recent multi-gene phylogeny
by Recknagel et al. (2013) indicated the existence of five species groups within Zonosaurinae, but could not
resolve the generic status of Tracheloptychus.
Three species of Gerrhosaurus were described in 1837 in the genus Pleurotuchus Smith, but only the type
species Pleurotuchus typicus Smith, 1837―referred to the genus Gerrhosaurus by Duméril & Bibron (1839) soon
after its description―remains valid. The other two taxa, Pleurotuchus desjardinii Smith, 1837 and Pleurotuchus
chrysobronchus Smith, 1837, are junior synonyms of Gerrhosaurus flavigularis Wiegmann, 1828 (see Loveridge
1942). The genus Angolosaurus was erected by FitzSimons (1953) to accommodate the spade-snouted
deserticolous ecomorph Gerrhosaurus skoogi Andersson, 1916. A phylogeny based on mitochondrial markers
(Lamb et al. 2003) showed that the latter species was nested within, and should be returned to, the genus
Gerrhosaurus. Detailed analysis of the cranial osteology of G. skoogi and a parsimony analysis of 118 of these
characters within cordyliforms generated a phylogeny in which this species was identified as sister to all other
gerrhosaurids, including the Malagasy radiation (Nance 2003, 2007). Cautioning against generic re-assignments
based only on mitochondrial gene analysis, Nance (2007) proposed continued recognition of Angolosaurus. This
problem was revisited by Lamb & Bauer (2013) and mitochondrial, nuclear and combined gene analyses
corroborated and strengthened their earlier findings (Lamb et al. 2003), and also emphasized that the unique
morphological features of G. skoogi were autapomorphies associated with the species’ psammophilous habits.
Most species in the genus Gerrhosaurus have convoluted and complex taxonomic histories, although no new
taxa have been described for over 50 years. As many as 29 species and subspecies have been described (see
Loveridge 1942; Laurent 1954; Hellmich & Schmelcher 1956; Broadley 1960), but there remains considerable
confusion and difference of opinion regarding the status of several species. For example, in his description of
Gerrhosaurus multilineatus, Bocage (1866a) noted that this form may be a well-characterised variety of
Gerrhosaurus nigrolineatus Hallowell, 1857, and Loveridge (1942) subsequently considered it a mere colour
variant of the latter species. FitzSimons (1943) also treated G. multilineatus as a synonym of G. nigrolineatus.
Loveridge (1942) considered Gerrhosaurus auritus Boettger, 1887 to be a subspecies of G. nigrolineatus, while
FitzSimons (1943) and Mertens (1955) treated it as a full species. De Witte (1953) also treated G. auritus as a full
species, but his specimens were in fact referable to G. b u l s i , described as Gerrhosaurus auritus bulsi by Laurent
(1954) the following year. Broadley (1971) considered G. a. bulsi to be a synonym of G. multilineatus―which he
recorded from north-eastern Zambia―but it was later treated as a valid species (Haagner et al. 2000; Broadley &
Cotterill 2004; Broadley 2007). Confusion about the status of the above-mentioned three species is due partly to
the fact that the type series of the poorly-known G. multilineatus was destroyed in the 1978 fire at Museu Bocage in
Lisbon (Almaca & Neves 1987; Madruga 2012) and it has been difficult to establish the affinities and identities of
these taxa because of the lack of museum material, especially from Angola. As many as four subspecies of
Gerrhosaurus major Dumeril, 1851 were recognized by Loveridge (1942), but Broadley (1987) reviewed these and
came to the conclusion that only two subspecies were valid―a mainly tan-backed eastern form (Gerrhosaurus
major major) and a mainly black-backed form with rows of yellow dorsal spots (Gerrhosaurus major bottegoi Del
Prato, 1895). Juveniles of both subspecies are black dorsally with yellow speckling, but adults of the latter retain
the juvenile colour pattern as adults, except that the speckling becomes more stripe-like (Spawls et al. 2002).
The following nine species of Gerrhosaurus are currently recognized: G. flavigularis Wiegmann, 1828; G.
typus (Smith, 1837); G. validus Smith, 1849; G. major Duméril, 1851; G. nigrolineatus Hallowell, 1857; G.
Zootaxa 3750 (5) © 2013 Magnolia Press · 467
MOLECULAR PHYLOGENY OF GERRHOSAURUS
multilineatus Bocage, 1866a; G. auritus Boettger, 1887; G. skoogi Andersson, 1916; and G. b u l s i Laurent, 1954
(Adolphs 2006, 2013; Broadley 2007). Only G. validus (G. v. validus and G. v. maltzahni De Grys, 1938) and G.
major (G. m. major and G. m. bottegoi Del Prato, 1895) contain currently recognized subspecies.
Plated lizards are medium-sized (G. typicus attains a snout-vent length [SVL] of 140 mm) to large (G. validus
SVL 285 mm), diurnal, mainly insectivorous (vegetable matter is also eaten by some species) and oviparous
(Broadley 1966; Loveridge 1942; FitzSimons 1943; Branch 1998). Gerrhosaurus validus (rupicolous) and G.
skoogi (deserticolous) may form loosely-structured colonies, but the other species are usually solitary and mainly
terrestrial. Plated lizards often use burrows (at the base of a bush) for shelter, although the spade-snouted G. skoogi
frequently dives under sand in the Namib Desert (Loveridge 1942; FitzSimons 1953; Visser 1984a,b; Jacobsen
1989; Branch 1998). Gerrhosaurus typicus occurs only in the south-western part of the continent and was
considered Near Threatened by the IUCN (1996), but it was recently downgraded to Least Concern (Bates et al. in
press.). Gerrhosaurus skoogi is restricted to the Namib Desert in north-western Namibia and south-western
Angola, but most other species (range of G. multilineatus is unclear) are fairly widespread (e.g. Loveridge 1942;
Branch 1998) and none are currently regarded as being of conservation concern (e.g. Bates et al. in press.).
Lamb et al. (2003) conducted a molecular phylogenetic analysis of African and Malagasy gerrhosaurid genera
based on mitochondrial DNA sequence data. Their analysis included most known gerrhosaurid species from
mainland Africa, including seven of the currently recognized species (two with two subspecies each) of
Gerrhosaurus as listed above. They did not include samples of G. multilineatus, perhaps following Loveridge
(1942) who treated this species as a synonym of G. nigrolineatus, nor did they include samples of G. b ul s i
(described as G. auritus bulsi Laurent) which they may have considered a synonym of G. a u r i t u s . Among the
mainland African gerrhosaurids, three major clades emerged (Lamb et al. 2003). Gerrhosaurus major was shown
to be the sister clade to all other taxa, including other Gerrhosaurus which formed the sister clade to
Cordylosaurus/Tetradactylus. Lamb et al. (2003) suggested that the latter two genera may have evolved from
within Gerrhosaurus, but this relationship lacked statistical support at the relevant nodes. They also suggested that
the stout-bodied G. major constituted a separate genus, but indicated that verification would require a combination
of molecular and morphological data. Angolosaurus was shown to be nested within Gerrhosaurus and was
accordingly transferred to the latter genus. This and the other relationships noted by Lamb et al. (2003) were
corroborated and strengthened in a subsequent study―with the same taxa but a slightly supplemented
dataset―using combined multilocus mitochondrial, nuclear and combined gene analyses (Lamb & Bauer 2013).
We concur with the opinions, recommendations and proposals of Kaiser et al. (2013) regarding best taxonomic
practice, and therefore reject the unscientific taxonomy of Hoser (2013). We consider the nomenclatural changes to
the families Gerrhosauridae and Cordylidae proposed therein (Hoser 2013) to be ill-conceived and unethical, and
thus unavailable. We note specifically that the privately-published and personally-edited work of Hoser (2013) is in
direct violation of the spirit and intention of the Code (ICZN 1999) as indicated by, inter alia, Recommendation 8A
which explicitly encourages publication in “appropriate scientific journals or well-known monographic series”.
To examine evolutionary relationships and systematics of the Gerrhosauridae, we constructed a near-
comprehensive species-level phylogeny for the genus Gerrhosaurus with representative taxa from all other genera
in the family. The only species of Gerrhosaurus not included was G. multilineatus, which is known only from the
type locality, “district [presumably ‘region’] of Duque de Bragança, interior of Angola” (Bocage 1866a: 61).
Materials and methods
Samples. To determine the taxonomic placement of species within the Gerrhosauridae, multiple individuals from
across the range of species were (where possible) used, and samples collected near type localities were included
when available (Table 1). Data on type localities and geographical ranges of species are provided in the Discussion
below. Several samples used in the analysis are associated with vouchers housed at two South African museums:
Port Elizabeth Museum, Bayworld (examined by WRB), and National Museum, Bloemfontein (MFB).
In order to substantiate the taxonomic implications of our molecular phylogeny, we attempted to seek
morphological congruence for the various lineages. Specimens with matching tissue samples were identified using
diagnostic keys and other scalation and colour pattern data in FitzSimons (1943), Broadley (1971), Laurent (1954
& 1964, G. bulsi) and Branch (1998). Head length was measured from tip of snout to nearest part of ear opening. A
few additional specimens of G. nigrolineatus from Republic of the Congo and Gabon, not used in the molecular
BATES ET AL.
468 · Zootaxa 3750 (5) © 2013 Magnolia Press
analysis, were examined (by WRB) for comparison (Appendix I). Colour photographs of a few specimens of G.
nigrolineatus from Kouilou region, Republic of Congo, collected with the sampled specimens (including MBUR
02993, see Table 1; MBUR 02986, Tchiboula region; and PEM R20067, Appendix I), and images of the two
syntypes of G. nigrolineatus from Gabon in the collection of the Academy of Natural Sciences, Philadelphia
(ANSP 3729, adult; ANSP 8825, juvenile), were also examined (by MFB) to confirm the status of the latter species
in west-Central Africa. Voucher specimens of G. nigrolineatus from Republic of the Congo used for the molecular
analysis (see Table 1) are housed in the GERDIB (Groupe d’Etude et de Recherche sur la Diversité Biologique)
collection in Brazzaville, but could not be obtained on loan. Museum specimens from south-east Africa referable to
‘G. intermedius’ (see discussion below) were examined by WRB (Mozambique) and MFB (Limpopo Province,
South Africa), and data for specimen TM 80959 in the Ditsong National Museum of Natural History (Pretoria) was
provided by L. Mahlangu (Appendix I). Details of colour pattern, size and scalation are, where applicable,
presented in the discussion below.
In some cases tissue samples (tail tips) were removed and specimens released. For some of these latter
individuals, photographic evidence (examined by MFB and/or WRB) was used for follow-up identification; or the
taxa were readily identifiable in the field. The identity of most individuals for which Genbank sequences were used
was not checked, but only one sequence (AMB 8339 = G. flavigularis, not G. nigrolineatus as indicated by Stanley
et al. 2011; see Fig. 1 & Table 1) was represented in an unlikely position in the phylogeny.
Phylogenetic analyses. Tail tips or liver were removed from 51 individuals of the Gerrhosauridae (Table 1)
and stored in 70–99% ethanol. Total genomic DNA was extracted using standard salt extraction (Bruford et al.
1992) and portions of two mitochondrial markers (ND2, 732 bp and 16S, 576 bp) and one nuclear marker (PRLR,
538 bp) were sequenced. Amplifications were carried out in 25 µl reaction volumes containing 2 µl of extract (ca.
25 ng/µl), 0.25 µM of each primer, 0.2 mM dNTPs, 2.5 mM MgCl2, 1 X thermophilic Buffer (50 mMKCl, 10
mMTris–HCl, pH 9) and 0.25 unitSuper-ThermTaq polymerase using primers vMet and vTrp for ND2
(Cunningham & Cherry 2004), L2510 and H3080 for 16S (Palumbi et al. 1991), and F2 with R4 for PRLR (Portik
et al. 2012). The PCR profile was 95°C for 1 min, followed by 35 cycles of 35 s at 95°C, 30 s at 50–55°C and 1 min
at 72°C, with a final extension at 72°C for 30 s. PCR products were visualised on 0.8% agarose gels containing
ethidium bromide, and cycle-sequenced using a fluorescently-labelled dye-terminator kit (ABI, Foster City,
California, USA), purified with Sephadex spin columns, and analysed at Macrogen, Seoul Korea. Sequence
alignment was carried out in Geneious Pro v.4.8.5 (Drummond et al. 2009) using default parameters, and all gaps
were treated as missing data. All new sequences were deposited in GenBank (accession numbers KF717375 to
717496, Table 1).
The dataset also included 18 samples of gerrhosaurid taxa and two samples of outgroup taxa (Cordylus
cordylus and Smaug giganteus) available from GenBank, for a total of 71 samples (Table 1). Prior to analysis, the
partition homogeneity test was run in PAUP* v. 4.0 (Swofford 2002) to examine whether the two genomes
provided different phylogenetic signals, but this test indicated no conflict (P = 0.81), so the combined dataset was
used to produce a single phylogeny. Bayesian inference (BI) was run using MrBayes v.3.1.2 (Ronquist &
Huelsenbeck 2003) via remote upload at the CBSU cluster (cbsuapps.tc.cornell.edu). Prior to this analysis,
jModelTest v.0.0.1 (Posada 2008) was run for each marker to investigate the evolutionary model that best fits the
dataset using the AIC criterion. This suggested that the best fit was a model with six rate categories for each of the
markers: GTR I+G for 16S and ND2, GTR+G for PRLR. The phylogeny was therefore estimated with three data
partitions (one for each marker), uniform priors for all parameters, with each partition allowed to run
independently. The analysis was also run with data partitions based on codons (1+2, 3) for the two coding genes,
again using the models indicated by jModelTest: GTR I+G for both ND2(1+2) and ND2(3), GTR+G for both
PRLR(1+2) and PRLR(3). For each partitioning scheme, the MCMC was run twice in parallel for 10 million
generations, sampling trees every 1000 generations. Burn-in was determined by examining stationarity of log-
likelihood tree scores, and also ensuring that standard deviation of split frequencies approached <0.001. In
addition, we confirmed that the effective sample size (ESS) was more than 200 for all parameters (Tracer v.1.4.1:
Rambaut & Drummond 2007). For all runs, the first one million generations (1000 trees) was removed as burn-in
before constructing a 50% majority rule tree. Nodes with posterior probability >0.95 were considered supported.
A maximum likelihood (ML) search was run in RAXML v.7.2.7 (Stamatakis 2006; Stamatakis et al. 2008) via
CIPRES Science Gateway v.3.1 (http://www.phylo.org/sub_sections/portal/), with a search for the best scoring ML
tree, plus rapid bootstrapping. Three data partitions were set up (one for each gene), each with the GTR+I+G model
of evolution. This analysis was run three times to ensure that independent ML searches produced the same topologies.
Zootaxa 3750 (5) © 2013 Magnolia Press · 469
MOLECULAR PHYLOGENY OF GERRHOSAURUS
TAB L E 1 . Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of
Sciences, NMB = National Museum, Bloemfontein, NMZB = National History Museum of Zimbabwe, PEM = Port
Elizabeth Museum/Bayworld, TM = Ditsong National Museum of Natural History, UMMZ = University of Michigan
Museum of Zoology, USEC = University of Stellenbosch Ecology Collection) and GenBank accession numbers (16S,
ND2, PRLR) for individuals used in this study. Tetradactylus fitzsimonsi is treated as a full species following Bates et al.
(in press.).
Genus Species/
subspecies
Sample
number
Vo u c h e r
number
16S ND2 PRLR Locality
Cordylosaurus subtessellatus HB 007 KF717379 KF717425 NA Rooiberg, Western Cape, South Africa
Cordylosaurus subtessellatus HB 180 KF717380 KF717426 KF717466 Nieuwodtville, Northern Cape, South Africa
Cordylosaurus subtessellatus AMB 4649 HQ167167 HQ166956 HQ167496 near The Black Hills, Northern Cape, South
Africa (GenBank)
Cordylosaurus subtessellatus AMB 6928 HQ167168 HQ166957 HQ167497 7 km NW of Sesfontein, Namibia (GenBank)
Gerrhosaurus auritus NMZB
16027
NMZB
16027
AY167368 AY167402 KC954131 Ndau School, Western Province, Zambia
(GenBank)
Gerrhosaurus bulsi ANG 0116 PEM
R19479
KF717381 KF717427 KF717467 Luachimo village, L unda Norte, Angola
Gerrhosaurus bulsi ANG 0120 PEM
R19475
KF717382 KF717428 NA Lake Carumbo base camp, Lunda Norte,
Angola
Gerrhosaurus bulsi PEM
R18886
PEM
R18886
KF717383 KF717429 NA Kalumbila Village, Northwest Province,
Zambia
Gerrhosaurus flavigularis AMB 8339 HQ167245 HQ167023 HQ167574 62 km WNW of Mokopane, L impopo
Province, South Africa (GenBank)
Gerrhosaurus flavigularis ELS 020 KF717384 KF717430 KF717468 Malolotja, Swaziland
Gerrhosaurus flavigularis HB 056 KF717385 KF717431 KF717469 Handeni, Tanzania
Gerrhosaurus flavigularis JM 03176 KF717386 KF717432 KF717470 Mwatate, Keny a
Gerrhosaurus flavigularis MB 21407 PEM
R20379
KF717387 KF717433 KF717471 9 km SE of Dithakong, North West Province,
South Africa
Gerrhosaurus flavigularis MBUR
00201
PEM
R20380
KF717388 NA KF717472 Dullstroom area, Mpumalanga Province,
South Africa
Gerrhosaurus flavigularis MBUR
00249
PEM
R20381
KF717389 KF717434 KF717473 Malebogo Nat. Res., Limpopo Province,
South Africa
Gerrhosaurus flavigularis MBUR
00328
PEM
R20383
KF717390 KF717435 KF717474 Makgabeng area, Limpopo Province, South
Africa
Gerrhosaurus flavigularis MBUR
00378
PEM
R20387
KF717391 KF717436 KF717475 Makgabeng area, Limpopo Province, South
Africa
Gerrhosaurus flavigularis MBUR
00456
PEM
R20388
KF717392 KF717437 KF717476 Zitapile, Eastern Cape, South Africa
Gerrhosaurus flavigularis MBUR
00469
PEM
R20389
KF717393 KF717438 KF717477 Zitapile, Eastern Cape, South Africa
Gerrhosaurus flavigularis MBUR
00841
NMB
R10595
NA KF717439 KF717478 near Loo le Dam, Limpopo Province, South
Africa
Gerrhosaurus flavigularis MBUR
00947
NMB
R10890
KF717394 KF717440 KF717479 Tshipise area, Limpopo Province, So uth
Africa
Gerrhosaurus flavigularis PEM
R14985
PEM
R14985
KF717395 KF717441 KF717480 Bazarutho, Mozambique
Gerrhosaurus flavigularis PEM
R15454
PEM
R15454
KF717396 KF717442 NA Zambezi delta, Mozambique
Gerrhosaurus major bottegoi None AY167363 AY167397 KC954119 Atakpame, Togo (GenBank)
Gerrhosaurus major major HB 052 KF717375 KF717422 KF717462 Dodoma, Tanzania
Gerrhosaurus major major HB 054 KF717376 KF717423 KF717463 Arusha, Tanzania
Gerrhosaurus major major HB 055 KF717377 NA KF717464 Dodoma, Tanzania
Gerrhosaurus major major MBUR
01079
NMB
R10892
KF717378 KF717424 KF717465 Tshipise area, Limpopo Province, So uth
Africa
Gerrhosaurus major major NMZB
16199
NMZB
16199
AY167362 AY167396 KC954117 Chituripasi, Chipise, Zimbabwe (GenBank)
Gerrhosaurus nigrolineatus MBUR
02975
KF717399 KF717447 NA 4.0 km NE of Lo ulema, Kouil ou Dpt, Rep.
Congo
Gerrhosaurus nigrolineatus MBUR
02981
KF717400 KF717448 NA 3.9 km ESE of Loubanguila, Kouilou Dpt,
Rep. Congo
......continued on the next page
BATES ET AL.
470 · Zootaxa 3750 (5) © 2013 Magnolia Press
TAB L E 1 . (Continued)
Genus Species/
subspecies
Sample
number
Vo u c h e r
number
16S ND2 PRLR Locality
Gerrhosaurus nigrolineatus MBUR
02985
NA KF717449 KF717483 4.2 km SW of Tchizondi, Kouilou Dpt, Rep.
Congo
Gerrhosaurus nigrolineatus MBUR
02993
KF717402 KF717451 NA Tchiboula region, Kouilou Dpt, Rep. Congo
Gerrhosaurus nigrolineatus MBUR
02990
KF717401 KF717450 NA 5.5 km WNW of Tchizondi, Kouilou Dpt,
Rep. Congo
Gerrhosaurus nigrolineatus AJC 461 KF717397 KF 717443 NA Limpopo Province, South Africa
Gerrhosaurus nigrolineatus AJC 462 KF717398 KF 717444 NA Limpopo Province, South Africa
Gerrhosaurus nigrolineatus HB 057 NA KF717445 KF717481 Arusha, Tanzania
Gerrhosaurus nigrolineatus HB 061 NA KF 717446 KF717482 W of Skukuza, Mpumalanga Province, South
Africa
Gerrhosaurus nigrolineatus TM 80959 TM 80959 AY167367 AY167401 KC954130 Moebase Village, Mozambique (GenBank)
Gerrhosaurus skoogi CAS
206978
CAS
206978
AY167364 AY167398 KC954127 Ondonduiengo River, Namibia (GenBank )
Gerrhosaurus typicus KTH 05-06 KF717403 KF717452 NA Bo-Kruisfo ntein, Western Cape, South Africa
Gerrhosaurus typicus USEC-
H3504
USEC-
H3504
AY167365 AY167399 NA Unknown (GenBank)
Gerrhosaurus validus
validus
MBUR
00290
PEM
R20382
KF717407 NA KF717487 Makgabeng area, Limpopo Province, South
Africa
Gerrhosaurus validus
validus
MBUR
00892
KF717408 NA KF717488 Phalaborwa area, Limpopo Province, South
Africa
Gerrhosaurus validus
validus
MBUR
00901
NMB
R10645
KF717409 KF717456 KF717489 Phalaborwa area, Limpopo Province, South
Africa
Gerrhosaurus validus
validus
MBUR
01010
NMB
R10893
KF717410 KF717457 KF717490 Greater Kuduland Safaris, E of Tshipise,
Limpopo Pro vince, South Africa
Gerrhosaurus validus
validus
PEM R5935 PEM
R5935
KF717412 KF717459 NA Niassa, Mozambique
Gerrhosaurus validus
validus
RSP 352 KF717413 KF717460 NA Venitia Nat. Res., Limpopo Province, South
Africa
Gerrhosaurus validus
validus
MBUR
2809
KF717411 KF717458 KF717491 South Africa
Gerrhosaurus validus
validus
NMZB
16268
NMZB
16268
AY167361 AY167395 NA Chikwakwara, Chipise, Zimbabwe
(GenBank)
Gerrhosaurus validus
validus
AMB 6090 HQ167246 HQ167024 HQ16 7575 13 km NW of Hoe dspruit, Limpopo Provinc e,
South Africa (Genbank)
Gerrhosaurus validus
validus
ELS 004 KF717406 KF717455 KF717486 N Lebombo Mtns, Limpopo Province, South
Africa
Gerrhosaurus validus
maltzahni
CAS
206935
CAS
206935
AY167360 AY167394 KC954126 Usakos, Namibia (GenBank)
Gerrhosaurus validus
maltzahni
KTH09-
289A
PEM
R17984
KF717404 KF717453 KF717484 0.5 km S of Tambor, Iona, Angola
Gerrhosaurus validus
maltzahni
KTH09-
290A
PEM
R17985
KF717405 KF717454 KF717485 Omauha Lodge, Iona, Angola
Tetradactylus fitzsimonsi PEM
R14913
PEM
R14913
AY167370 AY167404 KC954123 13 km WNW of Humansdo rp, Eastern Cape,
South Africa (GenBank)
Tetradactylus fitzsimonsi PEM
R14912
PEM
R14912
KF717414 NA NA 13 km WNW of Humansdorp, Eastern Cape,
South Africa
Tetradactylus seps PEM
R17868
PEM
R17868
KF717418 KF717461 KF717495 Martin's River, Elandsberg, Eastern Cape,
South Africa
Tetradactylus seps USEC-
H3505
USEC-
H3505
AY167369 AY167403 KC954122 GenBank
Tetradactylus seps HB 083 KF717415 NA KF717492 Upper Kromme River, Eastern Cape, South
Africa
Tetradactylus seps HB 084 KF717416 NA KF717493 Upper Kromme River, Eastern Cape, South
Africa
Tetradactylus seps HB 085 KF717417 NA KF717494 near Joubertine dam, Eastern Cape, South
Africa
Tetradactylus tetradactylus HB 011 KF717419 NA NA Einsinheid, Western Cape, South Africa
......continued on the next page
Zootaxa 3750 (5) © 2013 Magnolia Press · 471
MOLECULAR PHYLOGENY OF GERRHOSAURUS
Nodes with a bootstrap value of >70% were considered supported in this analysis. Finally, sequence divergences
(uncorrected p-distances) between taxa were estimated using MEGA v.5.05 (Tamura et al. 2011).
Competing phylogenetic hypotheses of monophyly of species were investigated using a Shimodaira–
Hasegawa (SH) test (Shimodaira & Hasegawa 1999; Goldman et al. 2000) and the approximately unbiased (AU)
test (Shimodaira 2002) generating maximum likelihood scores for the trees (1000 replicates) using PAUP*
v.4.0b10 (Swofford 2002) and bootstrapping P values for the SH and AU tests in Consel (Shimodaira 2002).The
obtained maximum likelihood topology (see Results) was compared to a topology which constrained 1) G.
nigrolineatus to be monophyletic to the exclusion of G. auritus and G. bulsi and 2) Gerrhosaurus to be
monophyletic to the exclusion of Tetradactulus and Cordylosaurus.
Results
Both phylogenetic methods (ML and BI) produced the same topology with similar node support (Fig. 1). Several
taxonomic issues are notable within the phylogeny. Firstly, the phylogenetic analysis confirmed that the
Gerrhosaurinae (clades A to E) and Zonosaurinae (clade F) are sister taxa. Secondly, it revealed that Gerrhosaurus
is not monophyletic. In particular, two separate clades―G. major (clade E) and G. validus (clade D)―fall outside a
larger well supported clade containing all other Gerrhosaurus taxa (clade A) as well as Cordylosaurus (clade B)
and Tetradactylus (clade C). The latter three groups are all well-defined and deeply divergent. Gerrhosaurus major
differs from all gerrhosaurine taxa and populations by uncorrected p-distances of at least 19.4% (ND2) and 8.4%
(16S), while each of the two subspecies of G. v a l i d u s differs from the other taxa and populations by a minimum of
21.3% and 6.7% respectively (Table 2). Both the SH and AU tests reject a monophyletic Gerrhosaurus as presently
defined (obtained topology: –lnL = 15750.52, constrained topology: –lnL = 15769.64, SH test: P = 0.05; AU test:
P = 0.004) in favour of the observed topology.
One lineage of G. major was recovered suggesting the presence of a monophyletic species, without genetic
differentiation between the subspecies. The two subspecies of G. validus, however, are genetically distinct (Fig. 1).
While they are supported as sister clades within the phylogeny, sequence divergences (8.5% for ND2, 4.1% for
16S; Table 2) between these clades are similar to that usually found between other reptile species for these markers
(e.g. Tolley et al. 2006; Tilbury & Tolley 2009; Conradie et al. 2012).
In the Gerrhosaurus clade (A) each species is strongly supported, and there is also strong support for the
relationships between species (Fig. 1). The G. flavigularis clade contains at least three subclades, corresponding to
populations in Eastern South Africa, Northern South Africa and East Africa. Specimens identified as G.
nigrolineatus from East/southern Africa are supported as a monophyletic clade, but this clade is not the sister group
of G. nigrolineatus from west-Central Africa, which is more closely related to G. auritus. Therefore, G.
nigrolineatus as currently defined is not monophyletic. Sequence divergences between the west-Central Africa
versus East/Southern Africa clades of G. nigrolineatus are high (13.0% for ND2, 6.9% for 16S) and exceed that
TAB L E 1 . (Continued)
Genus Species/
subspecies
Sample
number
Vo u c h e r
number
16S ND2 PRLR Locality
Tetradactylus tetradactylus HB 012 KF717420 NA NA Einsinheid, Western Cape, South Africa
Tetradactylus tetradactylus RSP 205 KF717421 NA KF717496 Kammanassie Mountain, Western Cape,
South Africa
Tracheloptychus petersi UMMZ-
207157
UMMZ-
207157
AY167373 AY167407 KC954115 8 km from Ntolaria, N of Fitheranana River,
Madagascar (GenBank)
Tracheloptychus petersi AY140996 NA NA Madagascar (Genbank)
Zonosaurus laticaudatus UMMZ-
RAN 55301
UMMZ-
RAN55301
AY167372 AY167406 KC954116 7.5 km ENE of Hazofotsy, Madagascar
(GenBank)
Zonosaurus trilineatus None AJ416934 AY662560 NA consensus sequence, Madagascar (GenBank)
Outgroup
Cordylus cordylus PEM
R17464
PEM
R17464
HQ167190 HQ166956 HQ167519 24 km SE of Steytlerville, Eastern Cape,
South Africa (GenBank)
Smaug giganteus QP 0037
(MJC 5403)
HQ167193 HQ166981 HQ167522 5 km W of Harrismith, Free State, South
Africa (GenBank)
BATES ET AL.
472 · Zootaxa 3750 (5) © 2013 Magnolia Press
which is generally found between reptile species (e.g. Tolley et al. 2006; Tilbury & Tolley 2009; Conradie et al.
2012). Gerrhosaurus bulsi is shown to be a distinct species and the sister taxon to the G. nigrolineatus (west-
Central Africa)–G. auritus–G. nigrolineatus (East/Southern Africa) clade (Fig. 1). Neither the SH or AU test could
reject a monophyletic G. nigrolineatus as presently defined (observed topology: –lnL = 15750.52, constrained
topology –lnL = 15759.82, SH test: p = 0.23; AU test: P = 0.10) in favour of the observed topology, although the
observed topology has a better log-likelihood score than the constrained topology (Δ lnL = 9.3).
FIGURE 1. Maximum likelihood topology for Gerrhosauridae. Supported nodes are indicated by circles (black: ≥70%
likelihood bootstrap and ≥95% Bayesian posterior probabilities; grey: ≥65% likelihood bootstrap and ≥95% Bayesian
posterior probabilities). Genera and major clades are indicated, and corresponding letters A-F match the text.
0.06 substitutions/site
ANG 0120 G. bulsi
HB 007 C. subtessellatus
PEM R14912 T. fitzsimonsi
HB 085 T. seps
MBUR 00201 G. flavigularis
MBUR 02985 G. nigrolineatus
MBUR 00901 G. v. validus
HB 057 G. nigrolineatus (‘intermedius’)
AY140996 T. petersi
HB 084 T. seps
MBUR 00378 G. flavigularis
PEM R17868 T. seps
QP 0037 C. cordylus
MBUR 01079 G. major
NMZB 16268 G. v. validus
HB 061 G. nigrolineatus (‘intermedius’)
UMMZ RAN5 Z. laticaudatus
MBUR 02981 G. nigrolineatus
KTH 09-290A G. v. maltzahni
G. major (‘bottegoi’)
PEM R17464 S. giganteus
MBUR 02993 G. nigrolineatus
ELS 020 G. flavigularis
JM 03176 G. flavigularis
PEM R5935 G. v. validus
HB 012 T. tetradactylus
HB 056 G. flavigularis
AJC 461 G. nigrolineatus (‘intermedius’)
CAS 206935 G. v. maltzahni
TM 80959 G. nigrolineatus (‘intermedius’)
MBUR 02975 G. nigrolineatus
PEM R14913 T. fitzsimonsi
AMB 6090 G. v. validus
UMMZ 20715 T. petersi
USECH 3505 T. seps
PEM R18886 G. bulsi
HB 011 T. tetradactylus
MBUR 00290 G. v. validus
MBUR 00841 G. flavigularis
KTH 09-289A G. v. maltzahni
USEC H350 G. typicus
MBUR 00469 G. flavigularis
AMB 8339 G. flavigularis
AJC 462 G. nigrolineatus (‘intermedius’)
NMZB 16207 G. auritus
CAS 206978 G. skoogi
ELS 004 G. v. validus
HB 054 G. major
MBUR 00892 G. v. validus
ANG 0116 G. bulsi
MBUR 2809 G. v. validus
AMB 6928 C. subtessellatus
MBUR 01010 G. v. validus
PEM R14985 G. flavigularis
Z. trilineatus
MBUR 00456 G. flavigularis
PEM R15454 G. flavigularis
MBUR 00328 G. flavigularis
HB 180 C. subtessellatus
RSP 205 T. tetradactylus
HB 052 G. major
MBUR 02990 G. nigrolineatus
MBUR 00249 G. flavigularis
KTH 05-06 G. typicus
MBUR 00947 G. flavigularis
HB 055 G. major
HB 083 T. seps
AMB 4649 C. subtessellatus
RSP 352 G. v. validus
NMZB 16199 G. major
MB 21407 G. flavigularis
Eastern South Africa
Northern South Africa
East Africa
Southern Africa
West-Central Africa
East Africa
A
B
C
D
E
F
Matobosaurus
Bates & Tolley
gen. nov.
Zonosaurinae
Cordylosaurus
Tetradactylus
Broadleysaurus
Bates & Tolley
gen. nov.
Gerrhosaurus
Zootaxa 3750 (5) © 2013 Magnolia Press · 473
MOLECULAR PHYLOGENY OF GERRHOSAURUS
TAB L E 2 . Uncorrected p-distances among and within Gerrhosaurus, Cordylosaurus and Tetradactylus species for each
marker: a) ND2 in the top matrix and b) 16S in the bottom matrix. Within species p-distances are on the diagonal of each
matrix. Gerrhosaurus nigrolineatus and G. flavigularis are divided into geographic groups. na = within species p-
distances not available due to only one individual in the dataset. No ND2 data was available for Tetradactylus
tetradactylus.
Discussion
Taxonomy of the genus Gerrhosaurus
The phylogeny obtained (Fig. 1) is in broad agreement with other molecular phylogenies for gerrhosaurids (Lamb
et al. 2003; Lamb & Bauer 2013), and supports the treatment of Angolosaurus as a junior synonym of
Gerrhosaurus. As was the case in the latter two studies, our phylogeny also recovered G. skoogi (Fig. 2) and G.
typicus (Fig. 3) as sister taxa. Gerrhosaurus skoogi (type locality: Port Alexander [= Tombua], Angola) occurs in
the Namib Desert of south-western Angola and adjacent north-western Namibia; our sample was from the
Ondonduiengo River in Namibia. Gerrhosaurus typicus (type locality: “dry sandy flats of Little Namaqualand”,
Northern Cape, South Africa) occurs in Namibia and the Cape provinces of South Africa; our sample was from the
Western Cape (Smith 1837; Andersson 1916; FitzSimons 1953; Visser 1984a,b; Branch 1998; Bates et al. in
press.).
a) ND2 1 2345678910111213141516
Sthn A fr 1 G. nigrolineatus 0.020
East Africa 2 G. nigrolineatus 0 .005 na
W Centr Afr 3 G. nigrolineatus 0.130 0.132 0.019
4G. auritus 0.100 0.109 0.097 na
5G. bulsi 0.123 0.143 0.117 0.120 0.031
Northern SA 6 G. flavigularis 0.175 0.162 0.171 0.154 0.154 0.046
East Africa 7 G. flavigul aris 0.178 0.168 0.164 0.161 0.163 0.044 0.051
Eastern SA 8 G. flavigul aris 0.186 0.189 0.172 0.169 0.164 0.074 0.066 0.002
9G. v. validus 0.230 0.213 0.236 0.224 0.235 0.221 0.238 0.263 0.030
10 G. v. maltzahni 0.233 0.234 0.240 0.248 0.241 0.235 0.244 0.272 0.085 0.005
11 G. typicus 0.206 0.214 0.206 0.214 0.209 0.188 0.187 0.211 0.197 0.202 0.006
12 G. skoogi 0.214 0.223 0.204 0.217 0.219 0.198 0.206 0.225 0.207 0.234 0.167 na
13 G. major 0.205 0.197 0.229 0.194 0.221 0.204 0.203 0.231 0.217 0.228 0.200 0.196 0.036
14 C. subtessellatus 0.225 0.218 0.224 0.210 0.229 0.202 0.205 0.222 0.202 0.224 0.188 0.187 0.177 0.095
15 T. fitzsimonsi 0.226 0.257 0.250 0.260 0.262 0.212 0.207 0.227 0.216 0.245 0.252 0.237 0.227 0.206 na
16 T. seps 0.221 0.210 0.236 0.209 0.241 0.202 0.200 0.225 0.209 0.230 0.194 0.193 0.190 0.189 0.186 0.043
b) 16S 1 2 3 4 5 6 7 8 9 1011 121314151617
Sthn A fr 1 G. nigrolineatus 0.000
East Africa 2 G. nigrolineatus 0.013 na
W Centr Afr 3 G. nigrolineatus 0.072 0.076 0.012
4G. auritus 0.032 0.032 0.064 n a
5G. bulsi 0.044 0.051 0.059 0.057 0.020
Northern SA 6 G. flavigularis 0.040 0 .051 0.055 0.051 0.033 0.014
East Africa 7 G. flavigularis 0.045 0.056 0.053 0.056 0.043 0.008 0.016
Eastern SA 8 G. flavigularis 0.047 0.059 0.072 0.068 0.056 0.020 0.016 0.000
9G. v. validus 0.096 0.106 0.106 0.102 0.078 0.067 0.075 0.087 0.009
10 G. v. maltzahni 0.109 0.116 0.114 0.104 0.103 0.085 0.094 0.107 0.041 0.002
11 G. typicus 0.060 0.070 0.072 0.072 0.057 0.044 0.053 0.064 0 .065 0.087 0 .003
12 G. skoogi 0.084 0.086 0.083 0.078 0.080 0.061 0.070 0.082 0.088 0.100 0.062 na
13 G. major 0.119 0.124 0.126 0.130 0.108 0.090 0.090 0.100 0.090 0.110 0.084 0.129 0.025
14 C. subtessellatus 0.105 0.113 0.085 0.112 0.098 0.094 0.098 0.100 0.122 0.135 0.101 0.103 0.131 0 .024
15 T. fitzsimonsi 0.096 0.104 0.085 0.093 0.092 0.076 0.075 0.089 0.094 0.114 0.067 0.090 0.105 0.103 0.004
16 T. seps 0.092 0.099 0.088 0.102 0.091 0.075 0.083 0.093 0 .097 0.106 0 .079 0.088 0.086 0.086 0.051 0.017
17 T. tetradactylus 0.089 0.101 0.096 0.101 0.088 0.068 0.070 0.068 0.092 0.111 0.078 0.093 0.089 0.104 0.057 0.052 0.015
BATES ET AL.
474 · Zootaxa 3750 (5) © 2013 Magnolia Press
Our improved taxon sampling, moreover, has revealed several notable differences that have significant
implications for the taxonomy of the Gerrhosauridae and the genus Gerrhosaurus in particular. Firstly, we confirm
that the genus Gerrhosaurus is paraphyletic, and find that both G. major and G. validus represent deeply divergent,
well-supported lineages that are outside a clade comprising the remaining Gerrhosaurus + Tetradactylus +
Cordylosaurus. Gerrhosaurus major differs from all other gerrhosaurine taxa by p-distances of at least 19.4%
(ND2) and 8.4% (16S), and each of the two subspecies of G. validus differ from the other taxa by a minimum of
21.3% and 6.7% respectively (Table 2). Although similar results were obtained by Lamb et al. (2003) and Lamb &
Bauer (2013), their phylogenies lacked support at the relevant nodes and in order to maintain monophyletic
lineages, they did not undertake any taxonomic changes, merely treating G. major as incertae sedis within the
Gerrhosaurinae.
FIGURE 2. Young male Gerrhosaurus skoogi from about 20 km north of Terrace Bay, Skeleton Coast, Namibia. (Photo:
J. Harvey)
FIGURE 3. Gerrhosaurus typus from Worcester Valley, Western Cape Province, South Africa. (Photo: A. de Villiers)
Given the phylogenies obtained here and previously (Lamb et al. 2002; Lamb & Bauer 2013), Gerrhosaurus as
currently construed can only be maintained by subsuming Cordylosaurus and the mainly serpentine species
Zootaxa 3750 (5) © 2013 Magnolia Press · 475
MOLECULAR PHYLOGENY OF GERRHOSAURUS
currently included in Tetradactylus, into a large morphologically diverse genus (Gerrhosaurus sensu lato) that
includes the whole African radiation of gerrhosaurids. We are of the opinion that this obscures diversity and
evolutionary history within the family.
In order to maintain monophyletic lineages, and to continue recognition of Gerrhosaurus (sensu stricto) and
particularly the serpentine genus Tetradactylus, the two divergent clades containing, respectively, G. validus and G.
major, must be removed from Gerrhosaurus. Both species exhibit extremely deep divergences, as indicated by
topology tests, the presence of long branches (Fig. 1) and large p-distances (Table 2), and furthermore, they have
distinctive and easily distinguishable morphologies (see Loveridge 1942; FitzSimons 1943). As no suitable
replacement generic names are available within the synonymy of Gerrhosaurus, we take this opportunity to place
them in new genera (see below).
Systematics
Sauria: Gerrhosauridae Boulenger, 1884: Gerrhosaurinae Boulenger, 1884
Broadleysaurus Bates & Tolley gen. nov.
Type species: Gerrhosaurus major Duméril, 1851 here designated
Content: Broadleysaurus major (Duméril, 1851) comb. nov.
Etymology: Named in honour of Dr Donald George Broadley, Curator Emeritus at the Natural History
Museum, Bulawayo (Zimbabwe) for his numerous contributions to African herpetology (e.g. see Branch 1997). He
was recently ranked as the third most productive living author of reptile names (Uetz 2010). It is also fitting to
honour Dr Broadley with this name as his review (1987) of ‘Gerrhosaurus major’ is the most recent and
authoritative. The name Broadleysaurus is masculine, derived from Broadley + the Latinised word saurus meaning
lizard.
Diagnosis: The monophyly of Broadleysaurus is established on the basis of a suite of nuclear and
mitochondrial genetic characters (see above). This genus of terrestrial gerrhosaurids is differentiated from
members of the genus Gerrhosaurus and the new genus described below for ‘G. validus’ by the following
morphological characters: Body stout, cyclotetragonal to feebly depressed dorso-ventrally, and well armoured;
readily distinguished from the above-named genera by its much lower (31–38 versus 49–67) numbers of transverse
dorsal scale rows; also distinguished by having 9–10 longitudinal rows of ventrals (8 in Gerrhosaurus, but 10 in G.
typicus; 12–20 in the new genus described below for ‘G. validus’) (Loveridge 1942; FitzSimons 1943, 1953; De
Witte 1953; Laurent 1954, 1964; Broadley 1966; De Waal 1978; Jacobsen 1989).
Description: Head moderate, its length included in snout-vent length (SVL) 3.9-4.1 times (subadults) or 4.7–
5.2 times (adults); head shields rugose; rostral in contact with, or separated from, the frontonasal; frontonasal entire
or divided (longitudinally); prefrontals usually in broad, but sometimes narrow, contact (rarely separated);
supraoculars 4 (rarely 3); supraciliaries 5 (rarely 3, 4 or 6); tympanic shield narrow, band-like; body
cyclotetragonal or slightly depressed; dorsal scales strongly keeled, striated or rugose, in 14–21 longitudinal and
31–38 transverse rows (usually counted from row posterior to nuchals to row above vent); lateral scales keeled and
striated; ventral plates in 10 (rarely 9) longitudinal and 28–35 transverse rows (counted “from pectoral to anal
shields” according to Loveridge 1942; i.e. from axilla to row before enlarged ventral plate); femoral pores 8–17 per
thigh; fourth toe with 11–17 subdigital lamellae; largest known specimens: male 555 mm (240 mm SVL + 315 mm
tail length), female 518 mm (206 + 312), but another female had a SVL of 245 mm; tail 1.1 to 1.5 times SVL
(combination of features listed by: Loveridge 1942 for the subspecies Gerrhosaurus major major, G. m. bottegoi,
G. m. zechi Tornier and G. m. grandis Boulenger; Broadley 1966 for G. m. major in south-eastern Africa;
FitzSimons 1943 for G. m. grandis in southern Africa; and Jacobsen 1989 for G. m. major in north-eastern South
Africa).
Distribution: Found from Swaziland (Boycott 1992) and the north-eastern parts of South Africa (Bates et al.
in press.) northwards through Mozambique, Zimbabwe, Zambia, Malawi, Tanzania, Uganda, Kenya, southern
Sudan, Somalia and Ethiopia, with additional scattered populations in Central African Republic, Cameroun,
Nigeria, Benin, Togo, Ghana and Senegal (Loveridge 1942; Broadley 1987).
Note: We suggest the name Broadley’s Rough-scaled Plated Lizard for the sole member of this genus.
BATES ET AL.
476 · Zootaxa 3750 (5) © 2013 Magnolia Press
Matobosaurus Bates & Tolley gen. nov.
Type species: Gerrhosaurus validus Smith, 1849 here designated
Content: Matobosaurus validus (Smith, 1849) comb. nov.; Matobosaurus maltzahni (De Grys, 1938) comb.
nov.
Etymology: A masculine name derived from the Ndebele word matobo meaning ‘bald heads’, i.e. smooth
‘whaleback dwalas’ formed when granite is forced to the surface, and the Latinised word saurus meaning lizard.
The word matobo was the name given by Mzilikazi, founder of the Ndebele nation, to the Matobo (Matopo) Hills
area in southern Zimbabwe, characterized by granite hills and wooded valleys, representing prime habitat for
lizards in this genus (see Mertens 1955; Broadley 1966; Pienaar et al. 1983).
Diagnosis: The monophyly of Matobosaurus is established on the basis of a suite of nuclear and mitochondrial
genetic characters (see above). These large, well-armoured lizards have the head and body depressed.
Differentiated from the genera Broadleysaurus and Gerrhosaurus by higher numbers of ventral scale rows
longitudinally (12–20 versus 8–10) and larger size (maximum total length: sex unknown 690 mm [285 mm SVL +
405 mm tail length], male 681 mm, female 587 mm; versus sex unknown 613 mm, male 555 mm, female 518
mm)―although the tail is not particularly long, maximum SVL is distinctly greater than in the other taxa (285 mm
compared to 245 mm in Broadleysaurus and 213 mm in Gerrrhosaurus); also distinguished from Broadleysaurus
by its higher numbers of dorsal scale rows transversely (49–58 versus 31–38) and longitudinally (25–34 versus 14–
21), and usually higher numbers of femoral pores on each thigh (14–25 versus 8–17) (Loveridge 1942; FitzSimons
1943, 1953; De Witte 1953; Laurent 1954, 1964; Broadley 1966; De Waal 1978; Jacobsen 1989). These lizards are
largely rupicolous and often found in (sometimes large) colonies, compared to Broadleysaurus and Gerrrhosaurus
which are mostly terrestrial and found singly or in small groups (Loveridge 1942; FitzSimons 1943; Mertens 1955;
Broadley 1966; Visser 1984a; Jacobsen 1989).
Description: Head moderate, its length included in SVL 4.0–5.0 times; head shields smooth (juveniles),
striated (subadults) or rugose (adults); rostral separated from, but occasionally in contact with, the frontonasal;
prefrontals in broad contact; supraoculars 4; supraciliaries 5 (rarely 4 or 6); subocular excluded from lip by labial,
or in contact with lip; tympanic shield narrow and band-like (young) to broad and subtriangular (adult); body
cyclotetragonal or depressed; dorsal scales keeled, unicarinate (juveniles), tricarinate (subadults) or multicarinate
(adults), and serrated, in 25–34 longitudinal and 49–58 transverse rows (usually counted from row posterior to
nuchals to row above vent); lateral scales keeled and sometimes striated; ventral plates in 12–20 longitudinal and
34–45 transverse rows (counted “from pectoral to anal shields” according to Loveridge 1942; i.e. from axilla to
row before enlarged ventral plate); femoral pores 14–25 per thigh; fourth toe with 15–24 subdigital lamellae;
largest known specimens: sex unknown 690 mm (285 mm SVL + 405 mm tail length), male 681 mm (275 + 406),
female 587 mm (224 + 362), but another female had a SVL of 258 mm; tail 1.3 (young lizards) to 1.8 times SVL
(Loveridge 1942; FitzSimons 1943; Broadley 1966; Jacobsen 1989).
Distribution: Found in Malawi, Zambia, Mozambique, Zimbabwe, eastern Botswana, the north-eastern parts
of South Africa―mainly in the provinces of Limpopo, eastern Mpumalanga and northern KwaZulu-Natal―and
Swaziland (M. validus) and northern Namibia and southern Angola (M. maltzahni) (Loveridge 1942; FitzSimons
1943; Mertens 1955; Broadley 1966; Lang 1991; Branch 1998; Spawls et al. 2002; Adolphs 2006, 2013; Bates et
al. in press.).
Remarks: Although Ulber (1999) pointed out that the name ‘validus’ (for Gerrhosaurus) was in fact originally
spelled ‘vallidus’ by Smith (1849), the common usage of validus can be retained under ICZN (1999) Article
33.2.3.1 (“when an unjustified emendation is in prevailing usage and is attributed to the original author and date, it
is deemed to be a justified emendation”) to maintain nomenclatural stability.
Note: We suggest the name ‘giant plated lizards’ for all members of this genus, and the names Common Giant
Plated Lizard for M. validus and Western Giant Plated Lizard for M. maltzahni. In sub-Saharan Africa the only
lizards that are larger are the monitors (Varanus).
Gerrhosaurus Wiegmann, 1828
Pleurotuchus Smith, 1837
Zootaxa 3750 (5) © 2013 Magnolia Press · 477
MOLECULAR PHYLOGENY OF GERRHOSAURUS
Angolosaurus FitzSimons, 1953
Type species: Gerrhosaurus flavigularis Wiegmann, 1828
Content: Gerrhosaurus flavigularis Wiegmann, 1828; Gerrhosaurus typicus (Smith, 1837); Gerrhosaurus
nigrolineatus Hallowell, 1857; Gerrhosaurus multilineatus Bocage, 1866a; Gerrhosaurus auritus Boettger, 1887;
Gerrhosaurus intermedius Lönnberg, 1907 comb. nov.; Gerrhosaurus skoogi Andersson, 1916; Gerrhosaurus
bulsi Laurent, 1954.
Diagnosis: The monophyly of Gerrhosaurus is established on the basis of a suite of nuclear and mitochondrial
genetic characters (see above). These moderate-sized lizards are fairly well armoured and the head and body may
be cylindrical, cyclotetragonal or slightly depressed; differentiated from the genera Broadleysaurus and
Matobosaurus by its smaller size (maximum SVL 213 mm compared to 245 mm and 285 mm respectively for the
latter two genera) and less robust appearance; most species of Gerrhosaurus have only eight ventral scale rows
longitudinally (but 10 in G. typicus), whereas Broadleysaurus has 9–10 and Matobosaurus has 12–20; it also differs
from Broadleysaurus by having 49–67 versus 31–38 transverse dorsal scale rows (Loveridge 1942; FitzSimons
1943, 1953; De Witte 1953; Laurent 1954, 1964; Broadley 1966; De Waal 1978; Jacobsen 1989).
FIGURE 4. Broadleysaurus major from Arusha, Tanzania. (Photo: W.R. Branch)
Description: Head large, moderate or small, its length included in SVL 3.3–4.8 times (young lizards) to 4.0–
8.4 times (adults); head shields smooth or weakly striated; rostral in contact with, or separated from, the
frontonasal; prefrontals well separated, slightly separated, in narrow contact, or in broad contact; supraoculars 4;
supraciliaries 4–5 (rarely 3 or 6); tympanic shield narrow and band-like to broad and crescentic; body
cyclotetragonal, slightly depressed in some G. t y p i c u s , or almost cylindrical (G. skoogi); dorsal scales weakly to
strongly keeled, smooth or striated, in 20–28 (32–35 in G. skoogi) longitudinal and 49–67 transverse rows (usually
counted from row posterior to nuchals to row above vent); lateral scales keeled, striated or smooth; ventral plates in
8 or 10 (G. typicus only) longitudinal and 30–42 transverse rows (counted “from pectoral to anal shields” according
to Loveridge 1942; i.e. from axilla to row before enlarged ventral plate); femoral pores 9–27 per thigh; fourth toe
with 14–22 subdigital lamellae; largest known specimens: unknown sex 613 mm (213 mm SVL + 400 mm tail
length), male 485 mm (163 + 322), but another male had a SVL of 175 mm female: 475 (142 + 333), but another
BATES ET AL.
478 · Zootaxa 3750 (5) © 2013 Magnolia Press
female had a SVL of 157 mm; tail 1.0 to 2.5 times SVL (Loveridge 1942; FitzSimons 1943, 1953; De Witte 1953;
Laurent 1954, 1964; Broadley 1966; De Waal 1978; Jacobsen 1989).
Distribution: Widespread in Africa south of the equator, extending northwestwards into Gabon and Cabinda,
and north-eastwards through Uganda and Kenya to southern Sudan and Ethiopia (Loveridge 1942; FitzSimons
1943; De Witte 1953; Mertens 1955; Broadley 1966, 1971; De Waal 1978; Auerbach 1987; Jacobsen 1989; Lang
1991; Branch 1998; Spawls et al. 2002; Adolphs 2006, 2013; Bates et al. in press.).
Note: Lizards in this genus are commonly known as ‘plated lizards’.
Status of ‘Gerrhosaurus major’
The type locality of G. major (Fig. 4) is Zanzibar, an island off the coast of Tanzania, but G. m. major has an
extensive range in the eastern half of Africa, from northern KwaZulu-Natal in South Africa to Ethiopia; G. m.
bottegoi was described from Valley of Ghinda in Eritrea and has a fragmented distribution, extending from north-
east Africa (where it occurs together with the nominate subspecies in Kenya) across the continent to West Africa
(Duméril 1851; Del Prato 1895; Loveridge 1942; FitzSimons 1943; Broadley 1966; Jacobsen 1989; Branch 1998;
Spawls et al. 2002; Adolphs 2006, 2013; Bates et al. in press.). The two subspecies are distinguishable only by
their colour patterns (Broadley 1987). Our analysis included samples from southern and eastern Africa identifiable
as G. m. major and one sample from Atakpame in Togo referable to G. m. bottegoi (Table 1). The Togo sample is
embedded within samples of G. m. major. Based on our molecular data, plus the weak morphological differences
(i.e. colour variation) used for recognition of the two subspecies, we relegate G. bottegoi Del Prato, 1895 to the
synonomy of Broadleysaurus major (Duméril, 1851) comb. nov.
Status of ‘Gerrhosaurus validus’
The two currently recognized subspecies of G. va l i d u s each form separate monophyletic clades. In addition,
sequence divergences between these taxa are much larger than would be expected for subspecies and instead are at
the level of species (i.e. 8.5% ND2, 4.1% 16S). The two taxa are morphologically well differentiated (e.g.
subocular excluded from lip by a labial in validus, in contact with lip in maltzahni; longitudinal rows of dorsals 28–
34 in validus, 25–30 in maltzahni; longitudinal rows of ventrals 14–20 in validus, 12–14 in maltzahni; Loveridge
1942, FitzSimons 1943) and occur allopatrically. Gerrhosaurus v. validus occurs from Limpopo Province in South
Africa northwards to Mozambique, Zimbabwe, Zambia and Malawi, while G. v. maltzhani (type locality: Farm
Roidina, north of Omaruru, Namibia; De Grys 1938) is restricted to northern Namibia and southern Angola
(Loveridge 1942; FitzSimons 1943; Broadley 1966; Visser 1984a; Jacobsen 1989; Branch 1998; Spawls et al.
2002; Adolphs 2006, 2013; Bates et al. in press.). The two taxa appear to be separated by the Kalahari Desert
(Visser 1984a). Our samples of G. v. validus were from Limpopo Province in South Africa, Mozambique and
Zimbabwe; G. v. maltzahni was sampled in both Namibia and Angola (Table 1). The type locality for G. validus of
“towards the sources of the Garrep [Gariep], or Orange River” (Smith 1849, Appendix, p. 9), i.e. in Lesotho, must
be in error―as noted by FitzSimons (1943)―as the species is not known to occur anywhere south of 28o latitude
(Branch 1998; Bates et al. in press.). The combination of molecular, morphological and geographical evidence
suggests that the two taxa represent separate evolutionary lineages, and we therefore revive G. maltzahni De Grys,
1938 as a full species, as Matobosaurus maltzahni (De Grys, 1938) comb. nov. The two species in the genus are
illustrated in Figs 5 & 6.
Status of taxa in the Gerrhosaurus nigrolineatus species complex
The type locality of G. nigrolineatus is “Gaboon country, West Africa” (= Gabon; Hallowell 1857). This
species has now been collected at several localities in Gabon (Pauwels et al. 2006), confirming its occurrence there.
As currently understood it has a large distribution range, from Gabon and the lower Congo eastwards through
southern Democratic Republic of the Congo (D.R.C.) to Uganda and Kenya in the east, then southwards as far as
northern Namibia, northern Botswana and north-eastern South Africa (Loveridge 1942; FitzSimons 1943; De Witte
1953; Broadley 1966, 1971; Auerbach 1987; Jacobsen 1989; Branch 1998; Spawls et al. 2002; Bates et al. in
press.; Uetz 2013). Our samples were from Kouilou region, Republic of the Congo (west-Central Africa) adjacent
to Gabon, and Tanzania, Mozambique and South Africa (East and Southern Africa) (Table 1).
Our analysis showed that G. nigrolineatus as currently conceived is not monophyletic, although topology tests
could not reject a monophyletic G. nigrolineatus as presently defined. However, given the observed topology, the
Zootaxa 3750 (5) © 2013 Magnolia Press · 479
MOLECULAR PHYLOGENY OF GERRHOSAURUS
well-supported west-Central African clade of G. nigrolineatus is more closely related to G. auritus, rather than to G.
nigrolineatus from East and Southern Africa, and the nodes defining these groups are well-supported. Given the
node support, as well as other lines of evidence (see below), we suggest that there is reasonably strong support that
G. nigrolineatus as currently defined is not monophyletic. Although the phylogeny of Lamb et al. (2003) also
recovered a sister relationship between G. nigrolineatus and G. auritus, only a single G. nigrolineatus sample from
Mozambique was included. Because our analysis includes greater geographic coverage than previous studies, we
were able to evaluate the status of G. nigrolineatus. In addition to the lack of monophyly for G. nigrolineatus, the
west-Central African clade differs from the East and Southern African clade by large p-distances (13.0% ND2,
6.9% 16S). One individual (HB057, Arusha, Tanzania; Fig. 1) was found less than 140 km to the south-east of the
approximate type locality of Gerrhosaurus flavigularis intermedia Lönnberg, 1907 (i.e. “steppe near the Natron
lakes, Kibonoto”, northern Tanzania; p. 7). Taxonomic implications are that the East/Southern African clade
represents a separate species, for which the name Gerrhosaurus intermedius Lönnberg, 1907 comb. nov. is
available.
FIGURE 5. Matobosaurus validus (NMB R10893) from Greater Kuduland Safaris, east of Tshipise, Limpopo Province,
South Africa. (Photo: M. Burger)
Loveridge (1942) relegated G. f. intermedia to the synonymy of G. n. nigrolineatus without explanation.
Because of the similarity of taxa associated with the names G. flavigularis and G. nigrolineatus, the applicability of
the name G. intermedius for eastern populations previously referred to G. nigrolineatus requires explanation.
Although not mentioned in the text of Lönnberg’s (1907) description of G. f. intermedia, it is evident from his fig.
1b (left side of head) that there are four supraciliaries as in G. nigrolineatus (usually five in G. flavigularis;
Loveridge 1942, FitzSimons 1943). The proportions and scutellation of the head (fig. 1a) are also very similar to
FitzSimons’ (1943) fig. 157 of G. nigrolineatus. In addition, Lönnberg’s description mentions that the flank scales
of G. f. intermedia are strongly keeled, and minium red in colour with dark bars extending from the back. The
prefrontals are shown to be in good contact, with a long median suture (indicated in Lönnberg’s fig. 1a). All of
these features are rare or absent in G. flavigularis and often associated with G. nigrolineatus, including eastern
populations that we now refer to G. intermedius (Fig. 7).
BATES ET AL.
480 · Zootaxa 3750 (5) © 2013 Magnolia Press
FIGURE 6. Matobosaurus maltzahni (PEM R17984) from 0.5 km south of Tambor, Namibé district, Angola. (Photo:
W.R. Branch)
FIGURE 7. Gerrhosaurus intermedius from 9 km south of Skukuza, Kruger National Park, Mpumalanga Province,
South Africa. (Photo: W.R. Schmidt)
In the Congo and Gabon voucher specimens (G. nigrolineatus) examined (Appendix I) there were four
supraciliaries on either side of the head (e.g. PEM R20067, Fig. 8) in all but one specimen (PEM R20066, Congo)
which had five; flanks had weakly or moderately keeled scales in the two Congo specimens, weakly (5) or
Zootaxa 3750 (5) © 2013 Magnolia Press · 481
MOLECULAR PHYLOGENY OF GERRHOSAURUS
moderately (4) keeled in Gabon specimens; prefrontals in broad (PEM R20067) or moderate (PEM R20066)
contact in Congo specimens, in broad (5) to moderate (4) contact in Gabon specimens. We refer all of the above
specimens to G. nigrolineatus. The vouchered Mozambique sample of G. intermedius (TM 80959) from Moebase
Village had four supraciliaries on either side of the head; flanks with strongly keeled scales; and prefrontals in
broad contact.
Although Loveridge (1942: 511) was tempted to “separate an eastern race” of G. nigrolineatus, the only
character he found useful was the number of longitudinal rows of dorsal scales, which numbered 24–28 in “West
Africa” and 20–26 (but usually 22–24) in “East Africa”. Laurent (1954) later gave a count of 26 for a specimen
from Dundo in north-eastern Angola that he assigned to G. nigrolineatus. For southern Africa these counts were
given as 22–24 (usually 22) by FitzSimons (1943) and 20–24 (mostly 22–23) by Jacobsen (1989). The type
description of G. nigrolineatus (Hallowell 1857) refers to 25 longitudinal rows of dorsals, while the holotype of G.
flavigularis intermedia has 22 such rows (Lönnberg 1907). Laurent (1964) later referred a specimen from
Mayombe (lower Congo) with 25 such rows to G. n. nigrolineatus, and four specimens from Pweto in Katanga,
D.R.C., with 24–26 such rows to G. n. intermedius. The number of dorsal rows varied from 23 to 25 in both the
Congo (N = 2) and Gabon (N = 9) specimens examined. The vouchered southern African sample of G. intermedius
(TM 80959) had 24 longitudinal rows of ventrals. While there may be average differences in these counts between
western and eastern populations, there is also some overlap, and the usefulness of this feature for separating G.
nigrolineatus and G. intermedius requires further investigation.
FIGURE 8. Lateral view of the head of a Gerrhosaurus nigrolineatus (PEM R20067) from Koutou base camp for
Sintoukola Potash Project, Kouilou Province, Republic of the Congo. (Photo: M. Burger)
According to Broadley (2007), G. nigrolineatus from Gabon and the lower Congo region has ragged
dorsolateral stripes and smooth plantar scales, features which he felt may distinguish it from populations of this
species elsewhere in Africa. The plantar scales of eastern populations of G. nigrolineatus (= G. intermedius) are
reportedly keeled (smooth and tubercular in G. flavigularis) (FitzSimons 1943; Broadley 1966). In the Congo
specimens examined, the back and flanks were olive to light brown with distinct cream, black-bordered,
dorsolateral stripes, with a similarly coloured vertebral stripe that was continuous in one specimen (PEM R20066)
and broken in the other (PEM R20067). Gabon specimens examined were light brown with scattered black and
BATES ET AL.
482 · Zootaxa 3750 (5) © 2013 Magnolia Press
white lateral scales, and similar stripes, but the vertebral stripe was continuous in one specimen, broken in three
and absent in five. As shown in Fig. 9, MBUR 02993―a specimen sampled for the current analysis―also has
typical dorsolateral stripes as described above, with a broken vertebral stripe. The original description of G.
nigrolineatus refers to a yellow stripe on either side of the back, bordered internally (towards the centre of the
back) by a black band; and also mentions that the centre of the back contains black spots in the form of longitudinal
lines (Hallowell 1857). Colour photographs of the two syntypes of G. nigrolineatus indicated that both specimens
have faded somewhat, but their colour patterns were not dissimilar to the Congo and Gabon material described
above. ANSP 3729 had a pair of pale (cream) dorsolateral stripes with poorly defined black borders as well as a
similar vertebral stripe anteriorly (not visible beyond the nape; Fig. 10), while ANSP 8825 (juvenile) was similar
but lacked a discernible vertebral stripe (Fig. 11).
FIGURE 9. Gerrhosaurus nigrolineatus (MBUR 02993) from Tchiboula region, Kouilou Depratment, Republic of the
Congo. (Photo: M. Burger)
Donald G. Broadley (in litt. 21 March 2013) noted that a specimen of G. nigrolineatus from Ponte Denis in
Gabon in the collection of the Natural History Museum, Zimbabwe (Bulawayo) had smooth plantar scales,
differing somewhat from the weakly keeled plantar scales of PEM R20067 (a detailed photographic image was
used for comparison) from Republic of the Congo (Appendix I). In the Congo specimens examined, plantar scales
were almost smooth or weakly keeled, while in the Gabon sample they were weakly (7) or very weakly (2) keeled.
Based on photographs of one foot of each of the syntypes of G. nigrolineatus, the scales on the soles were weakly
keeled. The plantar scales of the sampled specimen (TM 80959) of G. intermedius were strongly keeled, while
those of 10 additional specimens from Mozambique were moderately keeled; two out of three specimens from
Limpopo Province in South Africa had moderately keeled palmar scales, while one had distinctly keeled scales
(Appendix I).
Although there was some variation in the extent and appearance of dorsal stripes and the keeling of plantar
scales, the Congo and Gabon samples (including material referred to by Broadley) are all considered conspecific
and referable to G. nigrolineatus. Nevertheless, the smooth to feebly keeled plantar scales in G. nigrolineatus from
Gabon and Congo is in contrast to the moderately to strongly keeled scales in populations referable to G.
intermedius (e.g. FitzSimons 1943), including those from Mozambique (e.g. TM 80959 and the other specimens
listed in Appendix I) as discussed above.
The minium red to vermillion flanks (with pale spots or bars) of adult eastern G. nigrolineatus (= G.
intermedius) differ from the light and dark barred or mostly brown flanks of G. flavigularis (see descriptions and
images in Jacobsen 1989; Branch 1998; Spawls et al. 2002; Alexander & Marais 2007). It should be noted
however, that according to Broadley (1966), G. flavigularis from Mozambique and adjacent parts of Zimbabwe
have vermillion flanks like G. nigrolineatus (= G. intermedius), although only in areas of allopatry. The same colour
pattern has been recorded in G. flavigularis from eastern Limpopo Department and eastern North West Province,
South Africa, where the underside of the head is blue-grey in males (Jacobsen 1989). The possibility that such
populations represent unique evolutionary lineages was not investigated in the present study, although some
genetic structuring is evident within G. flavigularis (Fig. 1).
Zootaxa 3750 (5) © 2013 Magnolia Press · 483
MOLECULAR PHYLOGENY OF GERRHOSAURUS
According to Loveridge (1942), the scales on the flanks of G. nigrolineatus (= G. intermedius) are striated,
keeled, or more-or-less smooth, whereas those of adult G. f. flavigularis are smooth. For southern African material,
FitzSimons (1943) noted that the laterals of G. nigrolineatus (= G. intermedius) are keeled and sometimes feebly
striated, while those of G. flavigularis are smooth or feebly keeled and striated. However, Loveridge (1942: 515)
also noted that in his “ill-defined race” G. flavigularis fitzsimonsi (a synonym of G. flavigularis) the laterals were
striated and keeled, although occasionally almost smooth, whereas the prefrontals were in broad contact. The latter
two features are consistent with G. nigrolineatus. However, Loveridge (1942: 515) noted that his new subspecies
had a short head (head length into SVL 4.75 times in young to 6 times in adults) as in G. f. flavigularis, and “should
not be confused with G. f. intermedia…which, from his [Lönnberg 1907] figure, is a synonym of the long-headed
G. n. nigrolineatus”. Head length into SVL was 4.7–5.0 times for the two Congo specimens examined, and 4.0–5.0
times (4.8–5.0 for three adults with SVL >100 mm, 4.0–4.6 for seven juveniles with SVL <80 mm) for the nine
Gabon samples. The vouchered Mozambique sample of G. intermedius (TM 80959) was similar with head length
into SVL 4.4 times. Therefore, we conclude that G. f. intermedia Lönnberg, 1907 is conspecific with eastern
populations currently referred to G. nigrolineatus Hallowell, 1857 and which we now refer to G. intermedius.
FIGURE 10. ANSP 3729, adult syntype of Gerrhosaurus nigrolineatus from Gabon. (Photo: N. Gilmore)
BATES ET AL.
484 · Zootaxa 3750 (5) © 2013 Magnolia Press
FIGURE 11. ANSP 8825, juvenile syntype of Gerrhosaurus nigrolineatus from Gabon. (Photo: N. Gilmore)
Zootaxa 3750 (5) © 2013 Magnolia Press · 485
MOLECULAR PHYLOGENY OF GERRHOSAURUS
In light of the phylogenetic and morphological differences mentioned above, we suggest that populations in
Gabon and lower Congo (including Kouilou region) are all referable to G. nigrolineatus, and that all East and
Southern African populations (Kenya, Uganda, Rwanda, Tanzania, Malawi, Mozambique, Zimbabwe and South
Africa) formerly identified as G. nigrolineatus be referred to G. intermedius. Accurate determination of
geographical boundaries for these two species, especially in Central Africa (Angola, D.R.C., Zambia, northern
Botswana, northern Namibia), will require additional sampling on a finer scale than presently available, as well as
additional morphological examination of specimens from throughout their extensive ranges. The assignment of
Angolan specimens referred to G. nigrolineatus (e.g. Hellmich 1957; Manaças 1963; Parker 1936; Schmidt 1933;
Laurent 1964), and their relationship to G. multilineatus, remains problematic.
FIGURE 12. Gerrhosaurus bulsi (PEM R18941) from Kalumbila Village, Northwest Province, Zambia. (Photo: W.R.
Branch)
FIGURE 13. Gerrhosaurus auritus from near Kome Pan, Botswana. (Photo: W.D. Haacke)
BATES ET AL.
486 · Zootaxa 3750 (5) © 2013 Magnolia Press
Gerrhosaurus bulsi, sister taxon to all other taxa in the G. nigrolineatus complex, is well supported as a distinct
lineage (Fig. 1), and is easily identifiable from others in the complex by its distinct, largely uniform, brown or grey
dorsal colour pattern in adults (Fig. 12). The type locality of G. bu l s i is Dundo, north-east Angola; the species also
occurs in Zambia and the D.R.C. (Laurent 1954, 1964; Broadley 1966; Haagner et al. 2000; Broadley & Cotterill
2004; Adolphs 2006, 2013). Our samples were from Kalumbila Village in North West Province, Zambia; and near
Lake Carumbo, Angola, i.e. about 100 km WSW of the type locality (Table 1).
Gerrhosaurus auritus appears to be closely related to G. nigrolineatus, but morphologically it is
distinguishable by its broad and crescentic (versus narrow) tympanic shield, smooth (versus keeled) lateral scales,
and lack (versus presence) of distinct dorsolateral stripes in adults (Loveridge 1942; FitzSimons 1943; Broadley
2007). Its back is usually pale brown, often with 3–4 narrow, pale, black-bordered dorsolateral stripes (Broadley
1966; Branch 1998; Fig. 13). The type locality of G. auritus is Ondonga, Ovamboland, northern Namibia, but the
species also occurs in southern Angola, south-western Zambia (where our single sample is from―Table 1),
Botswana, western Zimbabwe and northern Limpopo Province in South Africa (Loveridge 1942; FitzSimons 1943;
Broadley 1966; Visser 1984a; Broadley & Rasmussen 1995; Branch 1998; Broadley & Cotterill 2004; Adolphs
2006, 2013; Bates et al. in press.). The four species G. nigrolineatus, G. intermedius, G. auritus and G. bulsi, and
possibly the morphologically and geographically allied form G. multilineatus (if valid, see below), constitute the
‘G. nigrolineatus species complex’ with a widespread distribution in Africa.
FIGURE 14. Lateral view of the head and anterior part of the body of topotypical Gerrhosaurus multilineatus (BM
1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)
FIGURE 15. Dorsal view of the head and anterior part of the body of topotypical Gerrhosaurus multilineatus (BM
1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)
Zootaxa 3750 (5) © 2013 Magnolia Press · 487
MOLECULAR PHYLOGENY OF GERRHOSAURUS
Status of Gerrhosaurus multilineatus
The taxonomic status of G. multilineatus has been confused in the literature and remains uncertain. According
to Haagner et al. (2000), “Broadley (1999) notes that the taxon G. multilineatus Bocage is based on a hybrid
specimen. The name is therefore unavailable.” However, this was in fact a reference to an unpublished manuscript
(D.G. Broadley in litt. 8 February 2012). According to Article 17.2 of the Code (ICZN 1999), even if the specimen
was a hybrid, the name would in fact still be available.
In his description of G. multilineatus, based mainly on colour pattern, Bocage (1866a) noted that this form was
similar to G. nigrolineatus, of which it may be merely a well characterised variety. Loveridge (1942) and
FitzSimons (1943) subsequently relegated G. multilineatus to the synonymy of G. nigrolineatus. Although the type
series of G. multilineatus (Duque de Bragança district [region], interior of Angola) was destroyed in the 1978 fire at
Museu Bocage in Lisbon (Almaca & Neves 1987; Madruga 2012), we examined colour photographs of two
‘virtual’ topotypes (‘Duque de Bragança’) in the collection of the Natural History Museum (London). In terms of
morphology and colour pattern (e.g. Figs 14 & 15) these specimens agree well with Bocage’s description. Although
somewhat faded, cream coloured longitudinal stripes, with black borders, are present on the back, at least
anteriorly. In BM 1904.5.2.32 there are dorsolateral stripes as well as a vertebral stripe (Fig. 15), as described by
Bocage (1866a), whereas BM 1904.5.2.33 appears to have only dorsolateral stripes. The two specimens (about 170
mm and 150 mm SVL respectively) appear to be adults.
FIGURE 16. Juvenile Gerrhosaurus bulsi from Lake Carumbo base camp, Lunda Norte Province, Angola (PEM
R19475). (Photo: W.R. Branch)
Laurent (1964) presented data for a large series of Gerrhosaurus from Angola which he referred to G. bulsi,
contrasting these with a specimen from ‘Mayombe’ (may refer to the region from western Gabon southwards to
western D.R.C., or to the Mayombe massif in Republic of Congo) which he referred to G. nigrolineatus
nigrolineatus (because of its “blackish colouration”, p. 54), and four specimens from Pweto at the northern end of
Lake Mweru in Katanga Province, D.R.C. which he referred to G. nigrolineatus intermedius. Laurent (1964: 54)
noted that if the “type” of G. multilineatus was a young G. bulsi, the former name would have priority. According to
Laurent (1964), young G. bulsi have a (striped) dorsal colour pattern similar to that of G. nigrolineatus (striped
BATES ET AL.
488 · Zootaxa 3750 (5) © 2013 Magnolia Press
throughout life), but this pattern gradually fades and adult G. bulsi (13 cm SVL and larger) display a uniformly
coloured and unpatterned dorsum (Fig. 12). However, juvenile G. bulsi are not always striped: PEM R19475 from
Lake Carumbo base camp in Angola (Fig. 16), used for the molecular analysis, is brown with dark bands on the
flanks and scatterered dark scales on the back, but it lacks any distinct dorsolateral (and vertebral) stripes, whether
pale, black, or pale with black borders. Bocage’s (1866a) description of G. multilineatus was based on “three
specimens of identical colour” (see Bocage 1866b: 44, a paper preceding the description). However, although
Bocage (1866a) described all three syntypes as having olive backs with narrow yellow, black-bordered dorsolateral
and vertebral stripes, he noted that the largest (apparently adult) specimen (123 mm SVL + 250 mm tail length)
also had three similar but narrower stripes in each of the interspaces between dorsolateral and vertebral stripes,
while in the other (smaller) specimens these intermediate stripes were replaced by black markings. From the
available information it therefore seems that, at the very least, the largest specimen examined by Bocage (1866a) is
not conspecific with G. bulsi (adults are unstriped). Whether or not Bocage’s specimens are referable to G.
nigrolineatus, or a separate species, is unclear. However, the two topotypes of G. multilineatus do not have the very
large heads typical of adult G. nigrolineatus, G. intermedius and G. auritus (i.e. head length into SVL five times or
less; see Loveridge 1942, FitzSimons 1943). Based on scaled photographs of BM 1904.5.2.33 and BM
1904.5.2.32, head length is contained in SVL about 5.6 and 5.7 times, respectively.
Resolution of the taxonomic status of G. multilineatus must await the collection of material from the type
locality for molecular analysis, and a detailed morphological evaluation of the complex.
FIGURE 17. Gerrhosaurus flavigularis from Suikerbosrand Nature Reserve, Gauteng Province, South Africa. (Photo:
W.R. Schmidt)
Status of Gerrhosaurus flavigularis
While there is some sub-structuring within G. flavigularis (Fig. 17), with populations from Eastern South
Africa, Northern South Africa and East Africa (Table 1) all identifiable as subclades in the phylogeny (Fig. 1), we
consider this assemblage a single species pending a more detailed phylogeographical and morphological analysis.
The type locality for G. flavigularis of “South [or southern] Africa” (see Bauer et al. 1994) was restricted to an area
in the central Eastern Cape Province, South Africa (Bauer 2000), but the species occurs extensively from the
Zootaxa 3750 (5) © 2013 Magnolia Press · 489
MOLECULAR PHYLOGENY OF GERRHOSAURUS
Western Cape (South Africa) northwards through southern and eastern Africa to Ethiopia (Loveridge 1942).
Although not sampled for this study, should the apparently disjunct population in the Western and Eastern Cape
Provinces be found to represent a unique lineage, the name G. flavigularis would be applicable to it. If the other
population in southern and East Africa proves to be a separate species, the name Gerrhosaurus bibroni Smith, 1844
is available.
Acknowledgements
We thank the following individuals: Marius Burger (and the Southern African Reptile Conservation Assessment
project), Werner Conradie, Edward Stanley and John Measey for collecting specimens or tissue samples used in
this study; Don Broadley for drawing our attention to the existence of the two topotypes of Gerrhosaurus
multilineatus in the Natural History Museum (London); Patrick Campbell at the latter museum for providing
photographs of these specimens; Ned Gilmore at the Academy of Natural Sciences (Philadelphia) for providing
photographs of the two syntypes of G. nigrolineatus; Lauretta Mahlangu for recording morphological data for a
specimen of G. intermedius (TM 80959) in the Ditsong National Museum of Natural History (Pretoria); Marius
Burger for photographs published in this paper, as well as additional images of G. nigrolineatus from Kouilou
region, Republic of the Congo; Warren Schmidt, Wulf Haacke, James Harvey and Atherton de Villiers for
photographs used in this paper; and Klaus Adolphs and Aaron Bauer for supplying copies of old literature.
GERDIB (Groupe d’Etude et de Recherche sur la Diversité Biologique) in Brazzaville is thanked for supplying
collecting permits for Republic of the Congo. This work was funded by the National Research Foundation of South
Africa (South African Biosystematics Initiative Grant Number 65778).
References
Adolphs, K. (2006) Bibliotheca Cordyliformium. Squamata Verlag, Sannkt Augustin, 303 pp.
Adolphs, K. (2013) Squamata – Verlag für Herpetologie. Available from: http://www.squamata.de/ (accessed 25 June 2013)
Alexander, G. & Marais, J. (2007) A Guide to the Reptiles of Southern Africa. Struik Publishers, Cape Town, 408 pp.
Almaca, C. & Neves, A.M. (1987) The Museu Bocage and the new series of its Arquivos. Arquivos do Museu Bocage, Nova
Série, I (1), 1–8.
Andersson, L.G. (1916) Notes on the reptiles and batrachians in the Zoological Museum at Gothenburg with an account of some
new species. Göteborgs Kungliga Vetenskap och Vitter Hets-Samnalles Handlinger Sjatte Foljden, Series B, 4, (17, No. 5),
[Meddelanden fran Göteborgs Musei Zoologiska Afdelning (Göteborg), No. 9], 1–41.
Auerbach, R.D. (1987) The amphibians and reptiles of Botswana. Mokwepa Consultants, Gaborone, i–v, 295 pp.
Bates, M.F., Branch, W.R., Bauer, A.M., Burger, M., Marais, J., Alexander, G.J. & de Villiers, M. (Eds.) (in press.) Atlas and
Red List of the Reptiles of South Africa, Lesotho and Swaziland. Strelitzia, 32, xx–yy.
Bauer, A.M. (2000) Comments on the types and type localities of South African reptiles collected by Heinrich Bergius and
Ludwig Krebs. African Journal of Herpetology, 49 (1), 53–60.
http://dx.doi.org/10.1080/21564574.2000.9650016
Bauer, A.M., Lang, M. & Günther, R. (1994) An Annotated Type Catalogue of the Pygopods, Xantusiids, Gerrhosaurs and
Cordylids (Reptilia: Squamata: Pygopodidae, Xantusiidae, Gerrhosauridae and Cordylidae) in the Collection of the
Zoological Museum. Mitteilungen aus dem Zoologischen Museum in Berlin, 70 (1), 85–92.
http://dx.doi.org/10.1002/mmnz.19940700103
Bocage, J.V. Barboza du (1866a) Reptiles nouveaux ou peu connus recueillis dans les possessions portugaises de l'Afrique
occidentale, qui se trouvent au Muséum de Lisbonne. Jornal de Sciencias Mathematicas, Physicas e Naturaes (Lisboa), 1,
57–78.
Bocage, J.V. Barboza du (1866b) Lista dos reptis das possessões portuguezas d’Africa occidental que existem no Museu de
Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes (Lisboa), 1, 37–56.
Boettger, O. (1887) Zweiter Beitrag zur Herpetologie Südwestund Süd-Afrikas. Bericht über die Senckenbergische
naturforschende Gesellschaft in Frankfurt an Main (Frankfurt) (1886/1887), pp. 135–173, 5 tables.
Boycott, R.C. (1992) A herpetofaunal survey of Swaziland. M.Sc. dissertation, University of Natal, Durban, 13+175+43 pp.
Branch, W.R. (1997) A tribute to Don Broadley: Africa’s premier herpetologist. African Journal of Herpetology, 46 (2), 58–67.
http://dx.doi.org/10.1080/21564574.1997.9649979
Branch, W.R. (1998) Field Guide to Snakes and Other Reptiles of Southern Africa. Second Edition. Struik Publishers, Cape
Town, 399 pp.
Broadley, D.G. (1960) Sauria, Cordylidae: Gerrhosaurus nigrolineatus anselli n.subsp. Occasional Papers of the National
Museums of Southern Rhodesia, 24 (B), 433–434.
BATES ET AL.
490 · Zootaxa 3750 (5) © 2013 Magnolia Press
Broadley, D.G. (1966) The herpetology of south-east Africa. Unpublished Ph.D. dissertation, University of Natal,
Pietermaritzburg, 680 pp.
Broadley, D.G. (1971) The reptiles and amphibians of Zambia. The Puku, 6, 1–143.
Broadley, D.G. (1987) A review of geographical variation in Gerrhosaurus major Duméril (Sauria: Cordylidae).
Herpetological Journal, 1, 194–198.
Broadley, D.G. (2007) Book review: Bibliotheca Cordyliformium. Neues Quellenverzeichnis der Gurtelschweife und
Schildechsen (Reptilia, Cordylidae & Gerrhosauridae) by Klaus Adolphs. African Journal of Herpetology, 56 (1), 99–100.
Broadley, D.G. & Cotterill, F.P.D. (2004) The reptiles of southeast Katanga, an overlooked 'hot spot'. African Journal of
Herpetology, 53 (1), 35–61.
http://dx.doi.org/10.1080/21564574.2004.9635497
Broadley, D.G. & Rasmussen, G.S.A. (1995) Geographical distribution: Gerrhosaurus multilineatus auritus. African Herp
News, 23, 47.
Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Singlelocus and multilocus DNA fingerprint. In: Ha, R.
(Ed.), Molecular genetic analysis of populations: a practical approach. IRL Press, Oxford, pp. 225–270.
Conradie, W., Measey, G.J., Branch, W.R. & Tolley, K.A. (2012) Revised phylogeny of African sand lizards (Pedioplanis), with
the description of two new species from south-western Angola. African Journal of Herpetology, 61, 91–112.
http://dx.doi.org/10.1080/21564574.2012.676079
Cunningham, M. & Cherry, M. (2004) Molecular systematics of African 20–chromosome toads (Anura: Bufonidae). Molecular
Phylogenetics and Evolution, 32, 671–685.
http://dx.doi.org/10.1016/j.ympev.2004.03.003
De Grys, P. (1938) Gerrhosaurus Maltzahni spec. nov. Zoologischer Anzeiger (Leipzig), 124 (3/4), 58–60.
De Waal, S.W.P. (1978) The Squamata (Reptilia) of the Orange Free State, South Africa. Memoires van die Nasionale Museum,
Bloemfontein, 11, +i–iii, 1–160.
De Witte, G.F. (1953) Exploration du Parc National de l’Upemba. Vol. 6. Reptiles. Institut des Parcs Nationaux du Congo
Belge, Bruxelles, 322 pp.
Del Prato, A. (1895) Vertebrati Eritrea. Aggiunta al catalogo della collezione Eritrea Bottego. Atti della Societa Italiana di
Science Naturali e del Museo Civico di Storia Naturale in Milano (Milano), 35, 17–26.
Drummond, A.J., Ashton, B., Cheung, M., Heled, J., Kearse, M., Moir R., Stones-Havas, S., Thierer, T. & Wilson, A. (2009)
Geneious v4.6, Software available from: http://www.geneious.com/ (accessed 27 November 2013)
Duméril, A. (1851) Catalogue Méthodique de la Collection des Reptiles (Muséum d’Histoire Naturelle, Paris), Gide et Baudry,
Paris, i–iv, 224 pp.
Duméril, A.-M.-C. & Bibron, G. (1839) Erpétologie général ou Histoire naturelle complète des Reptiles. Vol. 5. Roret éd.,
Paris, [Nine volumes text, one volume plates], i–viii + 1–855, pls 43–58.
FitzSimons, V.F. (1943) The lizards of South Africa. Memoirs of the Transvaal Museum, 1, i–xv + 1–528, +24 pls + map.
FitzSimons, V.F. (1953) A new genus of gerrhosaurid from southern Angola. Annals of the Transvaal Museum, 22 (2), 215–
217.
Frost, D., Janies, D., Mouton, P. le F.N. & Titus, T. (2001) A molecular perspective on the phylogeny of the Girdled Lizards
(Cordylidae, Squamata). American Museum Novitates, 3310, 1–10.
http://dx.doi.org/10.1206/0003-0082(2001)310<0001:ampotp>2.0.co;2
Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third edition. Vences & Glaw
Verlag, Köln, 496 pp.
Goldman, N., Anderson, J.P. & Rodrigo, A.G. (2000) Likelihood-based tests of topologies in phylogenetics. Systematic Biology,
49, 652–670.
Haagner, G.V., Branch, W.R. & Haagner, A.J.F. (2000) Notes on a collection of reptiles from Zambia and adjacent areas of the
Democratic Republic of the Congo. Annals of the Eastern Cape Museums, 1, 1–25.
Hallowell, E. (1857) Notice of a collection of Reptiles from the Gaboon country, West Africa, recently presented to the
Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of
Philadelphia, 9, 48–72.
Hellmich, W. & Schmelcher, D. (1956) Eine neue Rasse von Gerrhosaurus nigrolineatus Hallowell (Gerrhosauridae).
Zoologischer Anzeiger, 156 (7/8), 202–205.
Hellmich, W. (1957) Die Reptilienausbeute der Hamburgischen Angola-Expedition. Mitteilungen aus dem Hamburgischen
Zoologischen Museum und Institut, 55, 39–80.
Hoser, R. (2013) A revised taxonomy for the Lizard Families Gerrhosauridae and Cordylidae. Australasian Journal of
Herpetology, 21, 3–32.
International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth Edition.
The International Trust for Zoological Nomenclature, The Natural History Museum, London, i–XXIX + 306 pp.
IUCN (1996) 1996 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland and Cambridge, UK, 448 pp.
Jacobsen, N.H.G. (1989) The Distribution and Conservation Status of Reptiles and Amphibians in the Transvaal. Final Report
Project TN 6/4/1/30. Chief Directorate of Nature and Environmental Conservation, Pretoria, i–xlii, 1621 pp.
Kaiser, H., Crother, B.I., Kelly, C.M.R., Luiselli, L., O’Shea, M., Ota, H., Passos, P., Schleip, W.D. & Wüster, W. (2013) Best
Practices: In the 21st Century, Taxonomic Decisions in Herpetology are Acceptable Only When Supported by a Body of
Evidence and Published via Peer-Review. Herpetological Review, 44 (1), 8–23.
Zootaxa 3750 (5) © 2013 Magnolia Press · 491
MOLECULAR PHYLOGENY OF GERRHOSAURUS
Lamb, T. & Bauer, A.M. (2013) To be or not to be Angolosaurus: a multilocus perspective on the phylogenetic position of
Africa’s desert plated lizard (Gerrhosauridae). Zoologica Scripta, 42 (4), 381–388.
http://dx.doi.org/10.1111/zsc.12016
Lamb, T., Meeker, A.M., Bauer, A.M. & Branch, W.R. (2003) On the systematic status of the desert plated lizard
(Angolosaurus skoogi): Phylogenetic inference from DNA sequence analysis of the African Gerrhosauridae. Biological
Journal of the Linnean Society, 78, 253–261.
http://dx.doi.org/10.1046/j.1095-8312.2003.00168.x
Lang, M. (1991) Generic relationships within Cordyliformes (Reptilia: Squamata). Bulletin de l'Institut recherches sciences
naturelles de Belgique, 61, 121–188.
Laurent, R.F. (1954) Reptiles et batraciens de la région de Dundo (Angola) (Deuxième note). Companhia de Diamantes de
Angola (Diamang), Serviços Culturais, Publicações Culturais, 23, 37–84.
Laurent, R.F. (1964) Reptiles et Amphibiens de l’Angola (Trosième contribution). Companhia de Diamantes de Angola
(Diamang), Serviços Culturais, Museu do Dundo, 67, 12–165.
Lönnberg, E. (1907) Reptilia and Batrachia. In: Sjostedt, Y. (Ed.), Wissenschaftliche Ergebnisse der Schwedischen
Zoologischen Expedition nach dem Kilimandjaro, dem Meru und dem umgebeden Massaisteppen 1905-1906 (Upsala), 4,
28 pp., pl. 1.
Loveridge, A. (1942) Revision of the African lizards of the family Gerrhosauridae. Bulletin of the Museum of Comparative
Zoology, 89 (11), 485–543.
Madruga, C. (2012) The zoological collection of the Museu de Lisboa and the networks of scientific correspondence and
exchange (1858–1898). In: Roca-Rosell, A. (Ed.), The Circulation of Science and Technology: Proceedings of the 4th
International Conference of the European Society for the History of Science, Barcelona, 18–20 November 2010. SCHCT–
IEC, Barcelona, pp. 928–933. Available from: http://www.academia.edu/783491/Networks_of_scientific_correspondence
_and_exchange_1858–1898_ (accessed 13 February 2013)
Manaças, S. (1963) Saurios de Angola. Memorias da Junta de Investigaçoes do Ultramar, 43 (2a series), 223–240.
Mertens, R. (1955) Die Amphibien und Reptilien Südwesafrikas. Aus den Ergebnissen einer im Jahre 1952 ausgeführten Reise.
Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 490, 1–172.
Nance, H. (2003) The cranial osteology of Angolosaurus skoogi (Squamata: Gerrhosauridae) with comments on the
morphology and phylogeny of Cordyliformes. Unpubl. M.Sc. dissertation, University of Texas, Austin, Texas.
Nance, H.A. (2007) Cranial osteology of the African gerrhosaurid Angolosaurus skoogi (Squamata: Gerrhosauridae). African
Journal of Herpetology, 56, 39–75.
http://dx.doi.org/10.1080/21564574.2007.9635552
Odierna, G., Canapa, A., Andreone, F., Aprea, G., Barucca, M., Capriglione, T. & Olmo, E. (2002) A phylogenetic analysis of
Cordyliformes (Reptilia: Squamata): Comparison of molecular and karyological data. Molecular Phylogenetics and
Evolution, 23, 37–42.
http://dx.doi.org/10.1006/mpev.2001.1077
Palumbi, S.R., Martin, A.P., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool’s guide to PCR.
In: Department of Zoology, Special Publication. University of Hawai'i, Honolulu, pp. 44.
Parker, H.W. (1936) Dr. Karl Jordan’s expedition to south-west Africa and Angola: herpetological collections. Novitates
Zoologicae, 40, 115–146.
Pauwels, O.S.G., Christy, P. & Honorez, A. (2006) Reptiles and national parks in Gabon, western Central Africa. Hamadryad,
30 (1&2), 181–196.
Pienaar, U. de V., Haacke, W.D. & Jacobsen, N.H.G. (1983) The Reptiles of the Kruger National Park. National Parks Board of
South Africa, Pretoria, 263 pp.
Portik, D., Wood, P., Jr., Grismer, J., Stanley, E. & Jackman, T. (2012) Identification of 104 rapidly-evolving nuclear protein-
coding markers for amplification across scaled reptiles using genomic resources. Conservation Genetics Resources, 4, 1–
10.
http://dx.doi.org/10.1007/s12686-011-9460-1
Posada, D. (2008) Jmodeltest: Phylogenetic model averaging. Molecular Biology and Evolution, 25, 1253–1256.
http://dx.doi.org/10.1093/molbev/msn083
Rambaut, A. & Drummond, A.J. (2007) Tracer. Version 1.4. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 27
November 2013)
Raselimanana, A.P., Noonan, B., Karanth, K.P., Gauthier, J. & Yoder, A.D. (2009) Phylogeny and evolution of Malagasy plated
lizards. Molecular Phylogenetics and Evolution, 50, 336–344.
http://dx.doi.org/10.1016/j.ympev.2008.10.004
Recknagel, H., Elmer, K.R., Noonan, B.P., Raselimanana, A.P., Meyer, A. & Vences, M. (2013) Multi-gene phylogeny of
Madagascar’s plated lizards, Zonosaurus and Tracheloptychus (Squamata: Gerrhosauridae). Molecular Phylogenetics and
Evolution, 69, 1215–1221.
http://dx.doi.org/10.1016/j.ympev.2013.06.013
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19,
1572–1574.
http://dx.doi.org/10.1093/bioinformatics/btg180
BATES ET AL.
492 · Zootaxa 3750 (5) © 2013 Magnolia Press
Schmidt, K.P. (1933) The reptiles of the Pulitzer Angola expedition. Annals of the Carnegie Museum, 22 (1), 1–15, 2 pls.
Shimodaira, H. (2002) An approximately unbiased test of phylogenetic tree selection. Systematic Biology, 51, 492–508.
Shimodaira, H. & Hasegawa, M. (1999) Multiple comparisons of log-likelihoods with applications to phylogenetic inference.
Molecular Biology and Evolution, 16, 1114–1116.
http://dx.doi.org/10.1093/oxfordjournals.molbev.a026201
Smith, A. (1837) The Characters of two new Genera of South African Reptiles, with descriptions of species belonging to each.
Magazine of Zoology and Botany, 1 (2), 141–145.
Smith, A. (1838–1849) Illustrations of the Zoology of South Africa; Consisting Chiefly of Figures and Descriptions of the
Objects of Natural History Collected During an Expedition into the Interior of South Africa, in the Years 1834, 1835, and
1836; Fitted Out by “The Cape of Good Hope Association for Exploring Central Africa.” 5. Reptilia. Smith, Elder and
Co., London, [3] + [188] + Appendix 28 pp., 78 pls.
Spawls, S., Howell, K., Drewes, R. & Ashe, J. (2002) A Field Guide to the Reptiles of East Africa. Academic Press, London,
543 pp.
Stamatakis, A. (2006) RAxML–VI–HPC: Maximum Likelihood-based Phylogenetic Analyses with Thousands of Taxa and
Mixed Models. Bioinformatics, 22 (21), 2688–2690.
http://dx.doi.org/10.1093/bioinformatics/btl446
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic
Biology, 57, 758–771.
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 and Evolution, 58, 53–
70.
http://dx.doi.org/10.1016/j.ympev.2010.08.024
Swofford, D.L. (2002) PAUP*.Phylogenetic Analysis Using Parsimony (*and Other Methods), 4.0 edn. Sunderland, MA, USA:
Sinauer Associates.
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M., & Kumar, S. (2011) MEGA5: Molecular Evolutionary Genetics
Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Molecular Biology and
Evolution, 28 (10), 2731–2739.
http://dx.doi.org/10.1093/molbev/msr121
Tilbury, C.R. & Tolley, K.A. (2009) A re-appraisal of the systematics of the African genus Chamaeleo (Reptilia:
Chamaeleonidae). Zootaxa, 2079, 57–68.
Tolley, K.A., Burger, M., Turner, A.A. & Matthee, C.A. (2006) Biogeographic patterns and phylogeography of dwarf
chameleons (Bradypodion) in an African biodiversity hotspot. Molecular Ecology, 15, 781–793.
http://dx.doi.org/10.1111/j.1365-294x.2006.02836.x
Uetz, P. (2010) The original descriptions of reptiles. Zootaxa, 2334, 59–68.
Uetz, P. (2013) (Ed.) The Reptile Database. Updated April 2013. Available from: http://www.reptile-database.org/ (accessed 22
May 2013)
Ulber, T.M. (1999) Sir Andrew Smith’s “Illustrations of the Zoology of South Africa”: An Annotated Table of Contents for the
"Reptilia" Volume. Smithsonian Herpetological Information Service, 120, 1–17.
Visser, J. (1984a) Gepantserde Akkedisse. 31. Net dinamiet sal dié kêrel laat roer. Landbou Weekblad, 346, 66–67, 69, 71. [19
October 1984]
Visser, J. (1984b) Gepantserde Akkedisse. 32. Geslepe klein bloustert ʼn kulkunstenaar. Landbou Weekblad, 347, 72–73, 75, 77.
[26 October 1984]
Wermuth, H. (1968) Liste der rezenten Amphibien und Reptilien. Cordylidae (Subfam. Cordylinae et Gerrhosaurinae). Das
Tierreich, 87, ++, VIII–X, 1–30.
Wiegmann, A.F. (1828) Beyträge zur Amphibienkunde. Isis von Oken (Leipzig), 21, 364–383.
APPENDIX I. List of Gerrhosaurus nigrolineatus and G. intermedius specimens examined for morphological
characters.
Gerrhosaurus nigrolineatus
GABON: Ogooué-Maritime Province―Loango National Park: PEM R5405 & R5437 (02°20'27"S, 09°35'33"E, O.S.G.
Pauwels & W.R. Branch, 8 and 2 Oct 2002 respectively), PEM R5411 (02°19'56"S, 09°35'32"E, O.S.G. Pauwels & W.R.
Branch, 11 Oct 2002), PEM R5435 (no co-ordinates available, O.S.G. Pauwels & W.R. Branch, 15 Oct 2002), PEM R5965
(02°20'27"S, 09°35'50"E, O.S.G. Pauwels & W.R. Branch, 10 Oct 2002); East of Gamba on road to Vera: PEM R5208 & MB
05848 (02°45'31"S, 10°08'13"E, M. Burger, 11 Jul 2001), PEM R5209 (02°44'41"S, 10°08'28"E, M. Burger, 11 Jul 2001);
Rabi, Shell Gabon: PEM R5322 (01°52'58"S, 09°50'26"E, J.A. Yoger & O.S.G. Pauwels, 10 Mar 2002). REPUBLIC OF THE
CONGO: Kouilou Department [region]―5.5 km west of Tchizondi: PEM R20066 (04°22'47.4"S, 11°36'17.2"E, M. Burger &
A.G. Zassi-Boulou, 12 May 2012); Koutou base camp for Sintoukola Potash: PEM R20067 (04°07'38.3"S, 11°41'37.0"E, M.
Burger & A.G. Zassi-Boulou, 16 Jul 2012).
Zootaxa 3750 (5) © 2013 Magnolia Press · 493
MOLECULAR PHYLOGENY OF GERRHOSAURUS
Gerrhosaurus intermedius
MOZAMBIQUE: Zambezia Province―between Namagure and Moebase Villages: PEM R13208 (16°58'26"S,
38°43'48"E, W.R. Branch, 13 May 1997); Moebase Village: PEM R13269, R13287, R13311, R13319 (16°58'58"S,
38°43'43"E, W.R. Branch, 21 May 1997), TM 80959 (16°59’05”S, 38°43’42.2”E, M.J. Whiting & W.R. Branch, 13 May
1997); Moleque River: PEM R13274 (17°02'31.2"S, 38°50'16"E, W.R. Branch, 24 May 1997); Tupuiko Village: PEM R15597,
R15670, R15671 (16°31'07"S, 39°38'38"E, W.R. Branch, 27 Feb 2000 and 18 Apr 2000); Cabo Delagado Province―Afungi
ENI site, Palma: PEM R20153 (10°48'51.1"S, 40°32'22.2"E, W. Conradie, 28 Aug 2012). SOUTH AFRICA: Limpopo
Province―Hermitage, Manyeleti Game Reserve: NMB R5987 (24°36’S, 31°27’E, G.V. Haagner, 8 Dec 1988); Dixie,
Manyeleti Game Reserve: NMB R5993 (24°42’S, 31°31’E, G.V. Haagner, 19 Dec 1988); Farm Glencoe 210, Phalaborwa
district: NMB R7588 (24°21’S, 30°52’E, G.V. Haagner, 9 Apr 1996).
