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An integrative taxonomic revision of the Tarentola geckos (Squamata, Phyllodactylidae) of the Cape Verde Islands

February 2012
Zoological Journal of the Linnean Society 164(2):328–360

February 2012
164(2):328–360

DOI:10.1111/j.1096-3642.2011.00768.x

Authors:

Raquel Vasconcelos

University of Porto

Ana Perera

David James Harris

University of Porto

Show all 5 authorsHide

Recent phylogeographical analyses using mitochondrial DNA (mtDNA) sequences indicate that the Tarentola geckos from the Cape Verde archipelago originated from a propagule that dispersed from the Canary Islands approximately 7.7 Mya and that underwent a fast evolutionary radiation. Molecular analyses carried out to date clearly show some incongruences with the current taxonomy of Tarentola from the Cape Verde Islands, with some species being paraphyletic or polyphyletic, and several independently evolving lineages needing formal taxonomic recognition. The aim of this study was to clarify the systematics of this group to unravel its taxonomy by applying an integrative approach based on information from three independent sources: mtDNA, nuclear genes, and morphology. As a result of this taxonomic revision, two novel species for the islands of S. Nicolau and Fogo are described and eight subspecies are upgraded to species level. Moreover, an identification key for the genus Tarentola from the Cape Verde archipelago is presented. This study reconciles taxonomy and phylogeny in this group, provides a better understanding of diversity patterns, new insights on evolutionary hypotheses, and supports the basic framework for the future management and conservation of this unique reptile radiation. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 328–360.

Map of the Cape Verde Islands showing the geographical location (latitudes and longitudes) and altitudes of the islands and the origins of the new Tarentola samples included in the genetic (circles) and morphological (diamonds) analyses (Geographic Coordinate System, Datum WGS 84). Island and taxa colours match the colours used on the network analyses. No specimens were found on Sal.

…

. Descriptive statistics for all the linear measurements and meristic variables of adult specimens of the different Tarentola taxa included in the multivariate analysis T. bocagei São Nicolau

…

. Continued T. darwini Santiago

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Phylogenetic relationships of endemic Cape Verde Tarentola taxa and their relatives from the Canary Islands modified from Vasconcelos et al. (2010) based on cytochrome b and 12S rRNA genes. The tree was inferred using maximum likelihood (ML) and GTR+I+G model of sequence evolution (log likelihood = −6468.896) and was rooted using Tarentola americana. Bootstrap support values above 60% for the ML analysis are shown below nodes. Posterior probability (PP) values higher than 95% for the Bayesian analysis are represented by an asterisk (*) and are shown above nodes. Names in bold follow the new taxonomic proposal and non-bold ones the taxonomy accepted in previous recent papers (Carranza et al., 2000; Jesus et al., 2002; Vasconcelos et al., 2010). For further details see Vasconcelos et al. (2010). Characters immediately to the right of island names correspond to the 15 evolutionarily significant units (ESUs) of A, B, C, and D clades recognized in the present work and represented in split green bars. Lines of evidence (in grey): 1, mitochondrial DNA (independent cyt b parsimony networks with a connection limit of 95%; see Appendix 3); 2, nuclear DNA (absence of shared haplotypes in MC1R); 3, morphology (detection of any diagnostic morphological character or a set of a unique combination of characters). Integration approaches (in red) from the most conservative to the most inflationist: ITC stands for integration by total congruence (all lines of evidence should be congruent), IPC stands for integration by partial congruence, retained in the present study (at least two lines of evidence are necessary); IC stands for integration by cumulation (one line of evidence is sufficient). Species are represented in split red bars and subspecies in yellow.

…

. Summary of the stepwise Canonical Discrimi- nant Function Analysis (CDFA) for the size/shape data set (obtained after using an isometric approach) and pholidosis

…

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An integrative taxonomic revision of the Tarentola

geckos (Squamata, Phyllodactylidae) of the Cape

Verde Islands

RAQUEL VASCONCELOS1,2, ANA PERERA1, PHILIPPE GENIEZ3, D. JAMES HARRIS1,2

and SALVADOR CARRANZA4*

1CIBIO-UP, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do

Porto, Campus Agrário de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal

2Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, R. Campo Alegre, s/n,

4169-007 Porto, Portugal

3EPHE – UMR 5175, Centre d’Ecologie Fonctionnelle et Evolutive, 1919 route de Mende, 34293

Montpellier cedex 05, France

4Institute of Evolutionary Biology (CSIC-UPF) – Passeig Marítim de la Barceloneta, 37-49, E-08003

Barcelona, Spain

Received 5 March 2011; revised 10 June 2011; accepted for publication 28 June 2011

Recent phylogeographical analyses using mitochondrial DNA (mtDNA) sequences indicate that the Tarentola

geckos from the Cape Verde archipelago originated from a propagule that dispersed from the Canary Islands

approximately 7.7 Mya and that underwent a fast evolutionary radiation. Molecular analyses carried out to date

clearly show some incongruences with the current taxonomy of Tarentola from the Cape Verde Islands, with some

species being paraphyletic or polyphyletic, and several independently evolving lineages needing formal taxonomic

recognition. The aim of this study was to clarify the systematics of this group to unravel its taxonomy by applying

an integrative approach based on information from three independent sources: mtDNA, nuclear genes, and

morphology. As a result of this taxonomic revision, two novel species for the islands of S. Nicolau and Fogo are

described and eight subspecies are upgraded to species level. Moreover, an identiﬁcation key for the genus

Tarentola from the Cape Verde archipelago is presented. This study reconciles taxonomy and phylogeny in this

group, provides a better understanding of diversity patterns, new insights on evolutionary hypotheses, and

supports the basic framework for the future management and conservation of this unique reptile radiation.

doi: 10.1111/j.1096-3642.2011.00768.x

ADDITIONAL KEYWORDS: Cape Verdean – geckonids – morphology – nDNA – species description.

INTRODUCTION

Delineating species boundaries is crucial because it is

the ﬁrst step toward discussing broader questions on

biogeography, ecology, conservation, or evolution. The

gap in communication between different disciplines

currently related to species recognition is an important

but often overlooked problem. According to de Queiroz

(2007) one of the main problems is that species delimi-

tation has long been confused with that of species

conceptualization, leading to a half a century of con-

troversy concerning both the deﬁnition of species cat-

egories and the methods for inferring the boundaries

and the number of species. Recent intellectual progress

in the ﬁeld has been made to identify a common

element among all the different species concepts in

order to propose a single, more general, concept of

*Corresponding author. E-mail:

salvador.carranza@ibe.upf-csic.es

Zoological Journal of the Linnean Society, 2012, 164, 328–360. With 7 ﬁgures

species known as the General Lineage Species Concept

(de Queiroz, 1998). This uniﬁed species concept con-

siders species as separately evolving metapopulation

lineages and treats this property as the single requisite

for delimiting species. Other properties, such as phe-

netic distinguishability, reciprocal monophyly, and pre-

and postzygotic reproductive isolation, are not part of

the species concept but serve as important lines of

evidence relevant to assess the separation of lineages

and therefore to species delimitation (de Queiroz,

2007). The divorce between conceptualization and

delimitation of species and the proposal of a uniﬁed

species concept has shifted emphasis away from the

controversy of species criteria, concentrating efforts in

the development of new approaches for species delimi-

tation, as for example with ‘integrative taxonomy’

(Dayrat, 2005; Cardoso, Serrano & Vogler, 2009; Padial

et al., 2010). The goal of integrative taxonomy is to

delimit the units of biotic diversity from multiple and

complementary disciplines (e.g. phylogeography, popu-

lation genetics, comparative morphology, or ecology).

Hence, molecular markers, population genetic tests,

morphological features and ecological characteristics

should be used as different complementary approaches

to achieve reliable identiﬁcations of species. All sets of

characters have the same weight during the process of

recognizing and diagnosing species and the goal is

to use as many as possible. Species delineation is

therefore regarded as an objective scientiﬁc process

that results in a taxonomic hypothesis. In this way,

the level of conﬁdence in the taxonomic hypothesis

supported by several independent character sets is

much higher than for species supported by only one

(Schlick-Steiner et al., 2010). Such an integrative view

is especially useful in the case of taxonomic groups that

are morphologically conservative such as the geckos

(Jesus, Brehm & Harris, 2002), where cryptic species

have probably been overlooked (Perera & Harris,

2010).

Tarentola is a genus of the family Phyllodactylidae

with around 20 species commonly called wall geckos. All

have robust bodies, non-divided subdigital lamellae,

and well-developed claws on the third and fourth digits

(Arnold & Ovenden, 2002) and, with the only exception

of Tarentola chazaliae (Mocquard, 1895), have a con-

servative morphology (Joger, 1984a; Carranza et al.,

2002; Harris et al., 2004). These climbing geckos are

mostly active by night and typically inhabit dry, open,

rocky areas but also artiﬁcial habitats (Arnold &

Ovenden, 2002). The genus is distributed across south-

ern Europe, Mediterranean islands, North Africa, and

on many islands of the Macaronesian region, namely

Madeira (including Selvagens), Canary Islands, and

Cape Verde Islands (Arnold & Ovenden, 2002; Sindaco

& Jeremcˇenko, 2008). On the other side of the Atlantic

Ocean, three species are accepted: T. americana (Gray,

1831), from Cuba and the Bahamas; the recently

described T. crombiei Díaz & Hedges, 2008 endemic to

Cuba; and the probably extinct T. albertschwartzi

Sprackland & Swinney, 1998, known from a single

specimen allegedly from Jamaica.

Tarentola members were divided into ﬁve different

subgenera based on anatomical, biochemical, immu-

nological, and phylogenetical data (Joger, 1984a; Car-

ranza et al., 2000): Sahelogecko and Saharogecko in

North Africa; Tarentola sensu stricto in North Africa,

southern Europe, and the eastern Canary Islands;

Neotarentola, which includes T. americana,T. crom-

biei and T. albertschwartzi (Weiss & Hedges, 2007);

and Makariogecko in the Macaronesian Islands (Car-

ranza et al., 2000; Weiss & Hedges, 2007). The sub-

genus Makariogecko presents a synapomorphy: the

supraciliar scales are larger than the remaining inter-

orbital scales and they are divided (Joger, 1984a).

Nevertheless, recent molecular phylogenies including

Tarentola chazaliae (previously Geckonia chazaliae)

do not seem to support the monophyly of this subge-

nus (Carranza et al., 2002). Within this subgenus, the

Tarentola from Cape Verde are especially interesting

as they originated from a single colonization event by

a propagule that rafted southwards from the western

Canary Islands (Carranza et al., 2000) around

7.7 Mya (Vasconcelos, Carranza & Harris, 2010).

The Cape Verde Islands are a volcanic archipelago

located approximately 500 km off the West African

coast with ten main islands, plus several islets, which

are topologically divided into north-western, eastern

and southern island groups (Fig. 1). The radiation of

the geckos after the single colonization event gave

origin to four currently accepted endemic species with

several subspecies, some of them exclusive to one

of these island groups: T. darwini Joger 1984b,

T. caboverdiana Schleich 1984, T. rudis Boulenger,

1906, and T. gigas (Bocage, 1875). However, the most

exhaustive recent revision regarding the genetic vari-

ability of Tarentola from the Cape Verde Islands using

mitochondrial markers recovered 15 evolutionary sig-

niﬁcant units (ESUs) arranged into four main groups

(Fig. 2; Vasconcelos et al., 2010) not completely congru-

ent with the current taxonomy (Schleich, 1987; Joger,

1993). The ﬁrst group included all T. darwini plus

T. rudis from Boavista, although both the bootstrap

and posterior probability (PP) values were low; the

second one grouped T. caboverdiana from São Vicente,

Santa Luzia, Branco, Raso, and Santo Antão; the third

one was exclusively formed by T. caboverdiana nico-

lauensis from São Nicolau; ﬁnally, the fourth group

included the remaining T. rudis populations. From all

the accepted Cape Verdean Tarentola, only T. gigas

and T. darwini are monophyletic based on mitochon-

drial data (Fig. 2), with T. rudis and T. caboverdiana

being poly- and paraphyletic, respectively (Vasconcelos

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 329

et al., 2010). As a result, previous molecular phyloge-

netic studies have always stressed that a review of the

systematics of the Cape Verdean Tarentola was needed

(Carranza et al., 2000, 2002; Vasconcelos et al., 2010).

In the case of T. rudis, the mitochondrial lineages of

each subspecies (T. r. boavistensis,T. r. rudis,T. r. pro-

togigas, and T. r. maioensis) seem to be quite divergent

among them and from all other forms. Moreover, the

T. r. boavistensis mitochondrial lineage is closer to the

T. darwini clade than to the other clade containing the

four remaining subspecies of T. rudis, and T. r. rudis

forms a clade with T. gigas, turning T. rudis into a

polyphyletic species. Also, T. ‘caboverdiana’nicolauen-

sis is more closely related to T. gigas and to the four

subspecies of T. rudis (T. r. rudis,T. r. protogigas T.

rudis hartogi, and T. r. maioensis) than to the other

T. caboverdiana subspecies (see Vasconcelos et al.,

2010). As effective conservation measures depend

largely on a good knowledge of the taxonomy of the

species (Mace, 2004), the present taxonomic revision is

clearly needed not only to clarify the systematics of

this group but also as a basic framework for the future

conservation management of the Tarentola geckos

from Cape Verde.

In order to describe new taxa, intraspeciﬁc variabil-

ity should be studied and a taxonomic revision should

be made, with all previous synonyms and chresonyms

identiﬁed (Dayrat, 2005). Genetic assessment regard-

ing the Tarentola geckos of the Cape Verde Islands

was accomplished in previous works (see Carranza

et al., 2000; Vasconcelos et al., 2010), although using

only mitochondrial markers. Therefore, in the present

work, information from mitochondrial DNA (mtDNA),

three nuclear markers, and morphology is used fol-

lowing an ‘integrative taxonomy’ approach to revise

the systematics of the genus Tarentola from the Cape

Verde archipelago and to fully reconcile taxonomy

with phylogeny. The results of this work are very

Figure 1. Map of the Cape Verde Islands showing the geographical location (latitudes and longitudes) and altitudes of

the islands and the origins of the new Tarentola samples included in the genetic (circles) and morphological (diamonds)

analyses (Geographic Coordinate System, Datum WGS 84). Island and taxa colours match the colours used on the network

analyses. No specimens were found on Sal.

330 R. VASCONCELOS ET AL.

relevant for a better understanding of diversity pat-

terns, for providing new insight into evolutionary

hypotheses, and for the conservation of this unique

island radiation.

MATERIAL AND METHODS

ORIGIN OF TISSUE SAMPLES AND SPECIMENS

A total of 135 sequences of Cape Verdean Tarentola

were included in the genetic analyses of the nuclear

data and 92 in the multivariate morphological analy-

sis. All specimens were identiﬁed in the ﬁeld using

diagnostic characters published by Joger (1984b,

1993) and Schleich (1987) and a piece of tail was

removed and stored in 96% ethanol. Before the

animals were released, digital photographs (from

dorsal, ventral, and lateral parts) were taken to quali-

tatively analyse the colour pattern characteristics

that may disappear in preserved specimens and to

perform pholidotic counts a posteriori. Then, 2051 of

these photographs were deposited in MorphoBank

(http://www.morphobank.org/; see Appendix 1).

Figure 2. Phylogenetic relationships of endemic Cape Verde Tarentola taxa and their relatives from the Canary Islands

modiﬁed from Vasconcelos et al. (2010) based on cytochrome band 12S rRNA genes. The tree was inferred using maximum

likelihood (ML) and GTR+I+G model of sequence evolution (log likelihood =-6468.896) and was rooted using Tarentola

americana. Bootstrap support values above 60% for the ML analysis are shown below nodes. Posterior probability (PP)

values higher than 95% for the Bayesian analysis are represented by an asterisk (*) and are shown above nodes. Names

in bold follow the new taxonomic proposal and non-bold ones the taxonomy accepted in previous recent papers (Carranza

et al., 2000; Jesus et al., 2002; Vasconcelos et al., 2010). For further details see Vasconcelos et al. (2010). Characters

immediately to the right of island names correspond to the 15 evolutionarily signiﬁcant units (ESUs) of A, B, C, and D

clades recognized in the present work and represented in split green bars. Lines of evidence (in grey): 1, mitochondrial

DNA (independent cyt bparsimony networks with a connection limit of 95%; see Appendix 3); 2, nuclear DNA (absence

of shared haplotypes in MC1R); 3, morphology (detection of any diagnostic morphological character or a set of a unique

combination of characters). Integration approaches (in red) from the most conservative to the most inﬂationist: ITC stands

for integration by total congruence (all lines of evidence should be congruent), IPC stands for integration by partial

congruence, retained in the present study (at least two lines of evidence are necessary); IC stands for integration by

cumulation (one line of evidence is sufficient). Species are represented in split red bars and subspecies in yellow.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 331

A total of 352 specimens of all taxa of Cape Verdean

Tarentola were observed and/or measured and a total

of 119 Cape Verdean voucher specimens of all taxa

were also examined. Examined vouchers are depos-

ited at the British Museum of Natural History

(BMNH), London, at the Centre d’Ecologie Fonction-

nelle et Evolutive, Montpellier, but previously housed

at the Laboratoire de Biogéographie et Ecologie des

Vertebrés collection (BEV), Departamento de Biología

de la Universidad de Las Palmas de Gran Canaria

(DBULPGC), and at the Museum National d’Histoire

Naturelle (MNHN), Paris. Identiﬁcation codes, locali-

ties, and GenBank and MorphoBank accession

numbers of the live and voucher specimens examined

are listed in Appendix 1. In addition, specimen data

from other authors used in the taxonomic revision are

included in the ‘Additional material and references’

section under each taxon and duly cited to be distin-

guished from data gathered on the present study.

MUSEUMS ACRONYMS

Laboratoire de Biogéographie et Ecologie des Verte-

brés collection (BEV), Montpellier; British Museum

of Natural History (BMNH), London; Centro de Zoo-

logia, Instituto de Investigação Cientíﬁca Tropical

(IICT), Lisbon; Departamento de Biología de la

Universidad de Las Palmas de Gran Canaria (DB-

ULPGC), Canary Islands; Museum der Universitat

Helsingfors (MUH), Helsinki; Gabinete d’Ajuda (GA),

Lisbon; Hessisches Landesmuseum Darmstadt

(HLMD), Darmstadt; Hessisches Landesmuseum

Wiesbaden (HLMW), Wiesbaden; Jon Boone collection

(JB); Museu Civico ‘G. Doria’ di storia Naturale

de Genova (MSNG), Genoa; Museu di Zoologia

dell’Università degli Studi di Torino (MZUT), Turin;

Muséum National d’Histoire Naturelle (MNHN),

Paris; Institut Fondamental d’Afrique Noire (IFAN),

Dakar; Rijksmuseum van Natuurlijke Historie

(RMNH), Leiden; Senckenberg-Museum Fors-

chungsinstitut (SMF), Frankfurt; Universidade da

Madeira (CCBG), Funchal; Zoologische Staatssam-

mlung München (ZSM and ZSMH), Munich; Zoologis-

ches Forschungsinstitut und Museum Alexander

Koenig (ZFMK), Bonn; Zoologisches Museum Berlin

(ZMB), Berlin; Zoologisches Museum der Universität

Hamburg (ZMH), Hamburg.

GENETIC ANALYSES

Total genomic DNA was extracted from small pieces

of tail using standard methods. Three fragments of

nuclear genes were analysed: phosducin (PDC), ace-

tylcholinergic receptor M4 (ACM4), and melanocortin

1 receptor (MC1R). The sets of primers used were:

PHOR1 and PHOF2, and Tg-F and Tg-R (Gamble

et al., 2008) for the PDC and ACM4 fragments,

respectively, and MC1R-F and MC1R-R (Pinho et al.,

2010) for the MC1R fragment. For ampliﬁcation of

these three fragments, an initial denaturation step of

95 °C for 90 s was used, followed by 35 cycles of 95 °C

for 30 s, 50 °C (annealing temperature) for 45 s and

72 °C (extending temperature) for 90 s, and a ﬁnal

extension at 72 °C for 7 min. Ampliﬁed nuclear DNA

(nDNA) fragments were sequenced from both strands

with the same primers used in the ampliﬁcation

process. Sequences were aligned manually using

BIOEDIT v.7.0.4. (Hall, 1999). The Bayesian algo-

rithm implemented in the program PHASE 2.1.1

(Stephens, Smith & Donnelly, 2001) was used to

reconstruct haplotypes from population genotyped

data. Sequence pairs with probability lower than 0.7

were not included in posterior analyses.

POPULATION ANALYSES

The genealogical relationships between taxa were

assessed with haplotype networks constructed using

statistical parsimony (Templeton, Crandall & Sing,

1992), implemented in the program TCS v.1.21

(Clement, Posada & Crandall, 2000), with a connec-

tion limit of 95%. Haplotypes were then arranged in

groups based on the 15 ESUs recovered in the mito-

chondrial study by Vasconcelos et al. (2010). Genetic

differentiation between ESUs for the three nuclear

genes was calculated using the nearest neighbour

statistic, Snn (Hudson, 2000), implemented in the

program DnaSP v.5 (Rozas et al., 2003) and tested

with 1000 permutations. Additionally, estimates of

evolutionary divergence (p-dist) over 302 bp of cyto-

chrome b(cyt b) sequences among the 15 ESUs were

calculated with MEGA4 (Tamura et al., 2007). All cyt

bsequences used (GenBank accession numbers

Q380699–Q381129) were from Vasconcelos et al.

(2010).

The IMa software (Hey & Nielsen, 2007), which

takes into account population divergence and gene

ﬂow in the same framework, was used to disentangle

the relative effects of isolation and migration in

shaping the patterns of variation among diverging

very similar species occurring on the same island and

sharing nuclear haplotypes, as was the case of the

two Tarentola from S. Nicolau. This software uses a

Markov Chain Monte Carlo (MCMC) sampling of gene

genealogies to PP distributions of rates of migration

in either direction (m1and m2) and time of divergence

(t), among other parameters. The assumption made

by IMa of no recombination was tested with DnaSP

v.5 (Rozas et al., 2003) coalescent simulations. After

two experimental runs to assess appropriate para-

meter settings and ensure proper mixing, IMa was

run three times for the two-species data set for 50

332 R. VASCONCELOS ET AL.

million steps along the Markov Chain after 10 million

steps of burn-in with ten Metropolis-coupled chains

with linear heating. The mixing properties of the

MCMC were checked by monitoring the values of

the parameters and the trend-line plots of the

parameters.

MORPHOLOGICAL ANALYSES

A multivariate analysis of the three populations pre-

viously described as T. ‘darwini’ from the islands of

Fogo, S. Nicolau, and Santiago (Fig. 2) was performed

to assess if diversity existed and, if so, which level of

morphological distinctiveness these populations pre-

sented. Several morphological characters from 153

individuals (ind.) of other groups were also measured

to disentangle complex relationships detected at the

mitochondrial level, such as between T. protogigas

from Fogo and Brava islands (Appendix 2).

As ﬁxation and preservation in museums may

deform bodies or some body parts, hampering the

comparison with live specimens (Vervust, Dongen &

Van Damme, 2009), no vouchers were included in this

analysis, and only live adult specimens that had been

genetically conﬁrmed using the cyt bmitochondrial

marker were used. Sex was determined by the pres-

ence of enlarged spurs and more developed cloacal

pouches in males (Barbadillo et al., 1999) and by their

larger body size and robustness (Arnold & Ovenden,

2002). Details on the live specimens analysed are

listed in Appendix 1. Morphological variation was

assessed using both morphometric and meristic vari-

ables (14 linear body measurements and seven pholi-

dotic variables, respectively). Bilateral variables

(Table 1 and Appendix 2) were taken from the same

side of the animals whenever possible.

All 14 linear body measurements were recorded in

the ﬁeld by the same person (A.P.) using a ruler (for

snout–vent length, SVL, with accuracy to the nearest

0.1 mm) and a digital calliper (all the remaining

variables with accuracy to the nearest 0.01 mm) and

were expressed in millimetres. Trunk length (TrL)

was measured from the posterior edge of forelimb

insertion to the anterior edge of hindlimb insertion

and the tail width (TW) was recorded at its widest

point. The total lengths of front (FLL) and hindlimbs

(HLL) from the longest toe to the base of the limb

were measured. Also, the partial lengths of front

(CFL) and hind (FFL) limbs were measured from the

tip of the longest toe to the elbow or knee inﬂexion

point, respectively. Head width (HW) was measured

at its widest part, usually at the level of the temporal

region, and maximum head height (HH) was mea-

sured from occiput to jaws. Ear length (EL) and eye

diameter (OD) were considered the longest dimension

of ear and ocular orbit, respectively. Nostril–eye

(NED) and snout–eye distances (SED) were measured

from the anterior border of the ocular orbit to the

posterior margin of the right nostril and snout,

respectively. Ear–eye distance (EED) was measured

from the anterior border of the ear to the posterior

border of the ocular orbit.

Pholidotic (meristic) variables recorded included

the number of supra- and infra-labial scales (SLS and

ILS, respectively) counted until the limit of the mouth

opening, and the number of non-divided, enlarged

side-to-side lamellae under the fourth hind toe (Lam).

The number of transverse (Trow and Srow) and lon-

gitudinal (Tline and medS) tubercles and scales in the

dorsum, respectively, were counted paramedially. The

number of small scale rows (Srow) in the vertebral

line was counted in the midbody, in a midline between

the front and hindlimbs, between the upper and lower

rows of tubercles. The number of small scales lines

(medS) was estimated by the mean number of scales

between tubercles on the intersection of the midbody

line with the vertebral line.

Prior to the analysis, linear measurements were log

transformed and checked for homoscedasticity (Lillie-

ford test) and normality (Levene test). As linear body

measurements are correlated with body size (P<0.05

in all cases), body-size-corrected variables were esti-

mated using an isometric correction (Somers, 1986) to

investigate the existence of possible differentiation

patterns not related to body size. For this, an isomet-

ric vector was created in which all linear measure-

ments (log transformed) were projected, in order to

obtain a multivariate representation of the isometric

size of each individual (SIZE). After that, each vari-

able was regressed on this isometric vector. The

obtained residuals for each variable were used as

size-corrected variables (Kaliontzopoulou, Carretero

& Llorente, 2010). The multivariate representation of

the isometric size (SIZE) was used as size estimator,

while the remaining size-corrected variables were

considered as shape estimators.

MANOVAs were used to analyse the effect of sex,

population, and their interaction (sex*population) on

all linear (both sets, raw log-transformed and size-

corrected) and pholidotic variables.

To assess the generalised morphological patterns

within the different populations previously assigned

as T. ‘darwini’, a stepwise Canonical Discriminant

Function Analysis (CDFA) was performed on all mer-

istic and size-corrected linear variables. Due to the

different degree of sexual dimorphism observed

between populations in some of the variables, multi-

variate analysis was performed on males and females

separately. This multivariate approach maximizes dif-

ferences between a priori deﬁned groups from differ-

ent island populations (mtDNA clades A2–A4) and

classiﬁes the individuals based on CDFs. Only 30 of

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 333

Table 1. Descriptive statistics for all the linear measurements and meristic variables of adult specimens of the different

Tarentola taxa included in the multivariate analysis

T. bocagei

São Nicolau

Males (N=19) Females (N=14) All (N=33)

Mean ± SD Range Mean ± SD Range Mean ± SD Range

SVL 60.00 ± 3.45 54.00–65.50 55.82 ± 3.34 49.00–63.50 58.23 ± 3.95 49.00–65.50

TrL 23.04 ± 1.64 19.40–26.19 21.12 ± 1.31 19.17–24.47 22.22 ± 1.77 19.17–26.19

TW 7.23 ± 0.65 5.86–8.06 6.25 ± 0.70 5.08–7.40 6.82 ± 0.83 5.08–8.06

FLL 20.64 ± 1.37 18.27–24.13 18.89 ± 1.61 15.94–21.09 19.90 ± 1.70 15.94–24.13

CFL 13.06 ± 0.98 11.16–15.06 11.98 ± 1.09 10.08–13.76 12.60 ± 1.15 10.08–15.06

HLL 25.18 ± 1.44 22.52–27.97 23.86 ± 1.70 20.6–26.34 24.62 ± 1.67 20.60–27.97

FFL 14.61 ± 0.95 12.97–16.45 13.81 ± 1.10 12.18–15.58 14.27 ± 1.08 12.18–16.45

HW 13.56 ± 0.78 12.48–15.32 12.45 ± 0.75 11.10–13.85 13.09 ± 0.94 11.10–15.32

HH 8.28 ± 0.48 7.17–9.17 7.43 ± 0.47 6.67–8.19 7.94 ± 0.63 6.67–9.17

OD 3.64 ± 0.21 3.38–4.02 3.67 ± 0.20 3.16–3.85 3.65 ± 0.20 3.16–4.02

EL 2.73 ± 0.26 2.05–3.20 2.68 ± 0.26 2.22–3.20 2.71 ± 0.26 2.05–3.20

NED 5.73 ± 0.48 5.07–6.93 5.22 ± 0.39 4.50–5.68 5.52 ± 0.50 4.50–6.93

SED 6.99 ± 0.71 5.93–8.30 6.41 ± 0.51 5.55–7.80 6.74 ± 0.69 5.55–8.30

EED 5.43 ± 0.40 4.62–6.01 4.96 ± 0.39 4.17–5.45 5.23 ± 0.46 4.17–6.01

SLS 11.05 ± 0.85 10–13 10.93 ± 0.83 10–12 11.00 ± 0.83 10–13

ILS 9.11 ± 0.74 8–10 8.43 ± 0.51 8–9 8.82 ± 0.73 8–10

Lam 9.32 ± 0.75 8–10 9.57 ± 0.65 8–10 9.42 ± 0.71 8–10

Trow 16.11 ± 1.20 14–18 15.57 ± 1.22 14–18 15.88 ± 1.22 14–18

Tline 20.74 ± 1.91 17–24 20.21 ± 1.31 18–22 20.52 ± 1.68 17–24

Srow 2.39 ± 0.49 1.50–3.00 2.71 ± 0.43 2.00–3.00 2.53 ± 0.48 1.50–3.00

medS 2.00 ± 0.24 1.50–2.50 2.13 ± 0.42 1.25–2.75 2.05 ± 0.33 1.25–2.75

T. fogoensis

Fogo

Males (N=16) Females (N=13) All (N=29)

Mean ± SD Range Mean ± SD Range Mean ± SD Range

SVL 61.84 ± 4.91 54.00–69.50 55.54 ± 5.81 48.50–69.00 59.02 ± 6.13 48.50–69.50

TrL 25.80 ± 2.82 20.98–32.57 23.23 ± 3.45 20.60–31.97 24.65 ± 3.33 20.60–32.57

TW 6.48 ± 1.09 4.80–8.64 5.07 ± 0.83 3.64–6.60 5.85 ± 1.20 3.64–8.64

FLL 22.78 ± 1.69 19.81–26.37 21.03 ± 1.68 19.22–24.66 22.00 ± 1.88 19.22–26.37

CFL 14.55 ± 1.23 12.10–16.36 13.52 ± 1.41 12.05–17.07 14.08 ± 1.39 12.05–17.07

HLL 29.11 ± 1.74 25.40–31.78 26.05 ± 2.28 21.77–31.07 27.74 ± 2.50 21.77–31.78

FFL 16.84 ± 1.15 14.61–18.58 14.86 ± 1.13 12.93–16.60 15.96 ± 1.50 12.93–18.58

HW 13.95 ± 1.20 12.22–15.97 12.49 ± 1.30 10.75–15.70 13.29 ± 1.43 10.75–15.97

HH 8.41 ± 0.70 7.67–10.16 7.44 ± 0.74 6.64–8.87 7.98 ± 0.86 6.64–10.16

OD 3.74 ± 0.23 3.39–4.16 3.46 ± 0.27 3.09–4.12 3.62 ± 0.28 3.09–4.16

EL 2.64 ± 0.30 2.07–3.05 2.33 ± 0.25 1.87–2.70 2.50 ± 0.31 1.87–3.05

NED 6.30 ± 0.44 5.43–7.00 5.81 ± 0.45 5.36–6.69 6.08 ± 0.50 5.36–7.00

SED 7.96 ± 0.57 6.69–8.67 7.23 ± 0.50 6.60–8.01 7.63 ± 0.65 6.60–8.67

EED 5.91 ± 0.66 4.92–7.24 5.21 ± 0.61 4.34–6.52 5.60 ± 0.72 4.34–7.24

SLS 10.73 ± 0.80 10–12 11.08 ± 0.79 10–12 10.89 ± 0.80 10–12

ILS 8.75 ± 0.58 8–10 9.15 ± 0.80 8–11 8.93 ± 0.70 8–11

Lam 10.07 ± 0.46 9–11 10.27 ± 0.79 9–11 10.15 ± 0.61 9–11

Trow 15.50 ± 1.37 14–18 15.08 ± 0.76 14–17 15.31 ± 1.14 14–18

Tline 22.94 ± 1.95 20–27 22.62 ± 2.36 20–27 22.79 ± 2.11 20–27

Srow 2.56 ± 0.36 2.00–3.00 2.38 ± 0.51 1.50–3.00 2.48 ± 0.43 1.50–3.00

medS 1.59 ± 0.40 1.00–2.50 1.69 ± 0.47 1.00–2.50 1.64 ± 0.43 1.00–2.50

334 R. VASCONCELOS ET AL.

the 88 individuals from Santiago were randomly

included in the analyses to avoid bias of results due

to uneven samples sizes. The leave-one-out option

was implemented to cross-validate the classiﬁcation

results. As this procedure generates individual clas-

siﬁcations using discriminant functions based on all

observations except the given case, it provides a more

accurate estimate of the classiﬁcation values. Statis-

tical analyses were performed using R (R Develop-

ment Core Team, 2010).

INTEGRATIVE APPROACH

For consistency, the same approach used in the taxo-

nomic revision of the endemic Cape Verdean skink

genus Chioninia (Miralles et al., 2010) was followed

in this study. The mitochondrial phylogenetic tree

(Fig. 2) adapted from Vasconcelos et al. (2010) was

used as a framework to investigate the taxonomy of

the Cape Verdean Tarentola. Three lines of evidence

have been deﬁned on the basis of the alleged inde-

pendence of their respective data sets (mtDNA,

nDNA, and morphology) to decide the taxonomic

status of each ESU (see Fig. 2). Each of these lines

represents equivalent, independent, and combinable

indicators able to detect splits between different

species: (1) mtDNA – presence of independent cyt b

parsimony networks with a connection limit of 95%

(see Hart & Sunday, 2007). The results of the cyt b

networks analyses are from Vasconcelos et al. (2010)

and are presented in Appendix 3. (2) nDNA – absence

of shared haplotypes in the MC1R nuclear gene (see

Monaghan et al., 2009). The other two genes (PDC

and ACM4) were not used as lines of evidence because

both presented a very low level of genetic variability

and a clear pattern of incomplete lineage sorting (see

below). (3) Morphology – detection of at least one ﬁxed

diagnostic character state (e.g. presence or absence

for qualitative characters or non-overlapping values

for meristic or allometric characters) or a set of a

unique combination of characters that might be

strong evidence of reduced or absence of gene ﬂow

(Wiens & Servedio, 2000).

Different possible integration approaches are pre-

sented in Figure 2, ranging from the most conserva-

tive to the most inﬂationist. Integration by total

congruence (ITC) was achieved by retaining only the

candidate species that are supported by all the three

lines of evidence, whereas integration by cumulation

(IC) was calculated considering that one line of

Table 1. Continued

T. darwini

Santiago

Males (N=15) Females (N=15) All (N=30)

Mean ± SD Range Mean ± SD Range Mean ± SD Range

SVL 55.77 ± 5.39 45.00–64.00 56.70 ± 2.81 51.00–60.00 56.23 ± 4.25 45.00–64.00

TrL 23.75 ± 3.07 18.40–30.00 24.49 ± 1.56 20.90–26.20 24.12 ± 2.42 18.40–30.00

TW 6.25 ± 1.04 4.30–7.60 5.92 ± 0.65 4.80–7.10 6.09 ± 0.87 4.30–7.60

FLL 19.71 ± 1.83 15.80–23.10 19.42 ± 1.06 18.00–21.30 19.56 ± 1.47 15.80–23.10

CFL 12.86 ± 1.52 10.40–15.00 13.06 ± 1.14 9.90–14.50 12.96 ± 1.32 9.90–15.00

HLL 24.03 ± 2.15 20.50–28.60 24.24 ± 1.59 21.90–26.90 24.13 ± 1.86 20.50–28.60

FFL 13.90 ± 1.69 11.20–16.60 14.12 ± 0.73 13.20–15.40 14.01 ± 1.28 11.20–16.60

HW 12.59 ± 1.52 9.80–14.80 12.71 ± 0.69 11.60–13.80 12.65 ± 1.16 9.80–14.80

HH 7.67 ± 1.04 5.70–9.00 7.76 ± 0.45 7.00–8.50 7.72 ± 0.79 5.70–9.00

OD 3.23 ± 0.28 2.70–3.60 3.32 ± 0.24 2.90–3.70 3.28 ± 0.26 2.70–3.70

EL 2.31 ± 0.32 1.90–2.80 2.20 ± 0.24 1.90–2.70 2.26 ± 0.28 1.90–2.80

NED 5.48 ± 0.55 4.60–6.50 5.61 ± 0.40 4.80–6.40 5.54 ± 0.48 4.60–6.50

SED 6.99 ± 0.67 6.00–8.20 7.10 ± 0.47 6.40–8.10 7.04 ± 0.57 6.00–8.20

EED 5.37 ± 0.64 4.20–6.60 5.44 ± 0.45 4.80–6.10 5.41 ± 0.54 4.20–6.60

SLS 10.29 ± 0.73 9–12 9.93 ± 0.88 9–12 10.1 ± 0.82 9–12

ILS 8.43 ± 0.51 8–9 8.33 ± 0.62 7–9 8.38 ± 0.56 7–9

Lam 9.73 ± 0.9 8–11 9.64 ± 0.81 8–11 9.68 ± 0.84 8–11

Trow 16.27 ± 0.88 15–18 15.67 ± 1.45 13–18 15.97 ± 1.22 13–18

Tline 23.2 ± 2.18 21–27 22 ± 2.65 17–26 22.6 ± 2.46 17–27

Srow 2.37 ± 0.55 1.5–3.5 2.17 ± 0.24 2–2.5 2.27 ± 0.43 1.5–3.5

medS 1.53 ± 0.3 1.0–2.0 1.58 ± 0.42 1.0–2.5 1.56 ± 0.36 1.0–2.5

For each variable, mean ± standard deviation (SD), range, and sample size (N) is given.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 335

evidence was sufficient for splitting taxa. However,

both methods have a tendency to under- and overes-

timate the number of species, respectively (see Padial

et al., 2010). Hence, a third approach was deﬁned,

integration by partial congruence (IPC), which is

intermediate between the two previous ones, as it

retains only candidate species that are supported by

at least two independent lines of evidence. This

approach does not preclude the discovery of species

supported by only one line of evidence. Also, as in

Miralles et al. (2010), splits supported by only one of

these three lines of evidence within infraspeciﬁc

allopatric ESUs have been considered as different

subspecies.

RESULTS

MOLECULAR DATA

The PDC and ACM4 networks recovered similar gene-

alogies, with a similar number of haplotypes (13 and

14, respectively; Fig. 3) and a different topology to the

one recovered with the MC1R fragment. PDC and

ACM4 network analyses recovered the central and

most common haplotype as the ancestral one, shared

by many different taxa and surrounded by several

singletons for most of the taxa groups. The only three

exceptions were found in the PDC gene, which pre-

sented three non-ancestral haplotypes shared by

geckos from S. Vicente and Santo Antão, specimens of

lineage D3 and D4 from Fogo and Brava, respectively,

and another one by some specimens from lineages A2

and C from S. Nicolau (see Fig. 3). On the other hand,

the MC1R network recovered a greater number of

haplotypes, 36 (including 23 for the same individuals

sequenced for the other genes), and an increased level

of substructuring among taxa, especially for each of

the three Tarentola from clade B that do not share

haplotypes and for the endemic Tarentola from

Boavista and T. darwini from Santiago (lineages A1

and A4 in Fig. 2, respectively; see Fig. 3). As expected

from the results of the other two nuclear markers,

MC1R also presents some sharing of ancestral hap-

lotypes between specimens from S. Nicolau and Fogo

(lineages A2 and A3 in Fig. 2, respectively) and also

between most of the specimens analysed from the two

species from S. Nicolau (lineages A2 and C in Fig. 2)

and some T. caboverdiana specimens from Santo

Antão, T. gigas specimens from Raso, and T. protogi-

gas specimens from Brava and Fogo (lineages B3, D1,

D3, and D4 in Fig. 2, respectively; see Fig. 3). More-

over, some recent haplotypes were shared by all speci-

mens of T. rudis from Santiago and the Tarentola

from Maio and by T. protogigas specimens from Fogo

and Brava, respectively.

Three independent runs using IMa software con-

verged on approximate marginal PP distributions.

Reliable estimates of m1,m2, and tbetween the two

Tarentola taxa occurring on S. Nicolau were obtained

to study the introgression versus ancestral polymor-

phism hypotheses. The migration rate curves pre-

sented a clear peak although their tails did not reach

zero, suggesting a high probability of no gene ﬂow in

either direction between these two populations and of

tdiffering from zero (see Appendix 4). This t-value

suggests that these two ESUs have indeed diverged.

MORPHOLOGICAL DATA

In general, males and females were different in size

(MANOVA P<0.001) but not in shape or pholidosis

(in both cases MANOVA P>0.05), while the three

populations (A2, A3, and A4) compared were different

in all the datasets analysed (size, shape, and pholi-

dosis, in almost all cases MANOVA P<0.001; see

Table 2). Populations had a similar degree of sexual

dimorphism (interaction sex*population) in pholidosis

and shape (MANOVAs P>0.005 in both cases), but

not in size (MANOVA P<0.01; see Table 2).

Regarding the linear measurements, the ANOVA

using raw log-transformed variables showed a clear

sexual dimorphism in all the variables, except in OD

(Table 2). However, such differences largely disap-

peared when body size-corrected variables were

compared, with the exception of TW and OD (Table 2).

Regarding the differences between the three popula-

tions compared, most raw log-transformed vari-

ables were signiﬁcantly different, even after

correcting them for body size (Table 2). The

sex*population interaction was signiﬁcant for most of

the characters using log-transformed but not size-

corrected variables. So, all raw log variables with the

exception of FLL, CFL, and EL were signiﬁcant, but

almost (with the exception of OD) all differences dis-

appeared when considering size-corrected variables

(Table 2).

Regarding the meristic variables, males and

females only differed in the number of dorsal trans-

verse rows of tubercles (Trow; Tables 1 and 2, Appen-

dix 2). All meristic variables, with the exception of

Trow and Srow, were statistically different between

populations (Table 2). However, all scale counts, with

the exception of ILS and Srow, did not differ when the

sex*population interaction was considered (Table 2).

The stepwise CDFA based on SIZE, shape, and

pholidosis showed good discrimination among the

three populations analysed. The ﬁrst canonical dis-

criminant function (CDF1) explained 66 and 76% of

the variation in males and females, respectively

(Table 3). The variables contributing most were TW

and HLL in males and TW, OD, and EL in females

(Table 3). Regarding CDF2 (34 and 24% of the male

and female variation, respectively), OD and SIZE in

336 R. VASCONCELOS ET AL.

Figure 3. Parsimony networks corresponding to the PDC, ACM4 and MC1R nDNA sequence variation in Tarentola from

the Cape Verde Islands. Lines represent a mutational step, circles haplotypes and dots missing haplotypes. The size of

circles is proportional to the number of haplotypes and colours to the number of individuals. The dotted circles represent

the most probable ancestral haplotype. Samples from the same island are similarly coloured but with different tonalities

for different taxa. For correspondences of sample and location codes see Appendix 1.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 337

males and TW, SLS, and OD in females were the most

important variables (Table 3). The graphical repre-

sentation of the factor scores across the two CDF axes

showed a good separation of the three populations

(Fig. 4). These results are conﬁrmed by the classiﬁ-

cation scores obtained, with 92.0% males and 90.5%

females correctly assigned to their populations

(Table 4). In males, the population from S. Nicolau

was the best discriminated (94.7% of the individuals

correctly classiﬁed), while the species from Fogo

(Table 4) presented the lowest classiﬁcation score,

although the values were still high (87.5%). Regard-

ing females, the population from S. Nicolau had the

highest score (100%), while T. darwini from Santiago

had the lowest (80.0%, Table 4).

INTEGRATIVE APPROACH

The IPC protocol recognizes the existence of 12

species within the Cape Verdean Tarentola (Fig. 2).

The distinctiveness of two species is supported by all

lines of evidence, eight species are supported by two,

and two are exceptionally supported by one (see Dis-

cussion for details). Also, two subspecies supported by

a single line of evidence are recognized for two out of

the 12 species. Based on these results, a new tax-

onomy for the genus Tarentola from Cape Verde is

proposed below. The different taxa are described fol-

lowing the order of the phylogenetic tree presented in

Figure 2 (from top to bottom).

ORDER SQUAMATA

FAMILY PHYLLODACTYLIDAE

TYPE GENUS TARENTOLA GRAY, 1825

TARENTOLA BOAVISTENSIS STAT.NOV.JOGER, 1993

(FIGS 1, 2A1, 3, 5A1, 6A1, 7A1)

MORPHOBANK M42539–M42659

Tarentola rudis boavistensis Joger, 1993: 438 (holo-

type: RMNH 24144, Boavista, Ilhéu Sal Rei, southern

part, paratypes: HLMD RA-1470, RMNH 24142,

24145-137, same locality and BMNH 1906.3.30.26,

Boa Vista, unknown locality.); Schleich, 1996: 125;

Andreone, 2000: 21, 25; Carranza et al., 2000: 641;

Köhler & Güsten, 2007: 279

Table 2. Summary of the ANOVA/MANOVA results regarding the effect of sex, population and their interaction

(sex*population) on the morphological variables using two different sets: raw variables (after log-transformation), and

size-corrected variables (using an isometric approach; SIZE)

Variable

Raw variables Size-corrected variables

Sex Population Sex*population Sex Population Sex*population

SIZE 16.06 ** 8.49 ** 5.78 **

SVL 12.14 ** 2.56 NS 5.68 ** 0.00 NS 32.54 ** 0.41 NS

TrL 5.73 * 9.40 ** 4.74 ** 1.11 NS 16.83 ** 0.23 NS

TW 25.73 ** 10.76 ** 3.65 * 18.34 ** 58.31 ** 0.16 NS

FLL 15.57 ** 19.94 ** 2.37 NS 0.27 NS 11.25 ** 2.70 NS

CFL 5.91 * 11.37 ** 2.84 NS 1.86 NS 7.06 ** 1.36 NS

HLL 12.77 ** 31.09 ** 5.39 ** 0.15 NS 22.43 ** 1.15 NS

FFL 10.77 ** 22.22 ** 6.19 ** 0.52 NS 15.24 ** 1.87 NS

HW 12.76 ** 2.30 NS 4.84 ** 0.61 NS 10.92 ** 0.21 NS

HH 15.77 ** 0.95 NS 6.11 ** 3.16 NS 11.02 ** 1.32 NS

OD 1.50 NS 23.05 ** 5.21 * 5.39 * 39.58 ** 4.11 *

EL 9.02 ** 20.64 ** 1.80 NS 1.60 NS 30.08 ** 1.95 NS

NED 9.34 ** 14.16 ** 5.12 ** 0.27 NS 5.28 * 1.26 NS

SED 9.20 ** 18.32 ** 4.25 * 0.24 NS 14.48 ** 0.18 NS

EED 10.36 ** 3.40 * 4.27 * 0.05 NS 5.80 ** 0.07 NS

SLS 0.45 NS 11.66 ** 1.58 NS

ILS 1.75 NS 6.06 ** 5.22 *

Lam 0.81 NS 8.84 ** 0.54 NS

Trow 4.26 * 2.78 NS 0.07 NS

Tline 1.92 NS 12.13 ** 0.38 NS

Srow 0.05 NS 2.96 NS 3.33 *

medS 0.28 NS 14.99 ** 0.05 NS

For each variable, F-values and level of signiﬁcances are provided (NS, not signiﬁcant, P>0.05; *, 0.01 <P<0.05; **,

P<0.01). See Material and methods for more details.

338 R. VASCONCELOS ET AL.

Tarentola delalandii delalandii: Boulenger, 1906:

200 (part.); Loveridge, 1947: 334 (part.); Schleich,

1982a: 246 (part.)

Tarentola delalandii: Angel, 1937: 1695 (part.)

Tarentola delalandii rudis: Mertens, 1954: 7 (part.)

Tarentola borneensis maioensis: Joger, 1984b: 102

(part.)

Tarentola maioensis: López-Jurado, Mateo &

Geniez, 1999: 11 (part.); López-Jurado, Mateo &

Fazeres, 2005: 101 (part.)

Tarentola maioensis boavistensis: López-Jurado

et al., 2005: 101

Specimens examined: 11 live specimens and six

voucher specimens (Appendix 1).

Additional material and references: Andreone (2000:

21, 25) refers to MSNG 49996, II.1898, and MSNG

37560. I.1898 (one and eight individuals, respectively,

all from Boavista, unknown locality, collected by

Fea); Carranza et al. (2000: 641) to BMNH 1998.344

(Boavista, Curral Velho), BMNH 1998.342, BMNH

1998.343 (Boavista, Vila de Sal Rei), and Köhler

& Güsten (2007: 279) refer to HLMD-RA-1470

(Boavista, Ilhéu Sal Rei).

Diagnosis: Medium to large-sized gecko (maximum

SVL 79.0 mm, 65.2 mm on average; cf. Appendix 2);

eye/ear opening ratio averages 1.59; ear–eye/eye–

snout distance ratio averages 0.83. Eight to 11

supralabials; seven to nine infralabials; nine to ten

enlarged lamellae under the 4th ﬁnger; 112–143

midbody scales (Joger, 1993); narrow central keeled

dorsal tubercles (Fig. 5A1) with 20–24 midbody lon-

gitudinal lines and 14–18 transverse rows; prominent

tubercle above and anterior to the ear opening. Light

orangey or yellowish to pinkish grey dorsal coloration

slightly translucent with reduced pattern in adults

(Figs 6A1, 7A1) and whitish below. A light vertebral

stripe, interrupted or complete, appears on most indi-

viduals. Eye iris generally orange to orangey brown,

contrasting with the rest of the head coloration. Juve-

niles with black tails with strongly marked white

stripes. Most specimens with thin brown streaks

arranged in different angles in front of and behind the

ear. First supra- and infralabials white followed by

labials with very dark spots.

It differs from other taxa from clade A by presenting

keeled dorsal tubercles and having an orangey, yel-

lowish to pinkish grey dorsal coloration that is

slightly translucent, and an orangey eye iris. It differs

from T. caboverdiana, clade B, and clade C by having

a light, reduced dorsal pattern. It differs from T. gigas

by having smaller SVL and from T. ‘rudis’ from San-

tiago, Fogo, Brava, Rombos, and Maio (taxa from

clade D), by having a lower midbody scale count

(112–143) but a higher number of interorbital scales

(19–22) (Joger, 1993).

Genetic and phylogeographical remarks: Tarentola

boavistensis is monophyletic and phylogenetically not

related to T. ‘rudis’ as it branches in a completely

different clade (Fig. 2). It also shows a high level of

genetic divergence compared with its sister taxa from

clade A: A1–A2, A1–A3 and A1–A4 p-dist (cyt

b)=9.0 ± 1.5, 9.8 ± 1.6, and 10.7 ± 1.6%, respectively

(Table 5). The Snn test values for PDC, ACM4, and

MC1R performed with its sister taxa are all signiﬁ-

cant (Appendix 5). According to the presently selected

Table 3. Summary of the stepwise Canonical Discrimi-

nant Function Analysis (CDFA) for the size/shape data

set (obtained after using an isometric approach) and

pholidosis

Variable

Males Females

CDF1 CDF2 CDF1 CDF2

SIZE 0.190 0.374 0.196 0.125

TrL -0.118 -0.291 0.078 -0.199

TW -0.511 -0.074 0.468 -0.445

FLL 0.275 -0.036 -0.379 0.290

CFL 0.036 -0.050 -0.041 0.469

HLL 0.387 0.293 -0.075 0.028

FFL 0.367 0.182 -0.057 0.041

HW -0.102 -0.144 0.057 -0.199

HH 0.107 -0.131 0.043 -0.286

OD -0.137 0.511 0.442 0.362

EL -0.230 0.272 0.426 0.279

NED -0.247 -0.014 -0.220 0.143

SED 0.198 0.097 -0.454 0.331

EED -0.197 -0. 111 -0.069 -0.275

SLS -0.098 0.248 0.076 0.465

ILS 0.042 0.369 -0.295 0.298

Lam 0.190 0.196 0.039 0.142

Tline 0.246 -0.208 0.037 -0.319

Trow -0.298 -0.270 -0.284 0.024

medS -0.227 -0.004 0.016 0.189

Srow -0.287 0.005 0.007 0.237

Eigenvalues 4.35 2.22 5.97 1.87

% explained 66.30 33.70 76.10 23.90

% cumulative 66.30 100.0 76.10 100.0

For each analysis, the factor structure of the ﬁrst two

CDFs, eigenvalues, and total explained and cumulative

contribution (%) of each CDF to the total variation is also

given. Analyses were made separately for males and

females. Variables contributing the most (>0.35%) are

indicated in bold. Scores in italics indicate the variables

that were not selected by the stepwise CDFA. See text for

more details regarding the meaning of the abbreviations of

the variables.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 339

protocol of integration (IPC), all lines of evidence

clearly support the differentiation of the endemic

Tarentola from Boavista from other taxa from clade A

and from all the other Tarentola from Cape Verde (see

Figs 2–4 and Appendix 3). Consequently, this taxon is

upgraded to the species level.

Distribution: Boavista Island and Sal Rei Islet, Cape

Verde.

Conservation status: Listed as Data Deﬁcient

under the criteria of the First Red List of Cape Verde

(Schleich, 1996).

TARENTOLA BOCAGEI SP.NOV.

(FIGS 1, 2A2, 3, 4, 5A2, 6A2, 7A2)

MORPHOBANK M43461–M43781, M55889-M55901

Tarentola darwini Joger, 1984b: 96 (part.), 1993: 443

(part.); Schleich, 1996: 125 (part.); Carranza et al.,

2000: 641 (part.); López-Jurado et al., 2005: 101

(part.); Köhler et al., 2007: 76 (part.)

Holotype: MNHH 2011.0201, male from S. Nicolau

Island (Cape Verde), Carriçal oasis, in the eastern

part of the island (16.555289′N, 24.082165′W,

WGS84), collected on 3 October 2009 by Vasconcelos,

Figure 4. Discriminant analyses for males and females previously called T. darwini. The total contribution of each of the

two Canonical Discriminant Functions (CDF1 and CDF2) to explain the total morphological variation is also given. See

Material and methods for details.

Table 4. Classiﬁcation matrix retrieved from the canonical discriminant analyses (CDFA)

Taxa Correct classiﬁcation T. bocagei T. fogoensis T. darwini

Island Sex (% and N) S. Nicolau Fogo Santiago

T. bocagei Males 94.7 (19) 94.7 (18) 0 (0) 5.3 (1)

S. Nicolau Females 100.0 (14) 100.0 (14) 0 (0) 0 (0)

T. fogoensis Males 87.5 (16) 6.3 (1) 87.5 (14) 6.3 (1)

Fogo Females 92.3 (13) 0 (0) 92.3 (12) 7.7 (1)

T. darwini Males 93.3 (15) 0 (0) 6.7 (1) 93.3 (14)

Santiago Females 80.0 (15) 0 (0) 20.0 (2) 80.0 (13)

Total Males 92.0 (50) 38.0 (19) 30.0 (15) 32.0 (16)

Females 90.5 (42) 33.3 (14) 33.3 (14) 33.3 (14)

For each population the percentage (%) and frequency (N; in parentheses) of correctly classiﬁed individuals are provided.

340 R. VASCONCELOS ET AL.

Perera, and Harris (MorphoBank M43478-M43487).

Paratypes: MNHN 2011.0202 female, same data as

for holotype (MorphoBank M55889-M55893); BMNH

1998.346 juvenile, S. Nicolau, Juncalinho (Mor-

phoBank M55894-M55895).

Specimens examined: 40 live specimens and four

voucher specimens (Appendix 1).

Additional material and references: Joger (1984b: 96)

refers to ZSM 138/1981 (three individuals, two doubt-

ful, S. Nicolau, unknown locality).

Diagnosis: Medium-sized gecko (maximum SVL

65.5 mm, 58.2 mm on average; Table 1); eye/ear

opening ratio averages 1.37; ear–eye/eye–snout dis-

tance ratio averages 0.80. Ten to 13 supralabials;

eight to ten infralabials; eight to ten enlarged

lamellae under the 4th ﬁnger; 122–146 midbody

scales (Joger, 1984b); slightly keeled rounded dorsal

tubercles (Fig. 5A2) with 17–24 midbody longitu-

dinal lines and 14–18 transverse rows; no enlarged

tubercles between the eye and the ear opening.

Dorsal parts grey or greyish with four to six trans-

verse bands generally asymmetrical and frequently

Y-shaped on the ﬂanks (Figs 6A2, 7A2), most of

the dorsal tubercles darker than the ground colour

while several other tubercles white, especially in

subadults and young specimens, well-deﬁned ver-

tebral line without tubercles; pileus almost

uniform contrasting with densely marked dorsum,

two longitudinal light bands from snout to eye; labials

and sides of the throat uniformly whitish or yellow-

ish, without dark stains; eye iris blackish or dark

brown.

It is characterized by the same general features

as T. darwini (not presenting enlarged tubercles

between the eye and ear opening and not strongly

keeled dorsal tubercles), but in comparison with

taxa from clade A3 and A4 by having, relative to

SVL, a shorter trunk length (22.2 mm on average;

Table 1), larger ear opening, base of the tail propor-

tionally wider, distance between nostrils or snout

tip and eye signiﬁcantly shorter, higher average

number of small scales between dorsal tubercles (2.1

versus 1.6 for both A3 and A4 lineages; Table 1),

proﬁle of the forehead more concave, ventral part

Figure 5. Magniﬁed dorsal tubercles of Tarentola species of the Cape Verde Islands.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 341

more yellowish, and subdigital lamellae more grey

than taxa from clade A3 and A4. In comparison with

the strongly resembling Tarentola from clade C

living on the same island, dorsal tubercles are less

keeled and more rounded (Fig. 5), white tubercles

are less numerous, not transversally aligned and

usually placed on the ﬂanks, and iris more uni-

formly dark.

Etymology: The species epithet is a genitive Latin

noun to honour J. V. Barbosa du Bocage, one of the

ﬁrst naturalists to study the reptiles from the Cape

Verde Islands.

Genetic and phylogeographic remarks: Tarentola

bocagei is monophyletic (Fig. 2) in the mitochon-

drial phylogeny and presents a high level of genetic

Figure 6. Typical dorsal patterns of Tarentola species of the Cape Verde Islands (adapted from Joger, 1993).

342 R. VASCONCELOS ET AL.

divergence when compared with its sister taxa:

A2–A1, A2–A3, A2–A4 p-dist (cyt b)=9.0 ± 1.5,

9.0 ± 1.5, and 10.1 ± 1.6%, respectively (Table 5). The

Snn test values for PDC, ACM4, and MC1R between

its sister taxa are all signiﬁcant (Appendix 5). Accord-

ing to the presently selected protocol of integration

(IPC), a minimum of two lines of evidence clearly

support the differentiation of T. bocagei from other

taxa from clade A and from all the other Tarentola

from Cape Verde (see Figs 2–4 and Appendix 3). Con-

sequently, this taxon is considered a distinct species.

Distribution: Eastern part of S. Nicolau Island, Cape

Verde.

Figure 7. Photographs of the dorsal and lateral views of Tarentola of the Cape Verde Islands. A1, T. boavistensis; A2,

T. bocagei; A3, T. fogoensis; A4, T. darwini; B1, T. substituta; B2, T. raziana; B3, T. caboverdiana;C,T. nicolauensis; D1,

T. gigas (T. gigas brancoensis on the left and T. gigas gigas on the right); D2, T. rudis; D3, T. protogigas protogigas; D4,

T. p. hartogi from Brava Island; D5, T. p. hartogi from Rombos Islets; D6, T. maioensis.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 343

Description of the holotype: A male gecko having the

following morphometric features: SVL 63 mm, head

19.8 long, 13.7 mm wide, 8.29 mm high from occiput

to jaws, distance between anterior eye and snout tip

5.1 mm, distance between anterior ear and posterior

eye 6.13 mm, nostril–eye distance 16.69 mm, greatest

orbital diameter 4.02 mm, longest dimension of ear

2.89 mm, total forelimb length 19.87 mm, crus fore-

limb length from base of palm to elbow 12.23 mm,

hindlimb length 25.57 mm, crus length from base of

heel to knee 14.49 mm, partially regenerated tail

55.5 mm long (tip of the tail cut for DNA analyses)

and 7.69 mm wide at widest point. Dorsal tubercles

slightly longer than wider, with one longitudinal

smooth but well-deﬁned keel, a straight vertebral line

without tubercles of about three small scales wide, 14

longitudinal rows of dorsal tubercles at midbody, 19

tubercles along the vertebral line, these tubercles

separated on average by 2.25 small scales, 11 suprala-

bials on the left side, ten supralabials on the right

side, eight infralabials on the left and right side, 44

gular scales counted from a line between the anterior

margins of the ear openings to the mental scale, nine

enlarged lamellae under the fourth ﬁngers, nine

enlarged lamellae under the fourth toes, 22 interor-

bital scales, nostrils in contact with rostral, the ﬁrst

supralabial and the three nasals, nasal scales sepa-

rated by one scale, six tubercles on each verticillum.

Colour in live specimen: mid-grey on the dorsum with

four dark transverse bands, the third and fourth

indistinct Y-shaped on the ﬂanks, pileus with indis-

tinct darker marks on the back, iris eyes blackish,

scales bordering the anterior part of the eye light

yellow, two longitudinal dark-faded stripes from snout

to eye and one from snout to superior part of the ear

opening enclosing a lighter stripe on each side; the six

ﬁrsts supralabials yellowish, the four posterior ones

whitish; lighter not well-marked vertebral line, most

of the dorsal tubercles darker than ground colour,

except 47 whitish ones, all dorsal tubercles and small

scales dark dotted; upper part of the tail with three

whitish transverse bands with lighter grey marks;

ventral parts white-yellowish becoming yellow on the

back; subdigital lamellae greyish. GenBank accession

code JN185934.

Conservation status: Listed as Data Deﬁcient under

the criteria of the First Red List of Cape Verde (Schle-

ich, 1996).

TARENTOLA FOGOENSIS SP.NOV.

(FIGS 1, 2A3, 3, 4, 5A3, 6A3, 7A3)

MORPHOBANK M42945–M43220, M55902-M55919

Tarentola delalandii var. boettgeri: Boulenger, 1906:

200;

Table 5. Estimates of evolutionary divergence over cyt bsequence pairs between groups

Clade A1 A2 A3 A4 B1 B2 B3 C D1 D2 D3 D4 D5 D6

Clade Taxa Tv Tb Tf Td Ts Tz Tc Tn Tg Tr Tpp Tph Tph Tm

A1 Tv 1.5 1.6 1.6 1.4 1.4 1.5 1.6 1.6 1.6 1.6 1.6 1.7 1.7

A2 Tb 9.0 1.5 1.6 1.5 1.5 1.5 1.5 1.6 1.6 1.5 1.5 1.6 1.6

A3 Tf 9.8 9.0 1.4 1.4 1.4 1.4 1.6 1.6 1.6 1.6 1.5 1.6 1.8

A4 Td 10.7 10.1 8.1 1.6 1.6 1.5 1.6 1.6 1.7 1.7 1.6 1.7 1.8

B1 Ts 8.0 8.5 7.0 9.9 0.4 0.7 1.4 1.2 1.2 1.2 1.1 1.1 1.5

B2 Tz 7.6 8.5 6.6 9.7 0.9 0.8 1.4 1.2 1.2 1.2 1.1 1.1 1.5

B3 Tc 9.0 9.0 7.5 9.4 2.2 2.8 1.4 1.2 1.3 1.2 1.2 1.2 1.5

CTn 10.3 9.7 9.3 10.4 7.1 7.1 7.2 1.3 1.5 1.4 1.4 1.4 1.5

D1 Tg 9.1 9.2 9.1 10.7 4.6 4.6 5.8 6.4 0.8 0.9 0.9 0.9 1.0

D2 Tr 9.5 9.7 9.3 11.3 5.4 5.4 6.6 7.8 2.4 1.0 0.9 0.9 1.2

D3 Tpp 9.0 8.7 8.7 11.4 4.8 4.8 5.9 6.8 2.8 3.1 0.8 0.8 1.3

D4 Tph 9.1 8.8 8.1 10.3 4.2 4.3 5.3 6.5 2.6 2.6 2.1 0.3 1.2

D5 Tph 9.3 9.0 8.3 10.6 4.5 4.5 5.5 6.8 2.8 2.9 2.3 0.4 1.3

D6 Tm 10.9 10.2 11.9 13.1 7.4 7.5 8.0 8.7 3.9 5.3 5.7 5.3 5.6

The number of base differences per site from averaging over all sequence pairs between groups is shown (p-dist). All

results are based on the pairwise analysis of 459 sequences. Standard error estimates are shown in italic and were

obtained by a bootstrap procedure (1000 replicates). Analyses were conducted in MEGA4. All positions containing gaps

and missing data were eliminated from the dataset. There were a total of 302 positions in the ﬁnal dataset.

Tv, T. boavistensis; Tb, T. bocagei; Tf, T. fogoensis; Td, T. darwini; Ts, T. substituta; Tz, T. raziana; Tc, T. caboverdiana; Tn,

T. nicolauensis; Tg, T. gigas; Tr, T. rudis; Tpp, T. protogigas protogigas; Tph, T. protogigas hartogi; Tm, T. maioensis.

344 R. VASCONCELOS ET AL.

Tarentola darwini: Joger, 1984b: 96 (part.); Joger,

1993: 443 (part.); Schleich, 1987: 40 (part.); Schleich,

1996: 125 (part.); Carranza et al., 2000: 641 (part.);

Carranza et al., 2002: 247 (part.); Jesus et al., 2002:

49 (part.); López-Jurado et al., 2005: 101 (part.).

Holotype: MNHN 2011.0203, male from Fogo Island

(Cape Verde), Ilhéu de Contenda (14.983′N, 24.438′W,

WGS84), collected on 7 December 1999 by S.

Carranza (MorphoBank M55902-M55907). First

paratype: MNHN 2011.0204, female, same data as for

holotype (MorphoBank M55908-M55911). Second

paratype: BEV.11072, female, same data as for holo-

type (MorphoBank M55912-M55919).

Specimens examined: 31 live specimens and nine

voucher specimens (Appendix 1).

Additional material and references: Joger (1984b: 96;

1993: 443) refers to SMF 50015, 50016, BMNH

1906.3.30.27 (all from Fogo, Igreja) and HLMW 3280

(Fogo, S. Filipe), respectively; Carranza et al. (2000:

641, 2002: 247) to BMNH 1998.356 (Fogo, Ribeira

Ilhéu) and BMNH 1998.354 (Fogo, S. Filipe); and

Jesus et al. (2002: 49) to CCBG T23894 (Fogo, S.

Filipe).

Diagnosis: Medium-sized gecko (maximum SVL

69.5 mm, 59.0 mm on average; Table 1); eye/ear

opening ratio averages 1.46; ear–eye/eye–snout dis-

tance ratio averages 0.73. Ten to 12 supralabials;

eight to 11 infralabials; nine to 11 enlarged lamellae

under the 4th ﬁnger; 137–148 midbody scales (Joger,

1984b); small numerous smooth, rounded dorsal

tubercles (Fig. 5A3) with 20–27 midbody longitudinal

lines and 14–18 transverse rows; absence of enlarged

tubercles between the eye and the ear opening. Dorsal

parts grey or greyish and generally without a distinct

vertebral stripe, with usually ﬁve transverse bands,

indistinct or not, and sometimes Y-shaped on the

ﬂanks and sometimes forming one X-shape on the

midbody (Figs 6A3, 7A3); two longitudinal faded light

bands from snout to eye; ventral parts whitish or

slightly yellowish; labials and sides of the throat with

generally numerous dark stains; eye iris blackish and

slightly golden on the upperparts.

It is characterized by the same general features

of T. darwini (not presenting enlarged tubercles

between the eye and ear opening and not strongly

keeled dorsal tubercles), but differs from T. darwini

from Santiago and T. bocagei by having, relative to

SVL, a narrower tail, limbs considerably longer, dis-

tance between nostrils and eye proportionally longer,

proﬁle of the forehead not concave. Certain individu-

als present a dark ring mark at the back, not

observed in any other Cape Verdean Tarentola; ver-

tebral line absent or less deﬁned than in T. bocagei;

pileus frequently vermiculate or marbled (more

uniform in T. bocagei), sometimes ventral parts

slightly yellowish but less than in T. bocagei and

subdigital lamellae whiter.

Etymology: The species epithet is an adjective that

refers to the island where the taxon is found, Fogo.

Genetic and phylogeographical remarks: T. fogoensis

is monophyletic (Fig. 2) and presents a high level of

genetic divergence when compared with T. boavisten-

sis,T. bocagei, and T. darwini from Santiago: A3–A1,

A3–A2 and A3–A4 p-dist (cyt b)=9.8 ± 1.6, 9.0 ± 1.5,

and 8.1 ± 1.4%, respectively (Table 5). The Snn test

values for PDC, ACM4, and MC1R are all signiﬁcant

among this clade (Appendix 5). According to the pres-

ently selected protocol of integration (IPC), a

minimum of two lines of evidence clearly support the

differentiation of T. fogoensis from other taxa from

clade A and from all other Tarentola from Cape Verde

(see Figs 2–4 and Appendix 3). Consequently, this

taxon is considered a distinct species.

Distribution: Fogo Island, Cape Verde.

Description of the holotype: A male gecko having the

following morphometric features: SVL in alcohol

61 mm, head 21.0 long, 14.6 mm wide, 8.9 mm high

from occiput to jaws, distance between anterior eye

and snout tip 7.9 mm, distance between anterior ear

and posterior eye 6.8 mm, greatest orbital diameter

3.6 mm, longest dimension of ear 1.9 mm, forelimb

length 17.8 mm, forelimb length from base of palm to

elbow 10.6 mm, hindlimb length 21.9 mm, crus length

from base of heel to knee 11.0 mm, tail regenerated

(tip of the tail cut for DNA analyses), 8.3 mm wide at

widest point. Dorsal tubercles slightly longer than

wider, smooth and not keeled, not distinct vertebral

line, 14 longitudinal rows of dorsal tubercles at

midbody, 26 tubercles along the vertebral line, these

tubercles separated on average by 1.5 small scales,

nine supralabials on the left side, eight supralabials

on the right side, six infralabials on the left and right

side, 42 gular scales counted from a line between the

anterior margins of the ear openings to the mental

scale, nine enlarged lamellae under the fourth

ﬁngers, ten enlarged lamellae under the fourth toes,

20 interorbital scales, nostrils in contact with rostral,

the ﬁrst supralabial and the three nasals, nasal scales

separated by one scale, four to six tubercles on each

verticillium. Colour in preserved specimen: mid-grey

on the dorsum with ﬁve dark asymmetric transverse

bands, the third and fourth fusing into a X-shape on

the vertebral region, pileus with transverse darker

mark on the nape of the neck enclosing a distinct

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 345

ring, scales bordering the anterior part of the eye

lighter, supralabials grey light, dorsal tubercles with

same colour as ground, all dorsal tubercles and small

scales dark dotted, original part of tail with two

darker marks at the base, regenerated part uniformly

grey; ventral parts dirty whitish.

Conservation status: Listed as Low Risk under the

criteria of the First Red List of Cape Verde (Schleich,

1996).

TARENTOLA DARWINI JOGER, 1984B(RESTRICTED

TYPE SPECIES)(FIGS 1, 2A4, 3, 4, 5A4, 6A4, 7A4)

MORPHOBANK M44231–M44984

Tarentola darwini Joger, 1984b: 96 (part.) (holotype:

ZFMK 37256 Santiago, Tarrafal; paratypes: ZFMK

37255, ZSM 365/78, 146/1981, 147/1981, ﬁve individu-

als, MHNP 35–187,188, all from Santiago, around

Tarrafal), 1993: 443 (part.); Schleich, 1987: 40 (part.),

1996: 124 (part.); Brygoo, 1990: 51; Carranza et al.,

2000: 641 (part.); Jesus et al., 2002: 49 (part.); López-

Jurado et al., 2005: 101 (part.); Frazen & Glaw, 2007:

219.

Tarentola delalandii delalandii: Boulenger, 1906:

200 (part.); Loveridge, 1947: 334 (part.)

Tarentola delalandii rudis: Mertens, 1954: 6 (part.)

Tarentola sp. Schleich, 1982a: 246, 1984: 102

Tarentola delalandii boettgeri: Schleich, 1984: 102

Specimens examined: 88 live specimens and 21

voucher specimens (Appendix 1).

Additional material and references: Schleich (1984:

102, 1987: 40) refers to ZFMK 37256 (Schleich collec-

tion 1978), ZSM 365/78, 146/81, 147/81, 29/8 (22 indi-

viduals, all from Santiago, Tarrafal); Joger (1993:

443) to HLMW 3209 (Santiago, S. Domingos); Brygoo

(1990: 51) to MHNP 1935.187, 1935.188, G 944,

ZFMK 37256 (Santiago, Pico Antónia, collected by

Chevalier); Carranza et al. (2000: 641) to BMNH

1998.348 (Santiago, Rui Vaz), BMNH 1998.350

(Santiago, Assomada), BMNH 1998.351 (Santiago,

Tarrafal); Jesus et al. (2002: 49) to CCBG T23895

(Santiago, Tarrafal) and Frazen & Glaw (2007: 219) to

ZSM 365/1978 (adult, Santiago, Tarrafal, collected by

H.-H. Schleich in 1977), ZSM 147/1981/1-5 (ﬁve indi-

viduals, Santiago, Tarrafal, H.-H., collected by Schle-

ich in 09.1981), ZSM 146/1981/1-2 (two adults,

Santiago, 5 km South from Tarrafal, collected by

H.-H. Schleich, 09.1981).

Diagnosis: Medium-sized gecko (maximum SVL

64 mm, 56.2 mm on average; Table 1); eye/ear

opening ratio averages 1.49; ear–eye/eye–snout

distance ratio averages 0.77. Nine to 12 supralabials;

seven to nine infralabials; eight to 11 enlarged lamel-

lae under the 4th ﬁnger; 113–130 midbody scales

(Joger, 1984b); small numerous smooth, rounded

dorsal tubercles (Fig. 5A4) with 17–27 midbody lon-

gitudinal lines and 13–18 transverse rows; no

enlarged tubercles between the eye and the ear

opening. Dorsal pattern generally composed of ‘silky’

silver-grey diffuse dark or light spots, sometimes con-

densed to form an irregular marbling (Figs 6A4, 7A4)

but sometimes forming indistinct transverse stripes,

especially in juveniles; vertebral stripe absent or

narrow and diffuse; light ventral parts; many dark

spots on supralabials and some sublabials lighter but

spotted; eye iris blackish with upperparts slightly

silver.

It differs from T. boavistensis, from clade A, and

taxa from clades B, C, and D by the diffuse dorsal

pattern instead of composed of three to ﬁve dark or

light symmetrical cross marks or pattern of bands.

Moreover, it also differs from T. boavistensis and from

taxa from clade D by not presenting enlarged

tubercles between the eye and ear opening or strongly

keeled dorsal tubercles. Instead it has smooth, ﬂat

oval to round tubercles with aligned cilia that produce

a ‘silky’ silver-grey dorsal aspect (Schleich, 1987). It

differs from T. bocagei and T. fogoensis by presenting,

relative to SVL, an intermediate tail width at its

widest point and snout–eye distance; orbital diameter

and the longest dimension of the ear smaller. It has

comparatively fewer supralabial scales.

Genetic and phylogeographical remarks: Tarentola

darwini is monophyletic (Fig. 2) and presents a high

level of genetic divergence when compared with its

sister taxa from clade A, T. boavistensis,T. fogoensis,

and T. bocagei: A4–A1, A4–A2, and A4–A3 p-dist (cyt

b)=10.7 ± 1.6, 10.1 ± 1.6, and 8.1 ± 1.4%, respec-

tively (Table 5). The Snn test values for PDC, ACM4,

and MC1R are all signiﬁcant among this clade

(Appendix 5). According to the presently selected

protocol of integration (IPC), all lines of evidence

clearly support the differentiation of T. darwini

from other taxa from clade A and from all other

Tarentola from Cape Verde (see Figs 2–4 and Appen-

dix 3). Consequently, this taxon is considered a

distinct species.

Distribution: Santiago Island, Cape Verde.

Conservation status: Listed as Indeterminate under

the criteria of the First Red List of Cape Verde

(Schleich, 1996).

346 R. VASCONCELOS ET AL.

TARENTOLA SUBSTITUTA STAT.NOV.JOGER, 1984B

(FIGS 1, 2B1, 3, 5B1, 6B1, 7B1)

MORPHOBANK M44991–M44994, M55646–M55698

Tarentola caboverdiana substituta Joger, 1984b: 103

(holotype: ZMH-R 0167, S. Vicente, unknown locality;

paratypes: ZMH-R 01686, ZMH-R 01688-89, ZMNH

1935.5.11.1-8, 1922.11.23.11, 1970.2424-25; all from

S. Vicente, unknown locality); Schleich, 1987: 46;

Joger, 1993: 438; Schleich, 1996: 124; Andreone, 2000:

21, 25; Carranza et al., 2000: 641; Carranza et al.,

2002: 247; Jesus et al., 2002: 49; López-Jurado et al.,

2005: 101; Köhler et al., 2007: 76.

Tarentola delalandii: Boulenger, 1885: 199 (part.);

Bocage, 1896: 4 (part.); Bocage, 1902: 209 (part.);

Angel, 1937: 1695 (part.)

Tarentola delalandii var. rudis: Loveridge, 1947:

334 (part.)

Tarentola delalandei delalandei: Dekeyser & Villi-

ers, 1951: 1152 (part.)

Tarentola delalandii rudis: Mertens, 1954: 6 (part.)

Tarentola delalandii delalandii: Schleich, 1982a:

246 (part.)

Tarentola caboverdianus caboverdianus: Schleich,

1984: 98 (part.)

Specimens examined: 26 live specimens and 10

voucher specimens (Appendix 1).

Additional material and references: Boulenger (1885:

199) refers to BMNH (ﬁve individuals collected by Rev.

Lowe, J. Macgillivray and Dr Cunningham, all from S.

Vicente, unknown locality); Dekeyser & Villiers (1951:

1152) to IFAN 50-1-104 to 50-1-107, IFAN 50-1-108 to

50-1-120 (all from S. Vicente, Baía das Gatas, and S.

Pedro, respectively, and collected by J. Cadenat in

1950); Mertens (1954: 6) to MUH 30.11.1953 (S.

Vicente, B. de Norte), MUH 10.1.1954 (S. Vicente,

Mindelo), MUH 26.11./2.12.1953, 9./11.3.1954 (S.

Vicente, Ribeira Julião); Schleich (1984: 98, 1987: 46)

to ZSM 371/78; 01-10.140/81 (S. Vicente, 3 km west

from Madeiral); Joger (1993: 438) to RMNH 24118-122

(S. Vicente, S. Pedro Bay), HLMW 3279 (S. Vicente,

airport); Andreone (2000: 21, 25) to MSNG 29221,

MSNG 36007 (seven and ﬁve individuals, respectively,

S. Vicente, Mindelo) and MZUT R2555, R3233 (S.

Vicente, unknown locality); Carranza et al. (2000: 641;

2002: 247) to BMNH 1998.364 (S. Vicente, Baía das

Gatas), and Jesus et al. (2002: 49) to CCBG T23891-

T23892 (S. Vicente, Madeiral).

Diagnosis: Medium-sized gecko (maximum SVL

65.5 mm, 51.6 mm on average), eye/ear opening ratio

between 1.5 and 2 (Schleich, 1987); ear–eye/eye–snout

distance ratio ⱕ1 (Schleich, 1987). Eight to 11

supralabials and seven to nine infralabials (Schleich,

1987); eight to nine enlarged lamellae under the 4th

ﬁnger; 146–167 midbody scales (Joger, 1984b); oval to

round conical and saddle-like more-or-less keeled

dorsal tubercles (Fig. 5B1) with 14–20 longitudinal

lines (Schleich, 1987); no tubercles between the eye

and the ear opening. Dorsal pattern with symmetrical

butterﬂy- or X-shaped dark dorsal cross bands often

lined with whitish tubercles posteriorly; vertebral

stripe absent or reduced to a narrow light line

(Figs 6B1, 7B1); cream to yellowish ventral parts;

generally white labials; blackish eye iris with golden

upperparts.

Smaller scales than the other Tarentola species

from clades B and C, and greater number of scales

around midbody (Joger, 1984b). It differs from Taren-

tola from clade B from Desertas (clade B2 in Fig. 2) by

its larger SVL and higher number of dorsal bands;

four to ﬁve from the neck to the caudal region some-

times surrounded by white tubercles (Joger, 1984b);

and from Tarentola from Santo Antão (clade B3 in

Fig. 2) by the head length being longer than the

anterior limbs and by presenting a higher number of

interorbital scales, usually 21 or more, and from

specimens from clade C by a lower number of scales

and lamellae under the ﬁfth toe (Joger, 1984b).

Distribution: S. Vicente Island, Cape Verde. Probably

introduced to Santo Antaˇo Island, Cape Verde (see

Vasconcelos et al., 2010).

Genetic and phylogeographic remarks: Tarentola sub-

stituta is monophyletic in the mtDNA tree from

Figure 2, although levels of support are low. Genetic

divergence among taxa within clade B is lower than

among members of clade A and D: B1–B2, B1–B3

and B2–B3 p-dist (cyt b)=0.9 ± 0.4, 2.2 ± 0.7, and

2.8 ± 0.8%, respectively (Table 5), but most of the Snn

test values for PDC, ACM4, and MC1R are signiﬁcant

within this clade (Appendix 5; see Discussion below).

According to the presently selected protocol of inte-

gration (IPC), a minimum of two lines of evidence

support the differentiation of the different island

populations and of the endemic Tarentola from S.

Vicente from all the other Tarentola from Cape Verde

(see Figs 2, 3 and Appendix 3). Consequently, this

taxon is upgraded to the species level.

Conservation status: Listed as Data Deﬁcient under

the criteria of the First Red List of Cape Verde (Schle-

ich, 1996).

TARENTOLA RAZIANA STAT.NOV.SCHLEICH, 1984

(FIGS 1, 2B2, 3, 5B2, 6B2, 7B2)

MORPHOBANK M44995–M44500, M55699–M55714

Tarentola caboverdianus razianus Schleich 1984: 101

(holotype: ZSM 01.133/81, Santa Luzia, unknown

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 347

locality; paratypes: 02-10.133/81, Santa Luzia,

unknown locality, ZSM 01-10.134/81, Raso Islet)

Tarentola delalandii rudis: Mertens 1954: 6 (part.);

Schleich, 1982a: 246 (part.)

Tarentola delalandii delalandii: Schleich, 1982a:

246 (part.)

Tarentola sp. Schleich & Wuttke, 1983: 34, 42

Tarentola caboverdiana raziana: Joger, 1984b: 104;

Schleich 1987: 44; Joger 1993: 438; Schleich 1996:

124; Andreone 2000: 21, 25; Carranza et al. 2000: 641;

Jesus et al. 2002: 49; López-Jurado et al. 2005: 101;

Frazen & Glaw 2007: 219.

Tarentola caboverdiana: Mateo et al. 1997: 8

Specimens examined: 19 live specimens and 21

voucher specimens (Appendix 1).

Additional material and references: Mertens (1954: 6)

refers to MUH 3.12.1953 (Santa Luzia, Água Doce);

Joger (1984b: 104, 1993: 438) to ZSM 01/133/81

(Santa Luzia, unknown locality) and RMNH 24110-

111 (Raso Islet), respectively; Schleich (1987: 44) to

ZSM 01.133/81, 02-10.133/81 (Santa Luzia, unknown

locality), ZSM 01-10.134/81 (Raso Islet); Andreone

(2000: 21, 25) to MSNG 49273 (two individuals, Raso

Islet); Carranza et al. (2000: 641) to BMNH 1998.362

(Santa Luzia); Jesus et al. (2002: 49) to IICT317*

(Raso Islet) and Frazen & Glaw (2007: 219) to ZSM

133/1981/1 (given as ZSM 01.133/81 in the original

description, male, Santa Luzia), ZSM 133/1981/2-10

(given as ZSM 02-10.133/81 in the original descrip-

tion, nine individuals, same data), ZSM 134/1981/1-9

(given as ZSM 01-10.134/81 in the original descrip-

tion, nine individuals, Raso Islet).

Diagnosis: Smallest Cape Verdean wall-gecko

(maximum SVL <60 mm, on average 49.3 mm; Appen-

dix 2), eye/ear opening ratio >2 (Schleich, 1987); ear–

eye/eye–snout distance ratio clearly ⱕ1 (Schleich,

1987). Nine to 11 supralabials (often ten) and seven to

nine infralabials (often eight or nine) (Schleich, 1987);

seven to ten enlarged lamellae under the 4th ﬁnger;

116–156 midbody scales (Joger, 1984b); oval to round

conical and saddle-like more-or-less keeled dorsal

tubercles (Fig. 5B2) with 16–18 longitudinal lines

(Schleich, 1987); no tubercles between the eye and the

ear opening. Snout particularly pointed and forehead

concave. Dorsal pattern with only three (sometimes

four) symmetrical butterﬂy- or X-shaped broad dark

dorsal crossbands often lined with whitish tubercles

posteriorly (Figs 6B2, 7B2); light grey or beige to dark

brown olive dorsal parts and cream to yellowish

ventral parts; generally white labials; eye iris dark

golden with a broad black horizontal band.

Besides its smaller size, it has narrower ﬁngers,

smaller number of lamellae under the ﬁrst toe, and

smaller number of gular scales than other Tarentola

from clade B (Joger, 1984b); usually only three dorsal

transverse bands (Joger, 1984b).

Distribution: Santa Luzia Island, Raso and Branco

Islet, Cape Verde.

Genetic and phylogeographical remarks: Tarentola

raziana is monophyletic in the mtDNA tree from

Figure 2, although levels of support are low. Genetic

divergence among taxa within clade B is lower than

among members of clades A and D: B1–B2, B1–B3,

and B2–B3 p-dist (cyt b)=0.9 ± 0.4, 2.2 ± 0.7, and

2.8 ± 0.8%, respectively (Table 5), but most of the Snn

test values for PDC, ACM4, and MC1R are signiﬁcant

among this clade (Appendix 5; see Discussion below).

According to the presently selected protocol of inte-

gration (IPC), a minimum of two lines of evidence

clearly support the differentiation of the Tarentola

populations from Sta. Luzia, Raso, and Branco from

all the other Tarentola from Cape Verde (see Figs 2, 3

and Appendix 3). Consequently, it is upgraded to the

species level.

Conservation status: Listed as Low Risk on the archi-

pelago under the criteria of the First Red List of Cape

Verde (Schleich, 1996). Considered as Low Risk on

Santa Luzia Island and as Rare on Raso Islet under

this same criteria (Schleich, 1996).

TARENTOLA CABOVERDIANA STAT.NOV.SCHLEICH,

1984 (FIGS 1, 2B3, 3, 5B3, 6B3, 7B3)

MORPHOBANK M44501–M44514, M55715–M55761

Tarentola caboverdianus caboverdianus Schleich,

1984: 98 (part.) (holotype: ZSM 03.141/81, male,

Santo Antão, unknown locality; paratypes: 01-02.141/

81, 04.141/81-17.141/81, all from Santo Antão,

unknown locality)

Tarentola delalandii: Bocage, 1896: 4 (part.), 1902:

209 (part.); Angel 1937: 1695 (part.)

Tarentola delalandii var. rudis: Loveridge 1947:

334 (part.)

Tarentola delalandei delalandei: Dekeyser & Villi-

ers 1951: 1152 (part.)

Tarentola delalandii rudis: Mertens 1954: 6 (part.)

Tarentola delalandii delalandii: Schleich, 1982a:

246 (part.)

Tarentola caboverdiana caboverdiana: Joger, 1984b:

102; Schleich 1987: 42; Joger 1993: 443; Schleich

1996: 124; Jesus et al. 2002: 49; López-Jurado et al.

2005: 101; Frazen & Glaw 2007: 219 (part.).

Specimens examined: 24 live specimens and eight

voucher specimens (Appendix 1).

348 R. VASCONCELOS ET AL.

Additional material and references: Bocage (1896: 4,

1902: 209) refers to individuals from GA (collected by

Dr Hopffer and lost during a ﬁre); Dekeyser & Villiers

(1951: 1152) to IFAN 50-1-87 to 50-1-93 and IFAN

50-1-94 to 50-1-103 (Santo Antão, unknown locality

and Porto Novo, respectively, all collected by J.

Cadenat in 1950); Mertens (1954: 6) to MUH 1.1.1954

and MUH 4./7.1.1954, 3.1.1954 (Santo Antão, Monte

Conceição and Porto Novo, respectively); Joger

(1984b: 102) to SMF 500011 (Santo Antão, Porto

Novo); Schleich (1987: 42) to ZSM 03.141/81,

01-02.141/81, 04.141/81-17.141/81 (30 individuals,

Santo Antão, 4–10 km north of Porto Novo – Chã de

Morte road, given as Chã do Monte); Jesus et al.

(2002: 49) to CCBG T23855, CCBG T23839 (Santo

Antão, Ponta do Sol and Porto Novo, respectively);

Frazen & Glaw (2007: 219) to ZSM 141/1981/3

(female, given as ZSM 03.141/81 in the original

description, Santo Antão, unknown locality), ZSM

141/1981/1-2, ZSM 141/1981/4-18 (17 individuals

given as ZSM 01-02.141/81 and ZSM 04-17.141/81

[sic] in the original, same data).

Diagnosis: Medium-sized gecko [maximum SVL

around 73.0 mm (Joger, 1993), 56.7 mm on average;

Appendix 2], eye/ear opening ratio between 1.5 and 2

(Schleich, 1987); ear–eye/eye–snout distance ratio ⱕ1

(Schleich, 1987). Nine to 13 supralabials and seven to

ten infralabials (Schleich, 1987); eight to ten enlarged

lamellae under the 4th ﬁnger; 116–150 midbody

scales (Joger, 1984b); oval to round conical and

saddle-like more-or-less keeled dorsal tubercles

(Fig. 5B3) with 14–16 (often 16) longitudinal lines

(Schleich, 1987); no tubercles between the eye and the

ear opening. Dorsal pattern with symmetrical

butterﬂy- or X-shaped dark dorsal crossbands often

lined with whitish tubercles posteriorly; vertebral

stripe frequently present, but narrow and indistinct

(Figs 6B3, 7B3); cream to yellow ventral parts; gen-

erally white labials; eye iris blackish.

It differs from other Tarentola from clades B and C

by its tail length, which is smaller than SVL (Schle-

ich, 1984). It differs from T. raziana by its larger SVL

and higher number of dorsal bands; from T. substituta

by its lower number of interorbital scales and by the

head length being comparatively shorter than hind-

limb length (Joger, 1984b). It differs from specimens

from clade C by having a lower number of lamellae

under the ﬁfth toe (Joger, 1984b).

Distribution: Santo Antão Island, Cape Verde.

Genetic and phylogeographic remarks: Tarentola

caboverdiana is monophyletic in the mtDNA tree

(Fig. 2). Genetic divergence among taxa within clade

B is lower than among members of clades A and D:

B1–B2, B1–B3, and B2–B3 p-dist (cyt b)=0.9 ± 0.4,

2.2 ± 0.7, and 2.8 ± 0.8%, respectively (Table 5), but

most of the Snn test values for PDC, ACM4, and

MC1R are signiﬁcant (Appendix 5) within this clade.

According to the presently selected protocol of inte-

gration (IPC), a minimum of two lines of evidence

clearly support the differentiation of the different

island populations and of the endemic Tarentola from

Santo Antão from all the other Tarentola from Cape

Verde (see Figs 2, 3 and Appendix 3). Consequently, it

is upgraded to the species level.

Conservation status: Listed as Low Risk under the

criteria of the First Red List of Cape Verde (Schleich,

1996).

TARENTOLA NICOLAUENSIS STAT.NOV.SCHLEICH,

1984 (FIGS 1, 2C, 3, 5C, 6C, 7C)

MORPHOBANK M45653–M45992

Tarentola caboverdianus nicolauensis Schleich, 1984:

100 (holotype: ZSM 02.138/81, S. Nicolau, unknown

locality; paratypes: ZSM 01 and 03-11.138/81, all from

S. Nicolau, unknown locality)

Tarentola delalandii: Bocage, 1902: 209 (part.);

Angel, 1937: 1695 (part.)

Tarentola delalandii delalandii: Boulenger 1906:

200 (part.); Schleich, 1982a: 246 (part.)

Tarentola delalandii var. rudis: Loveridge 1947:

334 (part.)

Tarentola delalandii rudis: Mertens 1954: 6, 7

(part.)

Tarentola caboverdiana nicolauensis: Joger, 1984b:

104; Schleich 1987: 43; Joger 1993: 443; Schleich

1996: 124; Andreone 2000: 21, 25; Carranza et al.

2000: 641; Jesus et al. 2002: 49; López-Jurado et al.

2005: 101; Frazen & Glaw 2007: 219; Köhler et al.

2007: 76.

Specimens examined: 39 live specimens and seven

voucher specimens (Appendix 1).

Additional material and references: Bocage (1902:

209) refers to specimens from GA (S. Nicolau, Vila da

Ribeira Brava, collected by F. Newton in 1901 and lost

during a ﬁre); Mertens (1954: 6, 7) to MUH 13./

17.12.1954, S. Nicolau, Chã de Preguiça); Joger

(1984b: 104) to ZSM 02.138/81 (S. Nicolau, unknown

locality); Schleich (1987: 43) to ZSM 02.138/81, ZSM

01 and 03-11.138/81 (all from S. Nicolau, unknown

locality); Andreone (2000: 21, 25) to MSNG 49998

(three individuals, S. Nicolau, unknown locality, col-

lected by Fea in 1898); Carranza et al. (2000: 641) to

BMNH 1998.358 (S. Nicolau, Tarrafal), BMNH

1998.359 (S. Nicolau, Tarrafal-Ribeira Brava); Jesus

et al. (2002: 49) to CCBG T23848 (S. Nicolau, Ponta

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 349

Cachorro), CCBG T23849*, CCBG T23847 (S.

Nicolau, Vila da Ribeira Brava) and CCBG T23850*

(S. Vicente, Mindelo) and Frazen & Glaw (2007: 219)

to ZSM 138/1981/2 (given as ZSM 02.138/81 in the

original description, male, S. Nicolau, collected by

H.-H. Schleich & H.-J. Gruber, 02.1981), ZSM 138/

1981/1, ZSM 138/1981/3-11 (ten individuals, same

data, given as ZSM 01.138/81 and ZSM 03-11.138/81

in the original description).

Diagnosis: Medium-sized gecko (maximum SVL

71.0 mm, 58.6 mm on average; Appendix 2); eye/ear

opening ratio averages 1.53; ear–eye/eye–snout dis-

tance ratio averages 0.79. Nine to 12 supralabials;

eight to ten infralabials; eight to 11 enlarged lamellae

under the 4th ﬁnger; 133–155 midbody scales (Joger,

1984b); oblong asymmetrical angled dorsal tubercles

(Fig. 5C) with 18–26 midbody longitudinal lines and

14–18 transverse rows (Schleich, 1987); no enlarged

tubercles between the eye and the ear opening. Dorsal

pattern greyish, presenting ﬁve clear symmetrical

butterﬂy- or X-shaped dark dorsal crossbands often

lined with whitish tubercles posteriorly (Figs 6C, 7C);

white to yellowish light ventral parts; uniformly

white labials; dark eye iris with golden upperparts.

It differs from T. boavistensis,T. bocagei,T. fogoen-

sis, and T. darwini, species from clade A, by butterﬂy-

or X-shaped dorsal pattern and from all the species

from clade D, and T. boavistensis by not having

enlarged tubercles between the eye and ear opening.

It differs from T. caboverdiana and T. substituta by a

higher number of lamellae under the fourth and ﬁfth

toe (Joger, 1984b) and also from T. bocagei by gener-

ally presenting oblong apical tubercles (Schleich,

1987; this study). Finally, it differs from T. raziana by

having a more massive head and a higher number of

crossbands (generally ﬁve) on the dorsum.

Distribution: West and central part of S. Nicolau

Island, Cape Verde.

Genetic and phylogeographical remarks: Tarentola

nicolauensis is monophyletic (Fig. 2) and shows a

high level of genetic divergence with species from

clade B, T. substituta,T. raziana, and T. caboverdi-

ana, within which it was included before the present

taxonomic revision: C–B1, C–B2 and C–B3 p-dist (cyt

b)ª7.1 ± 1.4%, and with T. bocagei, C–A2 p-dist (cyt

b)=9.7 ± 1.5% (Table 5). It presents signiﬁcant Snn

test values for MC1R and most comparisons of PDC

with species from clade B and only of PCD with

T. bocagei (Appendix 5). According to the presently

selected protocol of integration (IPC), a minimum of

two lines of evidence support the differentiation of

T. nicolauensis from species from clade B and from all

the other Tarentola from Cape Verde (Figs 2, 3 and

Appendix 3). Consequently, it is upgraded to the

species level.

Conservation status: Listed as Low Risk under the

criteria of the First Red List of Cape Verde (Schleich,

1996).

TARENTOLA GIGAS (BOCAGE, 1875)

Diagnosis: Giant gecko with SVL above 100 mm

[maximum SVL 155 mm (Bocage, 1896), 103.6 mm on

average; (Schleich, 1987)]; eye/ear opening ratio 1.5–

2.0 (Schleich, 1987); ear–eye/eye–snout distance ratio

slightly ⱕ1 (Schleich, 1984, 1987). Eight to 12

supralabials and seven to nine infralabials (Schleich,

1984); eight to 12 enlarged lamellae under the 4th

ﬁnger; 160–195 midbody scales (Joger, 1984b); ﬂatter

apical dorsal tubercles (Fig. 5D1) with 16 transverse

rows (Schleich, 1984); several enlarged tubercles

between the eye and the ear opening. Grey dorsal or

olive greyish pattern with a broad, light well-deﬁned

middorsal line with generally ﬁve large saddle-like

marks (Figs 6D1, 7D1); cream ventral parts, yellow

on the lower parts; big dark spots on the labials,

creating an alternating light and dark pattern; eye

iris dark grey with a typical vertical light area around

the pupil, joining the upper and lower parts of the eye

which are also light.

It differs from other Tarentola from the same clade

D, T. ‘rudis’ from Santiago, Fogo, Brava, Rombos, and

Maio, besides from its size, by the absence of a keel on

dorsal tubercles. Unlike all other Cape Verdean

Tarentola, strong vocalisations play a clear role in

social behaviour (Schleich, 1982b, 1987). This species

avoids vertical surfaces presumably due to its weight,

and presents a robust body with typical extreme fat

storage (Schleich, 1987).

Distribution: Raso and Branco Islets, Cape Verde.

Genetic and phylogeographic remarks: Tarentola

gigas is monophyletic in the mtDNA tree from

Figure 2. Genetic divergence with other taxa within

clade D is higher than among taxa within clade B,

although lower than among members of clade A:

D1–D2, D1–D3, D1–D4, D1–D5, and D1–D6 p-dist

(cyt b)=2.4 ± 0.8, 2.8 ± 0.9, 2.6 ± 0.9, 2.8 ± 0.9, and

3.9 ± 1.0%, respectively (Table 5). Most of the Snn

test values for PDC, ACM4, and MC1R are not sig-

niﬁcant among this clade (Appendix 5). According to

the presently selected protocol of integration (IPC), a

minimum of two lines of evidence differentiate

T. gigas from all the other Tarentola from Cape Verde

except T. protogigas from which it differs only in

morphology (Fig. 2). Consequently, it is considered a

350 R. VASCONCELOS ET AL.

different species, although not fulﬁlling the rule in

respect to T. protogigas, due to several ecological,

behavioural and geographical differences (see

Discussion).

The two subspecies, T. g. gigas and T. g. brancoen-

sis, are not reciprocally monophyletic (Fig. 2) and the

level of genetic divergence is very low, p-dist (cyt

b)=0.2 ± 0.2% (data not shown). Only one of the

three lines of evidence (morphology) differentiates the

two island populations. Consequently, according to

the IPC protocol, these are considered distinct sub-

species (Figs 2, 3 and Appendix 3).

TARENTOLA GIGAS GIGAS (BOCAGE, 1875)

(FIGS 1, 2D1, 3, 5D1, 6D1, 7D1)

MORPHOBANK M45993–M45995

Ascalabotes gigas Bocage, 1875: 108 (holotype: from

GA, collected by Dr Hopffer in 1874, Raso Islet and

lost due to a ﬁre; paratype: ZMB Nr. 8998, Raso Islet,

following Mertens, 1954)

Tarentola gigas: Boulenger, 1885: 200, 414 (part.);

Bocage, 1896: 4; Bocage, 1897: 194; Bocage, 1902: 4;

Boulenger, 1906: 200; Angel, 1937: 1695 (part.);

Mateo et al., 1997: 9, 11 (part.); Gamble et al., 2008: 3

(part.)

Tarentola delalandii gigas: Loveridge, 1947: 330

(part.); Mertens, 1954: 7; Greer, 1976: 702 (part.);

Schleich, 1980: 147 (part.); Gruber & Schleich, 1982:

309; Schleich, 1982b: 82 (part.); Schleich & Wuttke,

1983: 83

Tarentola ‘delalandii’gigas: Schleich, 1982a: 246

(part.)

Tarentola borneensis gigas: Joger, 1984b: 100, 1993:

440

Tarentola borneensis: Joger, 1985: 308 (part.)

Tarentola gigas gigas: Schleich, 1984: 104, 1987: 48,

1996: 124; Andreone, 2000: 21, 25; Carranza et al.,

2000: 641; López-Jurado et al., 2005: 101.

Specimens examined: Two live specimens and one

voucher specimen (Appendix 1).

Additional material and references: Bocage (1896: 4,

1897: 194, 1902: 4) refers to specimens from GA (Raso

Islet, collected by Dr Hopffer and Newton in 1874 and

lost due to a ﬁre); Gamble et al. (2008: 3) to JB 45

(unknown islet); Mertens (1954: 7) to co-type ZMB

8998 (Raso Islet); Schleich (1980: 147) to ZSMH 362/

1978 (unknown islet); Joger (1984b: 100, 1993: 440) to

ZMB Nr. 8998 and RMNH 24148-163, respectively

(Raso Islet); Schleich (1984: 104, 1987: 48) to ZSM

131/1981 (Raso Islet) and Andreone (2000: 21, 25) to

MSNG 22150 (one individual, Raso Islet, collected by

Fea in X-XI.1898) and MSNG 37517 (one individual,

same data).

Diagnosis: Giant gecko, SVL larger than 100 mm

[maximum SVL 155 mm (Bocage, 1896), 109.5 mm on

average (Schleich, 1987]. It differs from T. g. bran-

coensis by the ratio between the width and length of

the fourth toe being generally lower than 1:5, by

presenting a higher scale count around midbody

(180–213 versus 160–195) and a longer snout (Schle-

ich, 1984; Joger, 1984b).

Distribution: Raso Islet, Cape Verde.

Genetic and phylogeographic remarks: See T. gigas,

above.

Conservation status: Listed as Endangered and so in

need of urgent protection under the criteria of the

First Red List of Cape Verde (Schleich, 1996). The

Cape Verde authorities later considered the status of

this population as Endangered (Anonymous, 2002).

TARENTOLA GIGAS BRANCOENSIS SCHLEICH, 1984

(FIGS 1, 2D1, 3, 5D1, 6D1, 7D1)

Tarentola gigas brancoensis Schleich, 1984: 104 (holo-

type: ZSM 01.362/78, Branco Islet; paratypes: 02.-

06.362/78, 01.-12.19/82, same data), 1987: 49, 1996:

124; Carranza et al., 2000: 641, 2002: 247; López-

Jurado et al., 2005: 101; Frazen & Glaw, 2007: 220.

Tarentola borneensis: Gray, 1845: 165 (part.)

(Borneo ex errore pro Branco, following Joger, 1984b);

Joger, 1985: 307

Tarentola gigas: Angel, 1937: 1695 (part.); Mateo

et al., 1997: 9, 11 (part.)

Tarentola delalandii gigas: Loveridge, 1947: 330

(part.); Greer, 1976: 702 (part.); Schleich, 1980: 147

(part.); Schleich, 1982b: 82 (part.); Schleich & Wuttke,

1983: 83

Tarentola ‘delalandii’gigas: Schleich, 1982a: 246

(part.)

Tarentola borneensis gigas: Joger, 1984b: 100

Tarentola borneensis borneensis: Joger, 1993: 443

Specimens examined: Two voucher specimens

(Appendix 1).

Additional material and references: Schleich (1980:

147) refers to ZSMH 362/1978 (unknown islet); Joger

(1984b: 100) to BMNH 1946.8.25.79-80 (Branco islet);

Schleich (1987: 49) to ZSM 01.362/78; 02.-06.362/78,

01.-12.19/82 (Branco Islet); Frazen & Glaw (2007:

220) to ZSM 362/1978/1 (female, Branco Islet, given

as ZSM 01.362/78 in the original description), ZSM

362/1978/2-8 (ﬁve adults, two juveniles, same data,

given as ZSM 02.-06.362/78 in the original descrip-

tion), ZSM 19/1982/1-7 (seven individuals, same data,

given as ZSM 01.-12.19/82 in the original description).

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 351

Diagnosis: Giant gecko with SVL above 100 mm

[maximum SVL 113 mm, 98.0 mm on average (Schle-

ich, 1987)].

It differs from T. g. gigas by its smaller body mass,

by the ratio between the width and length of the

fourth toe being generally higher than 1:5, by pre-

senting a lower scale count around midbody (160–195

versus 180–213) and a shorter snout (Schleich, 1984;

Joger, 1984b).

Distribution: Branco Islet, Cape Verde.

Genetic and phylogeographic remarks: See T. gigas,

above.

Conservation status: Listed as Endangered and so in

need of urgent protection under the criteria of the

First Red List of Cape Verde (Schleich, 1996). The

Cape Verde authorities later considered the status of

this population as Endangered (Anonymous, 2002).

TARENTOLA RUDIS STAT NOV.BOULENGER, 1906

(FIGS 1, 2D2, 3, 5D2, 6D2, 7D2)

MORPHOBANK M45996–M46036

Tarentola delalandii var. rudis Boulenger, 1906: 200

(part.) [lectotype: MCNG 28149/1, Santiago, unknown

locality; later terra typica restricted to Santiago,

Calheta de S. Martinho (Joger, 1984b: 101)].

Tarentola borneensis rudis: Joger, 1984b: 101

Tarentola rudis rudis: Schleich, 1984: 97 (part.),

1987: 36; Joger, 1993: 443; Schleich, 1996: 124;

Andreone, 2000: 21, 25; Carranza et al., 2000: 641,

2002: 247; López-Jurado et al., 2005: 101

Specimens examined: 25 live specimens and ten

voucher specimens (Appendix 1).

Additional material and references: Joger (1984b:

101) refers to lectotype MCNG 28149/1 and paralec-

totype MCNG 28149/2 (Calheta de S. Martinho);

Schleich (1984: 97, 1987: 36) to ZSM 372/1978 (one

individual, Santiago, Cidade Velha), ZSM 135/1981

(two individuals, Santiago, Praia), ZSM 139/1981

(three individuals, Santiago, Praia airport) and also

to MCNG 28149/2 (Santiago, Calheta de S. Martinho),

respectively; Andreone (2000: 21, 25) to MSNG 28149,

MSNG 37561 (two syntypes and two individuals,

respectively, Santiago, Calheta de S. Martinho) and

MSNG 49997 (one individual, Santiago, Pedra

Badejo), all collected by Fea in 1898; Carranza et al.

(2000: 641, 2002: 247) to BMNH 1998.369 (Santa

Maria islet), DB-ULPGC-GG-12, BMNH 1998.368

(Santiago, Praia), also BMNH 1998.365 (Santiago,

Cidade Velha).

Diagnosis: Medium to large-sized gecko [maximum

SVL around 88 mm (Schleich, 1987), 68.7 mm on

average; Appendix 2]; eye/ear opening ratio averages

1.92; ear–eye/eye–snout distance ratio averages 0.78.

Nine to 11 supralabials (generally ten or 11) and

seven to 11 infralabials (Schleich, 1984); nine to 13

enlarged lamellae under the 4th ﬁnger; 130–165

midbody scales (Joger, 1984b); conical to apical promi-

nent dorsal tubercles with a narrow central keel

(Fig. 5D2), especially on the tail, with 16–22 longitu-

dinal lines and 12–18 transverse rows (Schleich,

1984, 1987 and Appendix S2); several enlarged

tubercles between the eye and the ear opening. Grey

brownish-greenish dorsal pattern with a series of four

to ﬁve (usually four) light middorsal patches, each

preceded by a W-shaped dark mark, usually con-

nected by a light middorsal line, which is situated in

a tubercle-free space (Figs 6D2, 7D2); white ventral

parts; clearly marked big dark spots on the labials,

creating an alternating light and dark pattern; eye

iris light grey with a broad horizontal dark area. Note

that the insular specimens from Ilhéu Santa Maria

are less robust and have the middorsal line generally

more pronounced.

It differs from T. bocagei,T. fogoensis,T. darwini,T.

substituta,T. raziana,T. caboverdiana, and T. nico-

lauensis by presenting enlarged tubercles between

the eye and ear opening and prominent dorsal

tubercles with a narrow central keel and by present-

ing a W-shaped dorsal pattern limiting a white spot,

instead of symmetrical or asymmetrical butterﬂy- or

X-shaped dark dorsal crossbands or marbled patterns

(Figs 6, 7). It differs from T. gigas by its smaller SVL

(always below 100 mm), its smaller mass, and eye iris

coloration. It differs from T. boavistensis by generally

presenting greyer dorsal coloration with frequently

more contrasted pattern and eye iris not orangey, and

from other taxa from clade D by the coloration and

pattern of the labials (darker and/or more regularly

creating an alternated dark and light pattern than

Tarentola from Fogo, Brava, Rombos, and Maio). It

also differs from Tarentola from clade D from Brava,

Rombos, and Maio by four to ﬁve well-deﬁned

W-shaped dorsal bands (Fig. 6); from Tarentola from

Fogo, Brava, and Rombos of the same clade by a

whiter ventral coloration, and from Tarentola from

Maio by a higher number of scales and lamellae

under the ﬁfth toe [22–24, rarely 21 versus 19–21,

rarely 22 (Joger, 1984b)].

Distribution: South of Santiago Island and Santa

Maria Islet, Cape Verde.

Genetic and phylogeographical remarks: Tarentola

rudis is monophyletic (Fig. 2) and genetically differ-

entiated from other taxa from clade D: D1–D2,

352 R. VASCONCELOS ET AL.

D2–D3, D2–D4, D2–D5 and D2–D6 p-dist (cyt

b)=2.4 ± 0.8, 3.1 ± 1.0, 2.6 ± 0.9, 2.9 ± 0.9, and

5.3 ± 1.2%, respectively (Table 5). However, the Snn

test values for PDC, ACM4, and MC1R are not sig-

niﬁcant between T. rudis and Tarentola from Maio

(Appendix 5). According to the presently selected pro-

tocol of integration (IPC), a minimum of two lines of

evidence support the differentiation with sister taxa

from clade D and differentiation of T. rudis from all

the other Tarentola from Cape Verde (Figs 2, 3 and

Appendix 3). Consequently, T. rudis is considered a

distinct species.

Conservation status: Listed as Indeterminate and in

need of urgent protection under the criteria of the

First Red List of Cape Verde (Schleich, 1996). The

Cape Verde authorities later considered the status of

this population as Indeterminate (Anonymous, 2002).

TARENTOLA PROTOGIGAS JOGER 1984B

Diagnosis: Medium to large-sized gecko [maximum

SVL 98.5 mm (Schleich, 1987); 71.9 mm on average,

see Appendix 2]; eye/ear opening ratio averages 1.69;

ear–eye/eye–snout distance ratio averages 0.75. Eight

to 12 supralabials; seven to nine infralabials; ten to

13 enlarged lamellae under the 4th ﬁnger; 144–181

midbody scales (Joger, 1984b); conical to apical promi-

nent dorsal tubercles with a narrow central keel

(Fig. 5D4), especially on the sacral region, with 12–15

transverse rows and 15–21 longitudinal rows; several

enlarged tubercles between the eye and the ear

opening. Grey, brownish to yellowish dorsal pattern

with a series of four (sometimes ﬁve) light middorsal

patches, each preceded by a more indistinct and

lighter W-shaped dark mark, usually connected by a

light middorsal line (Figs 6D3–5 and 7D3–5); golden-

yellowish grey ventral parts; dark spots on the

labials, sometimes creating an alternating light and

dark pattern; eye iris grey with an indistinct broad

horizontal dark area.

It differs from T. bocagei,T. fogoensis,T. darwini,T.

substituta,T. raziana,T. caboverdiana, and T. nico-

lauensis by having prominent conical dorsal

tubercles, enlarged tubercles between the eye and ear

opening and a different dorsal pattern (Fig. 6), and

from T. gigas by the presence of a narrow well-

marked central keel, especially on the sacral region.

It also differs from T. gigas by having important mor-

phological, bioacustical, ecological, and behavioural

differences. It differs from T. boavistensis,T. rudis,

and Tarentola from Maio by its yellower ventral col-

oration. It also differs from T. rudis by a higher

number of scales around midbody and interorbital

scales [18–21 versus 16–19 (Joger, 1984b)], by having

four to ﬁve more indistinct and lighter W-shaped

dorsal bands (Fig. 6), fader spots on the labials and

less contrasted eye iris coloration (Fig. 7). It differs

from Tarentola from Maio by a higher number of

scales and lamellae under the ﬁfth toe [22–26 versus

19–21, rarely 22 (Joger, 1984b)] and interorbital

scales [19–21 versus 16–18 (Joger, 1984b)].

Distribution: The southern islands of Fogo, Brava,

and Rombos Islets, Cape Verde.

Genetic and phylogeographical remarks: Tarentola

protogigas is monophyletic (Fig. 2) and presents a

considerable level of genetic divergence from other

sister taxa from clade D, as T. gigas,T. rudis, and

Tarentola from Maio: D3–D1, D3–D2, and D3–D6

p-dist (cyt b)=2.5 ± 1.2, 2.6 ± 0.9, and 5.3 ± 1.2%,

respectively (Table 5). The population from Fogo pre-

sents a considerable level of genetic divergence with

the populations from Brava and Rombos: D3–D4 and

D3–D5 p-dist (cyt b)=2.1 ± 0.8 and 2.3 ± 0.8%,

respectively. However, the Snn test values for PDC,

ACM4, and MC1R are not signiﬁcant between T. pro-

togigas from Fogo versus Brava and Rombos (Appen-

dix 5). The population from Brava presents very low

values of genetic divergence with the population from

Rombos: D4–D5 p-dist (cyt b)=0.4 ± 0.3%. Therefore,

only one of the three lines of evidence (morphology)

differentiates the population of Fogo from Brava and

Rombos. Consequently, according to the IPC protocol,

T. p. protogigas and T. p. hartogi comb. nov. are con-

sidered only distinct subspecies (Fig. 2). The lack of

differentiation in at least two of the three lines

of evidence precludes any further differentiation

between the island populations from Brava and

Rombos.

TARENTOLA PROTOGIGAS PROTOGIGAS JOGER,

1984B(RESTRICTED TYPE SUBSPECIES)

(FIGS 1, 2D3, 3, 6D3–D5, 7D3)

MORPHOBANK M46037–M46055

Tarentola borneensis protogigas Joger, 1984b: 100

(part.) (restricted holotype: ZSM 01/145/81, Fogo,

unknown locality; paratypes: ZSM 02/145/1981, Fogo,

unknown locality; BMNH 1906.3.30.28-29; MCNG

C.E. 28149, all from Fogo, S. Filipe;); Frazen & Glaw,

2007: 219

Tarentola delalandii var. rudis: Boulenger, 1906:

200 (part.); Loveridge, 1947: 332 (part.)

Tarentola delalandii rudis: Mertens, 1954: 6 (part.)

Tarentola ‘delalandii’rudis: Schleich, 1982a: 246

(part.)

Tarentola rudis rudis: Schleich, 1984: 97 (part.)

Tarentola rudis protogigas: Schleich, 1987: 38

(part.), 1996: 124 (part.); Joger, 1993: 439 (part.), 443;

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 353

Andreone, 2000: 21, 25 (part.); López-Jurado et al.,

2005: 101 (part.); Frazen & Glaw, 2007: 219

Specimens examined: Two live specimens

(Appendix 1).

Additional material and references: Mertens (1954: 6)

refers to MUH 16./21. 2. 1954 (ﬁve individuals, Fogo,

unknown locality); Schleich (1984: 97, 1987: 38) to

ZMS 145/1981.1-11 (Fogo, S. Filipe or S. Lourenço);

Andreone (2000: 21, 25) to MSNG 28148 (one indi-

vidual, Fogo, Igreja), MSNG 37516, MSNG 37515,

MSNG 49249, MSNG 49250 (one, two, two, and three

individuals, respectively, all from Fogo, S. Filipe) all

collected by Fea in 1898 and Frazen & Glaw (2007:

219) to ZSM 145/1981/1 and ZSM 145/1981/2 (Fogo, S.

Filipe, given as ZSM 01/145/81 and ZSM 02/145/1981

in the original, respectively, collected by H. H. Schle-

ich & H.-J. Gruber in 01.1981).

Diagnosis: Large-sized gecko [maximum SVL

98.5 mm (Schleich, 1987), 80.0 mm on average,

Appendix 2]. It differs from T. protogigas hartogi by

its longer SVL, its less yellowish and more marbled

ventrum, and more distinct W-shaped dorsal marks

(Fig. 6).

Distribution: Fogo Island, Cape Verde.

Genetic and phylogeographical remarks: See T. proto-

gigas above.

Conservation status: Considered Low Risk on Fogo

Island under the criteria of the First Red List of Cape

Verde (Schleich, 1996).

TARENTOLA PROTOGIGAS HARTOGI COMB.NOV.

JOGER, 1993 (FIGS 1, 2D4–D5, 3, 5D4,

6D3–D5, 7D4–D5)

MORPHOBANK M46056–M46091

Tarentola rudis hartogi Joger 1993: 439 (holotype:

RMNH 24131, collected on Cima Island, Rombos

group – central plateau, in sandy area under shrub of

Malvaceae – on 23/24 August, 1986 by J. C. Den

Hartog; paratypes: HLMD RA-1471, RMNH 24116,

Cima Island, easternmost tip, under rock; RMNH

24130, same locality as holotype, SMF 50012, Luiz

Carneiro Islet, Rombos group); Schleich 1996: 124;

Andreone 2000: 21, 25; Carranza et al. 2000: 641;

López-Jurado et al. 2005: 101 (part.); Köhler &

Güsten 2007: 279.

Tarentola delalandii delalandii: Boulenger 1906:

200; Schleich, 1982a: 246 (part.)

Tarentola delalandii: Angel 1937: 1695 (part.)

Tarentola delalandii var. rudis: Loveridge 1947:

332 (part.)

Tarentola delalandii rudis: Mertens 1954: 6 (part.)

Tarentola ‘delalandii’rudis: Schleich, 1982a: 246

(part.)

Tarentola delalandii ssp.: Schleich, 1982a: 246

(part.)

Tarentola borneensis protogigas: Joger, 1984b: 100

(part.)

Tarentola rudis protogigas: Schleich, 1987: 38

(part.), 1996: 124; Joger, 1993: 439, 443; Andreone,

2000: 21, 25; Carranza et al., 2000: 641; López-Jurado

et al., 2005: 101

Specimens examined: 27 live specimens and 15

voucher specimens (Appendix 1).

Additional material and references: Mertens (1954: 6)

refers to MUH 22./26.2.1954 (ﬁve individuals, Brava,

unknown locality) and MUH 27.2.1954 (ﬁve individu-

als, Rombos, Luiz Carneiro and Cima Islet); Joger

(1984b: 100) to SMF 50013-014, Brava, Ilhéu de Con-

tenda (14.983 N, 24.438 W, WGS84), collected in 1984

by Joger, and SMF 50012, Luiz Islet, Rombos; Andre-

one (2000: 21, 25) to MSNG 28147, MSNG 49994,

MSNG 49995 (three, six, and one individual, respec-

tively, all from Brava, unknown locality, collected by

Fea in 1899) and to MSNG 37514 (ﬁve individuals,

Rombos, unknown locality, collected by Fea in 1898);

Carranza et al. (2000: 641) to BMNH 1998.374

(Brava, Porto da Furna), BMNH 1998.376, BMNH

1998.377 (Brava, Porto Ancião) and to BMNH

1998.372, BMNH 1998.373 (Rombos, unknown local-

ity); Köhler & Güsten (2007: 279) to HLMD-RA-1471

(Rombos, Cima Islet, southernmost tip).

Diagnosis: Medium to large-sized gecko (maximum

SVL 77.0 mm; 63.8 mm on average, Appendix 2). It

differs from T. protogigas protogigas by a shorter SVL,

more yellowish ventral coloration, and less distinct

W-shaped dorsal marks on adults.

Distribution: Brava Islands and Rombos Islet group,

Cima and Luiz Carneiro Islets, Cape Verde.

Genetic and phylogeographic remarks: See T. protogi-

gas above.

Conservation status: Listed as Data Deﬁcient on

Brava and Rombos under the criteria of the First Red

List of Cape Verde (Schleich, 1996).

TARENTOLA MAIOENSIS STAT.NOV.SCHLEICH, 1984

(FIGS 1, 2D6, 3, 5D6, 6D6, 7D6)

MORPHOBANK M46092–M46109

Tarentola rudis maioensis Schleich, 1984: 98

(holotype: ZSM 06.136/81, Maio, unknown locality;

354 R. VASCONCELOS ET AL.

paratypes: ZSM 01.136/81-05.136/8, 07.136/8,

09.136/8; all from Maio, unknown locality), 1987: 37,

1996: 124; Joger, 1993: 438; Frazen & Glaw, 2007: 220

Tarentola delalandii rudis: Schleich, 1982a: 246

(part.); Mertens, 1954: 6 (part.)

Tarentola delalandii ssp.. Schleich, 1982a: 246

Tarentola borneensis protogigas: Joger, 1984b: 102

(part.)

Tarentola maioensis maioensis: López-Jurado et al.

2005: 101

Specimens examined: 16 live specimens and ﬁve

voucher specimens (Appendix 1).

Additional material and references: Mertens (1954: 6)

refers to MUH 3.2.1954 (two individuals, Maio,

unknown locality); Joger (1984b: 102) to ZSM

06/136/81 (Maio, unknown locality); Schleich (1987:

37) to ZSM 136/81.1-9 (Maio, stream between Vila do

Maio and Morro); Joger (1993: 438) to HLMW 3281

(two individuals, Maio, unknown locality) and RMNH

24112-113 (Maio, North of Vila do Maio) and Frazen

& Glaw (2007: 220) to ZSM 136/1981/6 (male, Maio,

unknown locality, given as ZSM 06.136/81 in the

original description), ZSM 136/1981/1-5 and ZSM 136/

1981/7-9 (eight individuals, same data, given as ZSM

01.136/81-05.136/81 and 07.136/81-09.136/81, respec-

tively in the original description).

Diagnosis: Medium-sized gecko (maximum SVL

71.0 mm, 60.8 mm on average, Appendix 2) with a

wide and long head (Schleich, 1984); distinct eye/ear

opening ratio ⱖ2; ear–eye/eye–snout distance ratio

averages ⱕ1. Seven to nine supralabials and seven to

nine infralabials (Schleich, 1984, 1987); eight to ten

enlarged lamellae under the 4th ﬁnger; 129–149

midbody scales (Joger, 1984b); conical to apical promi-

nent dorsal tubercles with a narrow central keel

(Fig. 5D6) with 12–18 (often 14) transverse rows

(Schleich, 1987); several enlarged tubercles between

the eye and the ear opening. Light grey-brownish

dorsal coloration; dorsal pattern with a series of four

to ﬁve, faint light middorsal patches and/or a broad

light middorsal line, each preceded by wide brown

marks (Figs 6D6 and 7D6); white ventral parts;

usually faint dark spots on the labials sometimes

alternating dark and light; pale grey eye iris with a

faded horizontal darker area.

It differs from T. bocagei,T. fogoensis, and T. dar-

wini,T. substituta,T. raziana,T. caboverdiana and

T. nicolauensis by having conical dorsal slightly apical

prominent tubercles (Fig. 5), enlarged tubercles

between the eye and ear opening, and a different

dorsal pattern (Figs 6, 7). It differs from T. gigas,

T. boavistensis,T. rudis, and T. protogigas by a lower

maximum size (71 versus 115, 79, 83 and 99 mm,

respectively). Moreover, it differs from T. boavistensis

by a greyer dorsal and eye iris coloration and from

T. rudis and T. protogigas by generally presenting

lower number of scales and lamellae under the ﬁfth

toe [19–21, rarely 22 versus 22–26 (Joger, 1984b)]. It

also differs from T. rudis by presenting lighter dorsal

coloration with wider and fainter dorsal bands and

generally fainter coloration on the labials (Figs 6, 7).

Finally, it also differs from T. protogigas by a lower

number of interorbital scales [16–18 versus 19–21

(Joger, 1984b)] and the whitish ventral coloration.

Distribution: Maio Island, Cape Verde. Recently intro-

duced to S. Nicolau Island, Cape Verde (see Vascon-

celos et al., 2010).

Genetic and phylogeographical remarks: Tarentola

maioensis is a monophyletic lineage, genetically dif-

ferentiated from other members of its clade, T. gigas,

T. rudis, and T. protogigas: D6–D1, D6–D2, D6–D3/

4/5 p-dist (cyt b)=3.9 ± 1.0, 5.3 ± 1.2, and

ⱖ5.3 ± 1.2%, respectively (Table 5). Also Snn test

values for MC1R were signiﬁcant between T. maioen-

sis and all other species of clade D, except T. rudis,

and all for PDC except with the latter and T. gigas

(Appendix 5). According to the presently selected pro-

tocol of integration (IPC), all lines of evidence clearly

support the differentiation of this taxon from all other

taxa of clade D (see Figs 2, 3 and Appendix 3), with

the only exception of T. rudis, from which it differs by

only two lines of evidence (mtDNA and morphology).

Consequently, the endemic Tarentola from Maio is

upgraded to the species level.

Conservation status: Listed as Low Risk under the

criteria of the First Red List of Cape Verde (Schleich,

1996).

DISCUSSION

The results of the molecular and morphological analy-

ses are in accordance with previous reports of mito-

chondrial and morphological variation (Joger, 1984b,

1993; Schleich, 1984, 1987; Carranza et al., 2000;

Vasconcelos et al., 2010). There is a remarkable

degree of concordance between the units deﬁned

based on previously published mtDNA data and those

observed by morphological analyses and multilocus

nuclear data. The only exception is between T. sub-

stituta and T. raziana, which present low levels of

mtDNA divergence but signiﬁcant morphological and

nuclear differentiation. For this reason a large

number of samples were sequenced for MC1R

(N=58), conﬁrming that the absence of haplotype

sharing between T. substituta and T. raziana was not

a consequence of stochasticity due to low sample size.

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 355

These two taxa may have been in partial contact and

introgressed during the Pleistocene sea-level falls.

The gene ﬂow occurred as a consequence of the con-

nection of the Desertas island group with S. Vicente

during that period, and has left a signature in the

population genetic structure of low mitochondrial

divergence between T. substituta and T. raziana that

would be misleading if systematics was based on a

single line of evidence. Once those populations were

isolated again, the backcrossing of outnumbered

hybrids led to clear nuclear differentiation of each

island population. The persistence of introgressed

mtDNA after several backcrosses is explained by the

continuous passage of this information through the

maternal lineage. Other studies on birds and Taren-

tola have also shown that mtDNA alone can be mis-

leading (Zink & Barrowclough, 2008; Rato et al.,

2010). These examples highlight the importance of

multi-locus analyses and the general use of several

lines of evidence on the integrative taxonomy

approach. Alternatively, extra lines of evidence might

balance results differently, and thus further investi-

gation is needed.

Results of nuclear data analyses showed some dif-

ferentiation between T. bocagei,T. fogoensis, and

T. darwini and amongst the Tarentola from clade B,

especially on the MC1R gene. Nuclear data also sup-

ported the differentiation of T. boavistensis from all

taxa from clade D and between T. nicolauensis and

Tarentola from clade B. Conversely, it always pre-

IDENTIFICATION KEY

1. Adults larger than or around 100 mm SVL.................................................................T. gigas..................1⬘

1⬘Long snout; 180–213 midbody scales; present in Raso Islet.....................................................T. gigas gigas

Short snout; 160–195 midbody scales; present in Branco Islet.........................................T. gigas brancoensis

Adults smaller than 100 mm SVL.......................................................................................................2

2. Enlarged tubercles between the eye and ear opening and prominent keeled dorsal tubercles.........................3

No enlarged tubercles between the eye and ear opening and rounded to oval, smoother or less apical dorsal

tubercles........................................................................................................................................6

3. Dorsal pattern with a series of usually four light middorsal patches, each preceded by a dark mark, usually

connected by a broad light middorsal line and apical tubercles................................................................4

Reduced dorsal pattern with light yellowish to grey dorsal coloration and apical tubercles ...........................5

4. Well-deﬁned four to ﬁve W-shaped dorsal bands; white ventral coloration; 22–24 scales and lamellae under ﬁfth

toe; 83 mm maximum SLV; present in Santiago Island..................................................................T. rudis

Less contrasted W-shaped dorsal bands; yellowish ventral coloration; 22–26 scales and lamellae under ﬁfth toe;

99 mm maximum SLV; present in Fogo, Brava Island, and Rombos Islets................T. protogigas................4⬘

4⬘Maximum SVL of 98 mm; weak yellowish and marbled ventrum; present in Fogo ........T. protogigas protogigas

Maximum SVL of 77 mm, distinct yellowish and uniform ventrum; present in Brava Island and Rombos Islets

..........................................................................................................................T. protogigas hartogi

5. Orangey eye iris; 19–22 interorbital scales; present in Boavista Island....................................T. boavistensis

Pale grey eye iris; 16–18 interorbital scales; present in Maio Island...........................................T. maioensis

6. Dorsal pattern with clear symmetrical butterﬂy- or X-shaped dark crossbands often lined with whitish tubercles

posteriorly......................................................................................................................................7

Dorsal pattern different....................................................................................................................9

7. Oval to round dorsal tubercles; 18–21 lamella under ﬁfth toe..................................................................8

Oblong dorsal tubercles, 20–23 lamella under ﬁfth toe; present in S. Nicolau...........................T. nicolauensis

8. Maximum SLV above 60 mm; four to ﬁve dorsal bands; usually more than 21 interorbital scales; 146–167 midbody

scales; present in S. Vicente Island......................................................................................T. substituta

Maximum SLV below 60 mm; usually three dorsal bands; 19–22 interorbital scales; 116–156 midbody scales;

present in Santa Luzia and Raso Islet.....................................................................................T. raziana

Maximum SLV above 60 mm; four to ﬁve dorsal bands; 17–20 interorbital scales; 116–150 midbody scales; present

in Santo Antão Island....................................................................................................T. caboverdiana

9. Flat oval to round dorsal tubercles slightly keeled; short trunk length and distance between nostrils and eye or

snout tip; large ear opening; wide base of tail, proﬁle of forehead concave; present in S. Nicolau ....... T. bocagei

Smooth and ﬂat oval to round dorsal tubercles; long trunk length and distance between nostrils and eye or snout

tip; large ear opening; narrow base of tail, proﬁle of forehead not concave; present in Santiago or Fogo........10

10. Dorsal pattern composed of diffuse dark or light spots, sometimes condensed to form an irregular marbling;

113–130 midbody scales; present in Santiago............................................................................T. darwini

Dorsal pattern composed of diffuse dark crossbands and spots, sometimes with a dark ring mark on the back; long

limbs; 137–148 midbody scales; present in Fogo.......................................................................T. fogoensis

356 R. VASCONCELOS ET AL.

sented haplotype sharing among specimens of the

same species but belonging to different ESUs, such as

T. protogigas from Fogo and Brava. However, nuclear

genealogies do not support conclusively all the parti-

tions observed in mtDNA, especially differentiation

between T. bocagei and T. nicolauensis,T. rudis and

T. maioensis, and T. gigas and T. protogigas. Discrep-

ant results observed between mtDNA and nuclear

genealogies are probably explained by incomplete

lineage sorting of ancestral polymorphism, as nuclear

markers are evolving at slower rates than mitochon-

drial ones. Another possible explanation could involve

male-biased gene ﬂow. Further assessment on faster

evolving nuclear markers would be valuable to

analyse this.

When haplotype sharing exists between two dif-

ferent species from separated islands that were

never connected, it is probably due to ancestral

polymorphism, as gene ﬂow is greatly reduced by

the oceanic barrier. This is the case of haplotype

sharing between T. bocagei and T. caboverdiana/

T. protogigas/T. gigas/T. rudis, between T. rudis and

T. maioensis, and between T. nicolauensis and

several species of clade D. In the case of the two

Tarentola species present in S. Nicolau, levels of

gene ﬂow were estimated to discriminate between

the inﬂuence of ancestral polymorphism and migra-

tion scenarios in shaping the patterns of allele

sharing detected by the nuclear markers. The data

strongly suggest that the polyphyletic pattern of the

nDNA networks derives from the incomplete lineage

sorting of ancestral polymorphism as the most prob-

able migration rates inferred with IMa software

were zero (Appendix 4). When differentiation is

recent, as is the case here (see Carranza et al.,

2002; Vasconcelos et al., 2010), it is probable that

mitochondrial lineages may not be monophyletic

with respect to nuclear genealogies. Another line of

evidence is that, if gene ﬂow was the main cause for

the observed pattern, we expect both ancestral and

derived alleles (located in a central or marginal

position in the haplotype network, respectively) to

be equally transpeciﬁc, which is not the case (see

Fig. 3). Although possibly allopatric, probably due to

the geological history of S. Nicolau (see Vasconcelos

et al., 2010), T. bocagei and T. nicolauensis are very

similar species that are difficult to distinguish in

the ﬁeld. This is probably due to patristic similarity

or to convergence, as both species share evolution-

ary history and identical ecological pressures.

Further morphological analyses, including coloration

and other qualitative characters, are needed to

clearly identify these species in the ﬁeld. It would

also be interesting to focus on the possible contact

zone between the two species to assess if hybridiza-

tion is occurring.

Considering clade D, it is important to note that

despite the low differentiation in mtDNA between

T. gigas and other species of this clade and between

some of these species pairs in the nuclear genes, the

alternative possibility of considering all these mono-

phyletic lineages belonging to the same species has

been refuted by several authors (see Schleich, 1987;

Joger, 1993). The main reason for this is that T. gigas

presents important morphological, bioacoustical, eco-

logical, and behavioural differences and also a very

distinct geographical distribution (north-western

islands) compared with the remaining species, includ-

ing T. protogigas. That was the grounds for exception-

ally considering T. gigas and T. protogigas as different

species even though only supported by one line of

evidence.

A genetic divergence of 2.1% in the cyt bgene was

found between T. protogigas from Fogo and the popu-

lations from Brava and Rombos, while only 0.4% was

found between populations from Brava and Rombos

(Table 5), the latter ones even sharing a mitochon-

drial haplotype (see Appendix 3). Despite the fact

that populations from Brava and Rombos were

regarded as different subspecies based on morphology

(Joger, 1993), the evidence was weak. The analysis

was based on very variable pholidotic characters

(midbody, toe, and gular scale counts) with overlap-

ping values and from very few specimens (two from

Brava and ﬁve from Rombos). A reanalysis of four

additional voucher specimens from Rombos and 11

voucher specimens from Brava using several charac-

ters clearly showed that the morphological variation

of the individuals from Rombos falls within that of

the specimens from Brava (see Appendix 2). This

result coincides with the lack of genetic differentia-

tion between these two island populations and sup-

ports the conclusion that both populations should be

regarded as part of the same subspecies. On the other

hand, as shown in Table 5, Appendices 2, 3, and 5,

MorphoBank M46037-M46091 and explained in the

‘Diagnosis’ sections of the two subspecies of T. proto-

gigas,T. protogigas from Fogo differs morphologically

from the populations from Brava and Rombos and

also presents distinct haplotypes in mtDNA. Further-

more, the geographical affinities between Fogo and

those other populations are much weaker than

between Brava and Rombos. In the presence of

further evidence supporting the split between these

two lineages, T. protogigas and T. hartogi should be

regarded as candidate species.

MANOVA of the linear measurements indicated

that males and females of Tarentola present sexual

dimorphism in size but not in shape, as they gener-

ally became non-signiﬁcant after size correction. On

the other hand, for studying differences among taxa,

all linear measurements are important as these

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 357

analyses proved that differences among populations

are due to sizes and also shapes.

This taxonomic revision has considerable conserva-

tion implications for the Cape Verdean Tarentola as

some clades were subdivided and now present more

restricted areas of occupancy and extents of occur-

rence. Thus, a revision of the conservation status is

required. Presently, Tarentola is the most taxonomi-

cally diverse genus of all the endemic reptile genera

occurring on the Cape Verde archipelago (Hemidacty-

lus,Tarentola, and Chioninia) and hence efforts

should be made to ensure that the protected areas

that are going to be implemented in the near future

encompass all this richness.

ACKNOWLEDGEMENTS

R.V. is grateful to Xavier Santos for the T. substituta

photos; to S. Rocha, M. Fonseca, and J. C. Brito from

CIBIO, J. Motta, H. Abella, and A. Nevsky for help

during ﬁeldwork; to J. César, D. Andrade, O. Freitas,

J. Gonçalves, L. Carvalho, C. Dias, I. Delgado, and

staff from the Ministério da Agricultura e Ambiente

(MAA) and to I. Gomes and all staff from the Instituto

Nacional de Investigação e Desenvolvimento Agrário

(INIDA) for logistical aid and to J. Roca for laboratory

assistance. A.P. is grateful to A. Kaliontzopoulou for

her help with morphological analysis. P.G. is grateful

to L.F. López-Jurado and J.A. Mateo. Research was

supported by grants from Fundação para a Ciência e

Tecnologia (FCT): SFRH/BD/25012/2005 (to R.V.),

SFRH/BPD/26546/2006 (to A.P.), PTDC/BIA-BDE/

74288/2006 (to D.J.H.) and PTDC/BIA-BEC/105327/

2008 (to A.P.); from the Ministerio de Educación y

Ciencia, Spain: CGL2009-11663/BOS, Grup de

Recerca Emergent of the Generalitat de Catalunya:

2009SGR1462; and an Intramural Grant from the

Consejo Superior de Investigaciones Cientíﬁcas,

Spain: 2008301031 (to S.C.). Samples were obtained

according to licence no. 07/2008 by Direcção Geral

do Ambiente, MAA, Cape Verdean Government. We

appreciate the comments of two anonymous referees

that helped to improve the manuscript substantially.

REFERENCES

Andreone F. 2000. Herpetological observations on Cape

Verde: a tribute to the Italian naturalist LEONARDO FEA

with complementary notes on Macroscincus coctei (Duméril

& Bibron 1839) (Squamata: Scincidae). Herpetozoa 13:

15–26.

Angel F. 1937. Sur la faune herpétologique de l’Archipel du

Cap Vert. XII. Congrès International Zoologie Lisbonne 1935

9: 1693–1700.

Anonymous. 2002. Boletim Oﬁcial da República de Cabo

Verde 2002. N°. 37 Série I, Anexo II. Cabo Verde: Ministério

da Justiça.

Arnold EN, Ovenden D. 2002. A ﬁeld guide to the reptiles

and amphibians of Britain and Europe. London: Collins.

Barbadillo LJ, Lacomba JI, Pérez-Mellado V, Sancho V,

López-Jurado LF. 1999. Anﬁbios y reptiles de la Península

Ibérica Baleares y Canarias. Barcelona: Geoplaneta.

Bocage JV. 1875. Sur deux Reptiles Nouveaux de l’Archipel

du Cap-Vert. Jornal de Sciencias Mathematicas Physicas e

Naturaes, Academia Real das Sciencias de Lisboa 5: 287–

290.

Bocage JV. 1896. Reptis de algumas possessões portuguezas

d’África que existem no museu de Lisboa. Jornal de Scien-

cias Mathematicas Physicas e Naturae, Academia Real das

Sciencias de Lisboa 2: 1–9 (+2 plates).

Bocage JV. 1897. Mammiferos, Repteis e Batrachios d’Africa

de que existem exemplares typicos no Museu de Lisboa.

Jornal de Sciencias Mathematicas, Physicas e Naturaes,

Academia Real das Sciencias de Lisboa 4: 187–206.

Bocage JV. 1902. Aves e Reptis de Cabo Verde. Jornal de

Sciencias Mathematicas Physicas e Naturaes Academia Real

das Sciencias de Lisboa 14: 206–210.

Boulenger GA. 1885. Catalogue of the lizards in the British

Museum (Natural History). Volume 1. Geckonidae, Euble-

pharidae, Uroplatidae, Pygopodidae, Agamidae. London:

Trustees of the British Museum.

Boulenger GA. 1906. Report on the reptiles collected by the

late L. Fea in West Africa. Annali del Museo Civico di Storia

Naturale di Genova 3: 196–216.

Brygoo É. 1990. Les types de Gekkonidés (Reptiles, Sau-

riens) du Muséum national d’Histoire naturelle Catalogue

critique. Bulletin du Muséum National d’Histoire Naturelle

(Serie 4) 12: 19–141.

Cardoso A, Serrano A, Vogler AP. 2009. Morphological and

molecular variation in tiger beetles of the Cicindela hybrida

complex: is an ‘integrative taxonomy’ possible? Molecular

Ecology 18: 648–664.

Carranza S, Arnold EN, Mateo JA, Geniez P. 2002. Rela-

tionships and evolution of the North African geckos Gecko-

nia and Tarentola (Reptilia: Gekkonidae) based on

mitochondrial and nuclear DNA sequences. Molecular Phy-

logenetics and Evolution 23: 244–256.

Carranza S, Arnold EN, Mateo JA, López-Jurado LF.

2000. Long-distance colonization and radiation in gekkonid

lizards Tarentola (Reptilia: Gekkonidae) revealed by mito-

chondrial DNA sequences. Proceedings of the Royal Society

of London B 267: 637–649.

Clement M, Posada D, Crandall KA. 2000. TCS: a com-

puter program to estimate gene genealogies. Molecular

Ecology 9: 1657–1660.

Dayrat B. 2005. Towards integrative taxonomy. Biological

Journal of the Linnean Society 85: 407–415.

De Queiroz K. 1998. The general lineage concept of species,

species chteria, and the process of speciation – a conceptual

uniﬁcation and terminological recommendations. In:

Howard DJ, Berlocher SH, eds. Endless forms: species and

speciation. New York: Oxford University Press, 57–75.

358 R. VASCONCELOS ET AL.

De Queiroz K. 2007. Species concepts and species delimita-

tion. Systematic Biology 56: 879–886.

Dekeyser PL, Villiers A. 1951. Mission J. Cadenet aux Iles

du Cap Vert. Bulletin de L’Institute Français d’Afrique Noire

13: 1152–1158.

Frazen M, Glaw F. 2007. Type catalogue of reptiles in the

Zoologische Staatssammlung München. Spixiana 30: 201–

276.

Gamble T, Bauer AM, Greenbaum E, Jackman TR. 2008.

Out of the blue: a novel trans-Atlantic clade of geckos

(Gekkota, Squamata). Zoologica Scripta 37: 355–366.

Gray JE. 1845. Catalogue of the specimens of lizards in the

collection of the British Museum. London: Trustees of the

British Museum.

Greer AE. 1976. On the evolution of the giant Cape Verde

scincid lizard Macroscincus coctei.Journal of Natural

History 10: 691–712.

Gruber HJ, Schleich HH. 1982. Hemidactylus bouvieri

razoensis nov. ssp. von den Kapverdischen Inseln (Reptilia:

Sauria-Gekkonidae). Spixiana 5: 303–310.

Hall TA. 1999. Bioedit: a user friendly biological sequence

alignment editor and analysis program for Windows 95/98/

NT. Nucleic Acids Symposium Series 41: 95–98.

Harris DJ, Batista V, Lymberakis P, Carretero MA.

2004. Complex estimates of evolutionary relationships in

Tarentola mauritanica (Reptilia: Gekkonidae) derived from

mitochondrial DNA sequences. Molecular Phylogenetics and

Evolution 30: 855–859.

Hart MW, Sunday J. 2007. Things fall apart: biological

species form unconnected parsimony networks. Biology

Letters 3: 509–512.

Hey J, Nielsen R. 2007. Integration within the Felsenstein

equation for improved Markov chain Monte Carlo methods

in population genetics. Proceedings of the National Academy

of Sciences of the United States of America 104: 2785–

2790.

Hudson RR. 2000. A new statistic for detecting genetic

differentiation. Genetics 15: 2011–2014.

Jesus J, Brehm A, Harris DJ. 2002. Relationships of Taren-

tola (Reptilia: Gekkonidae) from the Cape Verde Islands

estimated from DNA sequence data. Amphibia-Reptilia 22:

235–242.

Joger U. 1984a. Taxonomische revision der Gattung Taren-

tola (Reptilia, Gekkonidae). Bonner Zoologische Beiträge 35:

129–174.

Joger U. 1984b. Die Radiation der Gattung Tarentola in

Makaronesien (Reptilia: Sauria: Gekkonidae). Courier For-

schungsinstitut Senckenberg 71: 91–111.

Joger U. 1985. The African gekkonine radiation: preliminary

phylogenetic results based on quantitative immonological

comparisons of serum albumins. Proceedings of the Interna-

tional Symposium on African Vertebrates Bonn 1985 36:

479–494.

Joger U. 1993. On two collections of reptiles and amphibians

from the Cape Verde Islands with descriptions of three new

taxa. Courier Forschungsinstitut Senckenberg 159: 437–444.

Kaliontzopoulou A, Carretero MA, Llorente GA. 2010.

Intraspeciﬁc ecomorphological variation: linear and geomet-

ric morphometrics reveal habitat-related patterns within

Podarcis bocagei wall lizards. Journal of Evolutionary

Biology 23: 1234–1244.

Köhler J, Güsten R. 2007. Herpetological type specimens in

the natural history collections of the museums in Darms-

tadt and Wiesbaden Germany. Spixiana 30: 275–288.

Köhler G, Hertz A, Sunyer J, Seipp R, Monteiro A. 2007.

Herpetologische Forschungen auf den Kapverden unter

besonderer Berücksichtigung des Kapverdischen Riesen-

skinks Macroscincus coctei.Elaphe 15: 75–79.

López-Jurado LF, Mateo JA, Fazeres AI. 2005. Chordata.

In: Arechavaleta M, Zurita N, Marrero MC, Martín JL, eds.

Lista Preliminar de Espécies Silvestres de Cabo Verde.

Hongos Plantas y Animales Terrestres. Islas Canárias: Gobi-

erno de Canárias Consejería de Médio Ambiente, 101.

López-Jurado LF, Mateo JA, Geniez P. 1999. Los Reptiles

de la Isla de Boavista (Archipiélago de Cabo Verde). Boletín

de la Asociación Herpetológica Española 10: 10–13.

Loveridge A. 1947. Revision of the African lizards of the

family Gekkonidae. Bulletin of the Museum of Comparative

Zoology at Harvard College 98: 3–469.

Mace GM. 2004. The role of taxonomy in species conserva-

tion. Philosophical Transactions of the Royal Society of

London B 359: 711–719.

Mateo JA, García-Márquez M, López-Jurado LF, Pether

J. 1997. Nuevas Observaciones Herpetológicas en las Islas

Desertas (Archipelago de Cabo Verde). Boletín de la Asoci-

ación Herpetológica Española 8: 8–11.

Mertens R. 1954. Die Eidechsen der Kapverden. Commenta-

tiones Biologicae 15: 1–17.

Miralles A, Vasconcelos R, Perera A, Harris DJ, Car-

ranza S. 2010. An integrative taxonomic revision of the

Cape Verdean skinks (Squamata, Scincidae). Zoologica

Scripta 40: 16–44.

Monaghan MT, Wild R, Elliot M, Fujisawa T, Balke M,

Inward DJG, Lees DC, Ranaivosolo R, Eggleton P,

Barraclough TG, Vogler AP. 2009. Accelerated species

inventory on Madagascar using coalescent-based models of

species delineation. Systematic Biology 58: 1–14.

Padial JM, Miralles A, De La Riva I, Vences M. 2010. The

integrative future of taxonomy. Frontiers in Zoology 7: 1–

14.

Perera A, Harris DJ. 2010. Genetic variability within the

Oudri’s fan-footed gecko Ptyodactylus oudrii in North Africa

assessed using mitochondrial and nuclear DNA sequences.

Molecular Phylogenetics and Evolution 54: 634–639.

Pinho C, Rocha S, Carvalho BM, Lopes S, Mourão S,

Vallinoto M, Brunes TO, Haddad CFB, Gonçalves H,

Sequeira F, Ferrand N. 2010. New primers for the ampli-

ﬁcation and sequencing of nuclear loci in a taxonomically

wide set of reptiles and amphibians. Conservation Genetics

Resources 2: 181–185.

R Development Core Team. 2010. R: a language and envi-

ronment for statistical computing. Vienna: R Foundation

for Statistical Computing. Available at: http://www.R-

project.org

Rato C, Carranza S, Perera A, Carretero MA, Harris DJ.

2010. Conﬂicting patterns of nucleotide diversity between

TAXONOMIC REVISION OF CAPE VERDEAN TARENTOLA 359

mtDNA and nDNA in the Moorish gecko, Tarentola mauri-

tanica.Molecular Phylogenetics and Evolution 56: 962–971.

Rozas J, Sánchez-DelBarrio JC, Messeguer X, Rozas R.

2003. DnaSP, DNA polymorphism analyses by the coales-

cent and other methods. Bioinformatics (Oxford, England)

19: 2496–2497.

Schleich HH. 1980. Der kapverdische Riesengecko Tarentola

delalandii gigas (Bocage, 1896). Spixiana 3: 147–155.

Schleich HH. 1982a. Vorlauﬁge Mitteilung zur Herpetofauna

der Kapverden. Courier Forschunginstitut Senckenberg 52:

245–248.

Schleich HH. 1982b. Letze Nachforschungen zum Kapverdis-

chen Riesenskinks Macroscincus coctei (Dúmeril & Bibron

1839) (Reptilia: Sauria: Scincidae). Salamandra 18: 78–85.

Schleich HH. 1984. Die Geckos der Gattung Tarentola der

Kapverden (Reptilia: Sauria: Gekkonidae). Courier Fors-

chungsinstitut Senckenberg 71: 95–106.

Schleich HH. 1987. Herpetofauna Caboverdiana. Spixiana

12: 1–75.

Schleich HH. 1996. Lista Vermelha para os Répteis (Rep-

tilia). In: Leyens T, Lobin W, eds. Primeira Lista Vermelha

de Cabo Verde, vol. 193. Frankfurt: Courier Forschungsin-

stitut Senckenberg, 122–125.

Schleich HH, Wuttke M. 1983. Die Kapverdishe eilande

Santa Luzia Branco und Razo – ein Reisebericht. Natur und

Museum 113: 33–45.

Schlick-Steiner BC, Steiner FM, Seifert B, Stauffer C,

Christian E, Crozier RH. 2010. Integrative taxonomy: a

multisource approach to exploring biodiversity. Annual

Review of Entomology 55: 421–438.

Sindaco R, Jeremcˇenko VK. 2008. The reptiles of the

Western Paleartic. Latina, Italy: Societas Herpetologica

Italica.

Somers KM. 1986. Multivariate allometry and removal of

size with Principal Components Analysis. Systematic

Biology 35: 359–368.

Stephens M, Smith NJ, Donnelly P. 2001. A new statistical

method for haplotype reconstruction from population data.

American Journal of Human Genetics 68: 978–989. Avail-

able at: http://www.stat.washington.edu/stephens/

software.html

Tamura K, Dudley J, Nei M, Kumar S. 2007. MEGA4:

Molecular Evolutionary Genetics Analysis (MEGA) software

version 4.0. Molecular Biology and Evolution 24: 1596–

1599.

Templeton AR, Crandall KA, Sing CF. 1992. A cladistic

analysis of phenotypic associations with haplotypes inferred

from restriction endonuclease mapping and DNA sequence

data. III. Cladogram estimation. Genetics 132: 619–633.

Vasconcelos R, Carranza S, Harris DJ. 2010. Insight into

an island radiation: the Tarentola geckos of the Cape Verde

archipelago. Journal of Biogeography 37: 1047–1060.

Vervust B, Dongen SV, Van Damme R. 2009. The effect of

preservation on lizard morphometrics – an experimental

study. Amphibia-Reptilia 30: 321–329.

Weiss A, Hedges SB. 2007. Molecular phylogeny and bioge-

ography of the Antillean geckos Phyllodactylus wirshingi

Tarentola americana and Hemidactylus haitianus (Reptilia,

Squamata). Molecular Phylogenetics and Evolution 45: 409–

416.

Wiens JJ, Servedio MR. 2000. Species delimitation in sys-

tematics: inferring diagnostic differences between species.

Proceedings of the Royal Society of London B 267: 631–636.

Zink RM, Barrowclough GF. 2008. Mitochondrial DNA

under siege in avian phylogeography. Molecular Ecology 17:

2107–2121.

SUPPORTING INFORMATION

Additional Supporting Information may be found in the online version of this article:

Appendix S1. Details of material (live and voucher animals) and sequences used in the present study.

Appendix S2. Descriptive statistics for all the linear measurements and meristic variables of adult specimens

of the different Tarentola taxa included in this study.

Appendix S3. Networks corresponding to cytochrome bsequence variation in endemic Cape Verde Tarentola

geckos (modiﬁed from Vasconcelos et al., 2010).

Appendix S4. Marginal probabilities of migration rates (m1 and m2) and time of divergence (t) between T.

bocagei and T. nicolauensis, present in S. Nicolau Island, obtained by ﬁtting the IM model to the three-locus

data (PDC, ACM4, MC1R) set.

Appendix S5. Estimates of genetic differentiation of the PDC, ACM4 and MC1R between ESUs using Snn test

values.

360 R. VASCONCELOS ET AL.

Appendix 1. Details of material (live and voucher animals) and sequences used in the present study.

Code (live)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

T661

16.10697

-22.89861

yes

M42539-M42547

GQ381016

yes

JN208926

JN208998

JN209070

T662

16.10645

-22.89908

yes

M42548-M42558

GQ381015

yes

JN208927

JN208999

JN209071

T663

16.10567

-22.89945

yes

M42559-M42572

GQ381014

yes

JN208928

JN209000

JN209072

T664

16.04173

-22.74916

yes

M42572-M42583

GQ381013

yes

JN208929

JN209001

JN209073

T665

16.04060

-22.70674

yes

M42584-M42592

GQ381012

yes

JN208930

JN209002

JN209074

T666

16.04135

-22.70532

yes

M42592-M42602

GQ381011

yes

JN208931

JN209003

JN209075

T667

16.10755

-22.81950

yes

M42605-M42616

GQ381010

yes

JN208932

JN209004

T668

16.10264

-22.81181

yes

M42617-M42625

GQ381009

yes

JN208933

JN209005

JN209076

T669

16.10264

-22.81181

yes

M42626-M42635

GQ381008

yes

JN208934

JN209006

JN209077

T672

16.07349

-22.72051

yes

M42636-M42645

GQ381007

yes

JN208935

JN209007

JN209078

T673

16.07349

-22.72051

yes

M42646-M42659

DB2532

16.61243

-24.12988

yes, CDFA

M43461-M43477

JN185933

DB2547

16.55529

-24.08217

yes, CDFA

M43478-M43487

JN185934

DB2561

16.55529

-24.08217

yes

M55879-M55888

JN185935

DB2596

16.61243

-24.12988

yes, CDFA

M43488-M43497

JN185936

DB2597

16.55529

-24.08217

yes, CDFA

M43498-M43506

JN185937

DB2607

16.61300

-24.15359

yes, CDFA

M43507-M43515

JN185938

DB2613

16.59156

-24.08601

yes, CDFA

M43516-M43524

JN185939

DB2622

16.61300

-24.15359

yes, CDFA

M43525-M43532

DB2765

16.55529

-24.08217

yes, CDFA

M43533-M43541

JN185940

DB2792

16.61243

-24.12988

yes, CDFA

M43542-M43550

JN185941

DB2796

16.55529

-24.08217

yes, CDFA

M43551-M43559

JN185942

DB2798

16.61140

-24.11905

yes, CDFA

M43560-M43568

DB2799

16.57828

-24.07568

yes, CDFA

M43569-M43577

JN185943

DB2800

16.61243

-24.12988

yes, CDFA

M43578-M43585

JN185944

DB2801

16.61243

-24.12988

yes, CDFA

M43586-M43594

JN185945

DB2803

16.61243

-24.12988

yes, CDFA

M43595-M43603

JN185946

DB2805

16.57828

-24.07568

yes, CDFA

M43604-M43612

JN185947

DB2808

16.59792

-24.09549

yes, CDFA

M43613-M43621

JN185948

DB2809

16.55529

-24.08217

yes, CDFA

M43622-M43630

JN185949

DB2812

16.61140

-24.11905

yes, CDFA

M43631-M43639

DB2815

16.61140

-24.11905

yes, CDFA

M43640-M43647

JN185950

DB2877

16.61953

-24.12920

yes, CDFA

M43648-M43657

JN185951

DB2881

16.55529

-24.08217

yes, CDFA

M43658-M43666

JN185952

DB2885

16.61140

-24.11905

yes, CDFA

M43667-M43675

JN185953

DB2888

16.61243

-24.12988

yes, CDFA

M43676-M43683

JN185954

DB2893

16.61300

-24.15359

yes, CDFA

M43684-M43691

JN185955

DB2898

16.61243

-24.12988

yes, CDFA

M43692-M43699

JN185956

DB2899

16.61300

-24.15359

yes, CDFA

M43700-M43707

JN185957

DB2934

16.59156

-24.08601

yes, CDFA

M43708-M43716

JN185958

DB2936

16.57828

-24.07568

yes, CDFA

M43717-M43724

JN185959

DB2938

16.61300

-24.15359

yes, CDFA

M43725-M43733

JN185960

DB2939

16.61243

-24.12988

yes, CDFA

M43734-M43742

JN185961

DB2950

16.59792

-24.09549

yes, CDFA

M43743-M43751

JN185962

DB2955

16.55529

-24.08217

yes, CDFA

M43752-M43760

JN185963

T302

16.55476

-24.08140

GQ380950

yes

JN208936

JN209008

JN209079

T304

16.59085

-24.08757

SVL

M43761-M43763

GQ380952

yes

JN208937

JN209009

JN209080

T349

16.62020

-24.12912

SVL

M43764-M43767

GQ380954

yes

JN208938

JN209010

JN209081

T362

16.59264

-24.06122

SVL

M43768-M43771

GQ380963

yes

JN208939

JN209011

JN209082

T364

16.60834

-24.09522

SVL

M43772-M43775

GQ380961

yes

JN208940

JN209012

JN209083

T365

16.60964

-24.09524

SVL

M43776-M43781

GQ380962

yes

JN208941

JN209013

T533

15.01382

-24.40431

yes, CDFA

M42945-M42952

GQ380784

yes

JN208942

JN209014

JN209084

T534

14.88365

-24.41666

yes, CDFA

M42953-M42963

GQ380785

T536

14.88400

-24.41676

yes, CDFA

M42964-M42973

GQ380786

T537

14.88530

-24.41710

yes, CDFA

M42974-M42983

GQ380787

yes

JN208943

JN209015

JN209085

T538

14.86594

-24.44522

yes, CDFA

M42984-M42992

GQ380788

T539

14.86594

-24.44522

yes, CDFA

M42993-M43002

GQ380789

T540

14.86753

-24.44612

yes, CDFA

M43003-M43012

GQ380790

yes

JN208944

JN209016

JN209086

T578

15.01291

-24.41703

yes, CDFA

M43013-M43021

GQ380792

T579

15.01291

-24.41703

yes, CDFA

M43022-M43030

GQ380793

T581

14.97345

-24.42914

yes, CDFA

M43031-M43039

GQ380794

T583

14.97379

-24.45443

yes

M43040-M43046

GQ380795

yes

JN208945

JN209017

JN209087

T584

15.04369

-24.33996

yes, CDFA

M43047-M43055

GQ380796

yes

JN208946

JN209018

JN209088

T585

15.04397

-24.33767

yes, CDFA

M43056-M43063

GQ380797

T586

15.02519

-24.31852

yes, CDFA

M43064-M43072

GQ380798

T587

15.02519

-24.31852

yes, CDFA

M43073-M43080

GQ380799

T588

14.96933

-24.29293

yes, CDFA

M43080-M43087

GQ380800

Code (live)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

T589

14.96884

-24.29262

yes, CDFA

M43088-M43096

GQ380801

yes

JN208947

JN209019

JN209089

T590

14.96884

-24.29262

yes, CDFA

M43097-M43105

GQ380802

T591

14.91507

-24.34401

yes, CDFA

M43106-M43115

GQ380803

T592

14.91362

-24.34490

yes, CDFA

M43116-M43124

GQ380804

T593

14.91362

-24.34490

yes, CDFA

M43125-M43133

GQ380805

T594

14.90025

-24.35563

yes, CDFA

M43134-M43143

GQ380806

T595

14.90025

-24.35563

yes, CDFA

M43144-M43152

GQ380807

T596

14.89915

-24.35590

yes, CDFA

M43153-M43161

GQ380808

yes

JN208948

JN209020

JN209090

T597

14.84548

-24.32733

yes, CDFA

M43162-M43170

GQ380809

T599

14.86309

-24.44382

yes, CDFA

M43171-M43179

GQ380810

yes

JN208949

JN209021

JN209091

T600

14.89229

-24.30076

yes, CDFA

M43180-M43188

GQ380811

yes

JN208950

JN209022

JN209092

T601

14.89229

-24.30076

yes, CDFA

M43189-M43197

GQ380812

T602

14.89283

-24.30111

yes, CDFA

M43198-M43206

GQ380813

yes

JN208951

JN209023

JN209093

T603

14.83589

-24.39089

yes

M43207-M43209

GQ380814

yes

JN208952

JN209024

JN209094

T606

14.90883

-24.41889

yes, CDFA

M43210-M43218

GQ380817

T370

15.03740

-23.62620

yes

M44231-M44239

GQ380825

yes

JN208953

JN209025

JN209095

T373

14.91247

-23.59675

yes, CDFA

M44240-M44248

GQ380827

T374

14.91247

-23.59675

yes, CDFA

M44249-M44255

GQ380831

T375

14.91247

-23.59675

yes

M44256-M44265

GQ380826

T378

15.00739

-23.52359

yes

M44266-M44273

GQ380863

yes

JN208954

JN209026

JN209096

T389

15.10945

-23.51747

yes, CDFA

M44274-M44281

GQ380837

T390

15.10945

-23.51747

yes

M44282-M44288

GQ380834

T392

14.94052

-23.67154

yes

M44289-M44296

GQ380843

yes

JN208955

JN209027

JN209097

T394

14.94524

-23.67117

yes, CDFA

M44297-M44304

GQ380835

T395

14.95536

-23.67062

yes, CDFA

M44305-M44312

GQ380822

yes

JN208956

JN209028

JN209098

T397

14.95567

-23.67068

yes

M44313-M44320

GQ380820

T398

14.92932

-23.63896

yes

M44321-M44329

GQ380842

T399

14.92932

-23.63896

yes, CDFA

M44330-M44337

GQ380823

yes

JN208957

JN209029

JN209099

T400

14.92871

-23.63700

yes, CDFA

M44338-M44345

GQ380824

yes

JN208958

JN209030

JN209100

T404

14.96614

-23.58241

yes

M44346-M44353

GQ380849

yes

JN208959

JN209031

JN209101

T405

14.96760

-23.58226

yes, CDFA

M44354-M44361

GQ380821

T406

14.96760

-23.58226

yes, CDFA

M44362-M44369

GQ380848

T409

14.94532

-23.55602

yes

M44370-M44377

GQ380844

yes

JN208960

JN209032

JN209102

T411

14.94532

-23.55602

yes

M44379-M44388

GQ380862

T412

15.06036

-23.47457

yes

M44389-M44396

GQ380850

T413

15.06079

-23.47494

yes

M44397-M44405

GQ380885

yes

JN208961

JN209033

JN209103

T414

15.05783

-23.47778

yes

M44406-M44414

GQ380876

T415

15.05433

-23.47058

yes

M44415-M44422

GQ380853

T416

15.05367

-23.47090

yes

M44423-M44430

GQ380857

T417

15.05367

-23.47090

yes, CDFA

M44431-M44438

GQ380858

T420

15.03371

-23.52336

yes

M44439-M44446

GQ380877

T421

15.03451

-23.52552

yes, CDFA

M44447-M44454

GQ380856

T422

15.15363

-23.56845

yes, CDFA

M44455-M44465

GQ380934

yes

JN208962

JN209034

JN209104

T423

15.15382

-23.56897

yes, CDFA

M44471-M44480

GQ380901

yes

JN208963

JN209035

JN209105

T424

15.15382

-23.56897

yes

M44481-M44488

GQ380893

yes

JN208964

JN209106

T425

15.15378

-23.56922

yes

M44489-M44496

GQ380900

T426

15.15492

-23.56569

yes

M44977-M44984

GQ380899

yes

JN208965

JN209036

JN209107

T428

15.15492

-23.56569

yes

M44497-M44504

GQ380894

T431

15.16898

-23.58145

yes

M44505-M44512

GQ380910

yes

JN208966

JN209037

JN209108

T432

15.16898

-23.58145

yes

M44513-M44521

GQ380942

T433

15.16814

-23.58249

yes, CDFA

M44522-M44528

GQ380919

T437

15.13990

-23.74731

yes

M44529-M44536

GQ380928

T438

15.13990

-23.74731

yes

M44537-M44543

GQ380911

T439

15.14013

-23.74904

yes

M44544-M44551

GQ380933

T440

15.14013

-23.74904

yes, CDFA

M44552-M44559

GQ380927

T444

15.15475

-23.63381

yes, CDFA

M44560-M44568

GQ380908

T445

15.15442

-23.63362

yes

M44569-M44576

GQ380920

T446

15.19741

-23.60256

yes, CDFA

M44577-M44584

GQ380902

yes

JN208967

JN209038

JN209109

T451

14.94624

-23.62375

yes

M44585-M44592

GQ380819

T452

14.94624

-23.62375

yes

M44593-M44600

GQ380881

T453

14.99202

-23.62272

yes

M44601-M44608

GQ380882

T462

14.94774

-23.49853

yes

M44609-M44616

GQ380846

T464

15.26524

-23.75379

yes

M44617-M44625

GQ380912

T468

15.06631

-23.60313

yes

M44626-M44634

GQ380865

T469

15.08512

-23.60028

yes, CDFA

M44635-M44643

GQ380828

T470

15.07021

-23.69503

yes, CDFA

M44644-M44651

GQ380915

T471

15.03814

-23.59595

yes

M44652-M44659

GQ380830

T472

15.05110

-23.57769

yes

M44660-M44669

GQ380866

T473

15.06663

-23.61430

yes

M44670-M44675

GQ380867

T474

15.09981

-23.71310

yes

M44676-M44683

GQ380914

Code (live)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

T475

15.09981

-23.71310

yes

M44684-M44692

GQ380917

T476

15.09592

-23.76615

yes

M44693-M44700

GQ380923

T477

15.09592

-23.76615

yes

M44701-M44708

GQ380925

T478

15.09592

-23.76615

yes

M44709-M44716

GQ380922

T480

15.10786

-23.76891

yes

M44717-M44725

GQ380916

T482

15.24671

-23.72318

yes, CDFA

M44726-M44733

GQ380946

yes

JN208968

JN209039

T483

15.24688

-23.72321

yes

M44734-M44741

GQ380945

T484

15.25059

-23.72422

yes

M44742-M44749

GQ380887

T486

15.25258

-23.72522

yes

M44750-M44758

GQ380897

T487

15.25247

-23.72516

yes

M44759-M44766

GQ380941

yes

JN208969

JN209040

JN209110

T488

15.30745

-23.70952

yes

M44767-M44775

GQ380891

T489

15.30745

-23.70952

yes

M44776-M44785

GQ380895

T490

15.30746

-23.70870

yes

M44786-M44794

GQ380898

yes

JN208970

JN209041

JN209111

T492

15.28715

-23.71260

yes

M44795-M44803

GQ380896

T493

15.28716

-23.71167

yes

M44804-M44812

GQ380889

yes

JN208971

JN209042

JN209112

T494

15.08660

-23.70932

yes, CDFA

M44813-M44821

GQ380948

T495

15.28076

-23.73057

yes

M44822-M44829

GQ380913

T496

15.28076

-23.73057

yes

M44830-M44838

GQ380947

T501

15.16521

-23.62645

yes

M44839-M44847

GQ380904

T502

15.16288

-23.62428

yes

M44848-M44856

GQ380905

T503

15.14243

-23.65662

yes

M44857-M44865

GQ380907

T505

15.11527

-23.62050

yes, CDFA

M44867-M44876

GQ380940

T508

15.18607

-23.67201

yes, CDFA

M44877-M44885

GQ380909

T509

15.18607

-23.67201

yes, CDFA

M44886-M44895

GQ380936

T510

15.18099

-23.67165

yes, CDFA

M44896-M44904

GQ380924

T511

15.18275

-23.67104

yes

M44905-M44913

GQ380939

T512

15.00521

-23.53196

yes

M44914-M44922

GQ380868

T513

15.00521

-23.53196

yes, CDFA

M44923-M44931

GQ380869

T516

15.06959

-23.55709

yes, CDFA

M44932-M44940

GQ380872

T523

15.05336

-23.46686

yes, CDFA

M44941-M44949

GQ380874

T524

15.05336

-23.46686

yes, CDFA

M44950-M44958

GQ380875

T526

15.06558

-23.76531

yes

M44959-M44967

GQ380929

T529

15.04916

-23.70036

yes, CDFA

M44968-M44976

GQ380932

T003

16.87172

-24.99760

SVL

GQ381038

yes

JN209113

T014

16.85089

-24.87269

SVL

M55646-M55649

GQ381040

yes

JN209114

T017

16.84454

-24.88425

SVL

M55651-M55652

GQ381041

yes

JN209115

T021

16.85035

-24.92702

SVL

GQ381042

yes

JN209116

T032

16.86289

-24.94440

SVL

GQ381044

yes

JN209117

T035

16.90569

-24.93826

SVL

M55653-M55656

GQ381046

yes

JN209118

T037

16.90390

-24.94287

SVL

M55657-M55660

GQ381047

yes

JN209119

T038

16.90492

-24.94089

SVL

M55661-M55663

GQ381048

yes

JN209120

T040

16.90492

-24.94089

SVL

yes

JN209121

T043

16.90851

-24.92862

SVL

GQ381049

yes

JN208972

JN209043

JN209122

T046

16.89993

-24.95152

SVL

M44991-M44994

GQ381051

yes

JN208973

JN209044

JN209123

T048

16.83013

-25.07208

SVL

M55664-M55666

GQ381053

yes

JN209124

T055

16.82982

-25.07634

SVL

M55667-M55670

GQ381055

yes

JN209125

T059

16.82848

-25.06429

SVL

M55671-M55672

GQ381056

yes

JN209126

T064

16.86241

-24.98141

SVL

M55673-M55676

GQ381057

yes

JN209127

T070

16.85721

-24.98110

SVL

M55677-M55676

GQ381058

yes

JN209128

T077

16.83474

-24.96736

SVL

M55677-M55683

GQ381059

yes

JN209129

T096

16.86031

-24.94325

SVL

M55684-M55686

GQ381064

yes

JN209130

T102

16.86817

-24.95305

SVL

M55687-M55689

GQ381066

yes

JN209131

T105

16.87582

-25.01886

SVL

GQ381067

yes

JN209132

T109

16.87695

-25.02152

SVL

M55690-M55692

GQ381068

yes

JN209133

T124

16.84348

-25.03513

SVL

GQ381072

yes

JN209134

T125

16.84348

-25.03513

SVL

M55693

GQ381073

yes

JN209135

T134

16.81443

-24.89439

SVL

M55694-M55696

GQ381079

yes

JN209136

T135

16.82938

-25.03060

SVL

GQ381080

yes

JN209137

T136

16.81633

-25.02315

SVL

M55697-M55698

GQ381081

yes

JN209138

T144

16.79072

-24.78505

SVL

M44995-M44997

GQ381027

yes

JN208974

JN209045

T145

16.78341

-24.77890

SVL

M44998-M45000

GQ381017

yes

JN208975

JN209046

JN209139

T150

16.61249

-24.60066

SVL

M55699-M55701

GQ381021

yes

JN209140

T151

16.79495

-24.77025

SVL

GQ381018

yes

JN209141

T154

16.78680

-24.76163

SVL

GQ381019

yes

JN209142

T158

16.76750

-24.77643

SVL

GQ381025

yes

JN209143

T162

16.76126

-24.76168

SVL

GQ381026

yes

JN209144

T164

16.74295

-24.74024

SVL

GQ381020

yes

JN209145

T166

16.74838

-24.73827

SVL

M55702-M55704

GQ381023

yes

JN209146

T170

16.73945

-24.71607

SVL

M55705-M55706

GQ381030

yes

JN209147

T172

16.74792

-24.70182

SVL

M55707-M55709

GQ381028

yes

JN209148

Code (live)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

T174

16.77977

-24.77000

SVL

M55710-M55711

GQ381024

yes

JN209149

T178

16.77584

-24.76580

SVL

M55712-M55714

GQ381031

yes

JN209150

Tra1

16.61249

-24.60066

SVL

GQ381032

yes

JN208976

JN209047

JN209151

Tra2

16.61249

-24.60066

SVL

GQ381033

yes

JN209152

Tra3

16.61249

-24.60066

SVL

GQ381029

yes

JN209153

Tra5

16.61249

-24.60066

SVL

GQ381034

yes

JN209154

Tra6

16.61249

-24.60066

SVL

GQ381035

yes

JN209155

Tra7

16.61249

-24.60066

SVL

GQ381036

yes

JN209156

T181

17.19484

-25.09118

SVL

M55715-M55720

GQ381097

yes

JN209157

T187

17.17077

-25.16326

GQ381108

yes

JN209158

T188

17.16766

-25.09881

SVL

M45001-M45003

GQ381117

yes

JN208977

JN209048

JN209159

T189

17.08340

-25.14861

SVL

M55727-M55729

GQ381100

yes

JN209160

T192

17.11002

-25.08886

SVL

M45004-M45007

GQ381119

yes

JN208978

JN209049

JN209161

T194

17.10095

-25.06384

SVL

M55721-M55723

GQ381088

yes

JN209162

T196

17.05217

-25.06745

SVL

M45008-M45010

GQ381089

yes

JN209050

T198

17.04673

-25.06845

SVL

M55724-M55726

GQ381090

yes

JN209163

T199

17.10948

-25.26420

SVL

GQ381091

yes

JN209164

T203

17.10122

-25.26699

GQ381093

yes

JN209165

T204

17.02456

-25.05634

SVL

M55730-M55732

GQ381111

yes

JN209166

T206

17.02392

-25.06249

SVL

M55733

GQ381094

yes

JN209167

T207

17.10742

-25.27155

SVL

M55734-M55736

GQ381095

yes

JN209168

T210

17.02278

-25.26347

SVL

GQ381096

yes

JN209169

T211

16.98320

-25.23454

SVL

M55737-M55739

GQ381109

yes

JN209170

T215

17.06053

-25.18749

SVL

GQ381102

yes

JN209171

T219

17.03717

-25.19258

SVL

M55740-M55742

GQ381118

yes

JN209172

T222

17.08197

-25.06486

SVL

M55743-M55745

GQ381112

yes

JN209173

T228

17.01747

-25.16001

SVL

M55746-M55748

GQ381114

yes

JN209174

T238

17.03588

-25.03919

SVL

M55749-M55755

GQ381086

yes

JN209175

T240

17.02712

-25.04083

SVL

M45011-M45014

GQ381116

yes

JN208979

JN209051

T248

17.02214

-25.32942

SVL

GQ381087

yes

JN209176

T256

16.96542

-25.31289

SVL

M55756-M55758

GQ381106

yes

JN209177

T259

16.95450

-25.30810

SVL

M55760-M55761

GQ381107

yes

JN209178

DB1544

16.61756

-24.27407

yes

M45653-M45661

JN185964

DB2422

16.61490

-24.39946

yes

M45662-M45671

JN185965

DB2535

16.65512

-24.31653

yes

M45672-M45681

JN185966

DB2540

16.61369

-24.29219

yes

M45682-M45689

JN185967

DB2562

16.58686

-24.32895

yes

M45690-M45698

JN185968

DB2580

16.59210

-24.39728

yes

M45699-M45707

JN185969

DB2589

16.58686

-24.32895

yes

M45708-M45716

JN185970

DB2591

16.61490

-24.39946

yes

M45717-M45726

JN185971

DB2605

16.61756

-24.27407

yes

M45727-M45733

JN185972

DB2610

16.59275

-24.30092

yes

M45734-M45743

JN185973

DB2624

16.61490

-24.39946

yes

M45744-M45752

JN185974

DB2773

16.65512

-24.31653

yes

M45753-M45761

JN185975

DB2794

16.66732

-24.37939

yes

M45762-M45770

JN185976

DB2795

16.65512

-24.31653

yes

M45771-M45780

JN185977

DB2804

16.61369

-24.29219

yes

M45781-M45789

JN185978

DB2806

16.61490

-24.39946

yes

M45790-M45798

JN185979

DB2816

16.58686

-24.32895

yes

M45799-M45807

JN185980

DB2817

16.66732

-24.37939

yes

M45808-M45816

JN185981

DB2824

16.61756

-24.27407

yes

M45817-M45825

JN185982

DB2828

16.61369

-24.29219

yes

M45826-M45834

JN185983

DB2829

16.61490

-24.39946

yes

M45835-M45843

JN185984

DB2840

16.56649

-24.28285

yes

M45844-M45852

JN185985

DB2880

16.58686

-24.32895

yes

M45853-M45861

JN185986

DB2884

16.56649

-24.28285

yes

M45862-M45870

JN185987

DB2892

16.66732

-24.37939

yes

M45871-M45880

JN185988

DB2894

16.61756

-24.27407

yes

M45881-M45889

JN185989

DB2895

16.61756

-24.27407

yes

M45890-M45899

JN185990

DB2902

16.56649

-24.28285

yes

M45900-M45908

JN185991

DB2903

16.59275

-24.30092

yes

M45909-M45917

JN185992

DB2932

16.66732

-24.37939

yes

M45918-M45926

JN185993

DB2935

16.65512

-24.31653

yes

M45927-M45935

DB2940

16.56649

-24.28285

yes

M45936-M45944

JN185994

DB2941

16.56649

-24.28285

yes

M45945-M45953

JN185995

DB2945

16.59275

-24.30092

yes

M45954-M45963

JN185996

DB2957

16.65512

-24.31653

yes

M45964-M45973

DB2958

16.65512

-24.31653

yes

M45974-M45983

JN185997

T288

16.56635

-24.34141

SVL

M45984-M45986

GQ380986

yes

JN208980

JN209052

T311

16.61661

-24.31657

SVL

M45987-M45989

GQ380972

yes

JN208981

JN209053

JN209179

Code (live)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

T315

16.66047

-24.31520

SVL

M45990-M45992

GQ380994

yes

JN208982

JN209054

JN209180

EU293662

Desertas

yes

EU293662

EU293707

Desertas

yes

EU293707

Tg02

Tgg

16.61249

-24.60066

colouration

M45993-M45994

GQ381127

yes

JN208983

JN209055

JN209181

Tg03

Tgg

16.61249

-24.60066

colouration

M45995

GQ381128

yes

JN208984

JN209056

JN209182

T371

14.91610

-23.60448

yes

M45996-M46002

GQ380725

yes

JN208985

JN209057

JN209183

T372*

14.91681

-23.60409

yes

GQ380841

T377

15.00821

-23.52470

yes

M46003-M46011

GQ380726

yes

JN208986

JN209058

JN209184

T380*

15.00948

-23.51746

yes

M46012-M46020

GQ380833

yes

JN208987

JN209059

JN209185

T381*

15.00948

-23.51746

yes

GQ380832

T382

15.00948

-23.51746

yes

M46021-M46028

GQ380727

yes

JN208988

JN209060

JN209186

T383*

15.00948

-23.51746

yes

GQ380836

T384*

15.02817

-23.57685

yes

M46029-M46036

GQ380886

yes

JN208989

JN209061

JN209187

T385

15.02817

-23.57685

yes

GQ380732

T386

15.03336

-23.59183

yes

GQ380730

T387

15.03336

-23.59183

yes

GQ380731

T401

14.95411

-23.56670

yes

GQ380742

T408

14.94532

-23.55602

yes

GQ380741

T410

14.94532

-23.55602

yes

GQ380733

T419

15.03371

-23.52336

yes

GQ380739

T435

15.04782

-23.61719

yes

GQ380854

T441*

14.99319

-23.52917

yes

GQ380861

T442

14.99319

-23.52917

yes

GQ380734

T443*

14.99220

-23.52856

yes

GQ380840

T454*

14.99202

-23.62266

yes

GQ380883

T461

14.94758

-23.49884

yes

GQ380740

T517

15.01704

-23.58025

yes

GQ380735

T518

14.99946

-23.57837

yes

GQ380736

T519

14.99883

-23.57908

yes

GQ380737

T520

14.99883

-23.57908

yes

T532

Tpp

14.91455

-24.45351

yes

M46037-M46046

GQ380781

yes

JN208990

JN209062

JN209188

T535

Tpp

14.88365

-24.41666

yes

M46047-M46055

GQ380782

yes

JN208991

JN209063

JN209189

T541

Tph

14.89053

-24.68965

yes

M46056-M46064

GQ380767

yes

JN208992

JN209064

JN209190

T542

Tph

14.89134

-24.68932

yes

T545

Tph

14.86478

-24.74425

yes

M46065-M46073

GQ380769

yes

JN208993

JN209065

JN209191

T546

Tph

14.86455

-24.74429

yes

T547

Tph

14.86341

-24.74533

yes

T548

Tph

14.87332

-24.73007

yes

M46074-M46082

GQ380770

yes

JN208994

JN209066

JN209192

T549

Tph

14.87241

-24.70266

yes

GQ380771

T553

Tph

14.83135

-24.73407

yes

T554

Tph

14.83135

-24.73407

yes

T555

Tph

14.83216

-24.73434

yes

T558

Tph

14.84299

-24.73560

yes

T559

Tph

14.84314

-24.73609

yes

T560

Tph

14.84299

-24.73560

yes

T561

Tph

14.85044

-24.72604

yes

M46083-M46091

GQ380774

yes

JN208995

JN209067

JN209193

T563

Tph

14.85233

-24.72713

yes

T564

Tph

14.85233

-24.72713

yes

T565

Tph

14.84556

-24.67676

yes

GQ380775

T566

Tph

14.84583

-24.67625

yes

T567

Tph

14.84568

-24.67580

yes

T568

Tph

14.84414

-24.67431

yes

T569

Tph

14.85590

-24.68782

yes

GQ380776

T571

Tph

14.83178

-24.70087

yes

GQ380778

T572

Tph

14.83148

-24.70070

yes

T573

Tph

14.83148

-24.70070

yes

T574

Tph

14.83706

-24.71593

yes

T575

Tph

14.83691

-24.71569

yes

GQ380779

T576

Tph

14.84658

-24.71339

yes

GQ380780

T607

15.23536

-23.21131

SVL

M46092-M46100

GQ380743

yes

JN208996

JN209068

JN209194

T610

15.18570

-23.17992

SVL

GQ380744

T613

15.19017

-23.15439

SVL

GQ380746

T619

15.17609

-23.22275

SVL

GQ380747

T621

15.32239

-23.18520

SVL

GQ380748

T625

15.13488

-23.18450

SVL

GQ380749

T628

15.17714

-23.13047

SVL

GQ380750

T631

15.14746

-23.12474

SVL

GQ380751

T634

15.27249

-23.16318

SVL

GQ380752

T637

15.28472

-23.15853

SVL

GQ380753

T640

15.19690

-23.11949

SVL

M46101-M46109

GQ380754

yes

JN208997

JN209069

JN209195

Code (live)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

T643

15.20626

-23.11115

SVL

GQ380755

T645

15.24676

-23.11164

SVL

GQ380756

T647

15.27751

-23.11127

SVL

GQ380757

T650

15.27997

-23.11126

SVL

GQ380758

T653

15.22301

-23.14336

SVL

GQ380760

Code (vouchers)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

BEV.11065

16.15

-22.90

yes

BEV.11066

15.99

-22.88

yes

BEV.11067

15.99

-22.81

yes

DB-ULPGC-GG-1

16.18

-22.92

yes

AF185009

DB-ULPGC-GG-2

15.99

-22.79

yes

AF185008

MNHN 2011.0213

16.18

-22.92

yes

BEV.11068

16.61

-24.13

yes

BMNH 1998.346†

16.61

-24.13

yes

M55894-M55895

AF185036

MNHN 2011.0201††

16.56

-24.08

yes

M55896-M55901

JN185934

MNHN 2011.0202†

16.56

-24.08

yes

M55889-M55893

JN185935

BEV.11069

15.03

-24.32

yes

BEV.11070

14.86

-24.39

yes

BEV.11071

14.89

-24.49

yes

BEV.11072†

14.98

-24.44

yes

M55912-M55919

BEV.11073

14.98

-24.44

yes

BEV.11074

14.98

-24.44

yes

DB-ULPGC-GG-6

14.90

-24.50

yes

AF185044

MNHN 2011.0203††

14.98

-24.44

yes

M55902-M55907

MNHN 2011.0204†

14.98

-24.44

yes

M55908-M55911

BEV.11075

yes

BEV.11076

yes

BEV.11077

yes

BEV.11078

yes

BEV.11079

yes

BEV.11080

yes

BEV.11081

yes

BEV.11082

yes

BEV.11083

yes

BEV.11084

yes

BEV.11085

yes

BEV.11086

yes

BEV.11087

yes

BEV.11088

15.28

-23.75

yes

BEV.11089

15.28

-23.75

yes

BEV.11090

15.28

-23.75

yes

DB-ULPGC-GG-3

15.09

-23.66

yes

AF185038

DB-ULPGC-GG-4

15.09

-23.66

yes

AF185040

DB-ULPGC-GG-5

15.25

-23.72

yes

AF185043

MNHN 2011.0205

yes

MNHN 2011.0206

yes

BEV.11091

16.87

-24.94

yes

BEV.11092

16.87

-24.94

yes

BEV.11093

16.87

-24.94

yes

BEV.11094

yes

BEV.11095

yes

BEV.11096

16.87

-24.94

yes

BEV.11097

16.87

-24.94

yes

BEV.11098

16.85

-24.87

yes

DB-ULPGC-GG-10

16.87

-24.94

yes

AF185030

MNHN 2011.0214

yes

BEV.11099

16.62

-24.59

yes

BEV.11100

16.62

-24.59

yes

BEV.11101

16.62

-24.59

yes

BEV.11102

16.62

-24.59

yes

BEV.11103

16.62

-24.59

yes

BEV.11104

16.62

-24.59

yes

BEV.11105

16.62

-24.59

yes

BEV.11106

16.62

-24.59

yes

BEV.11107

16.62

-24.59

yes

BEV.11108

16.62

-24.59

yes

BEV.11109

16.62

-24.59

yes

BEV.11110

16.62

-24.59

yes

BEV.11111

16.62

-24.59

yes

BEV.11112

16.77

-24.75

yes

BEV.11113

16.77

-24.75

yes

BEV.11114

16.77

-24.75

yes

BEV.11115

16.77

-24.75

yes

BEV.11116

16.77

-24.75

yes

BMNH 1998.361

16.62

-24.59

yes

AF185033

MNHN 2011.0207

16.62

-24.59

yes

Code (vouchers)

Taxa

ESU

Island

Lat

Long

Morphology

MorphoBank

mtDNA

nDNA

PDC

ACM4

MC1R

DB-ULPGC-GG-09

16.62

-24.59

yes

AF185033

BEV.11117

17.02

-25.07

yes

BEV.11118

16.99

-25.19

yes

GQ380712

BEV.11119

yes

BEV.11120

17.09

-25.14

yes

BEV.11121

17.09

-25.14

yes

BEV.11122

17.02

-25.09

yes

BEV.11123

17.11

-25.24

yes

MNHN 2011.0208

17.09

-25.14

yes

BEV.11124

16.61

-24.42

yes

BEV.11125

16.64

-24.32

yes

BEV.11126

16.64

-24.32

yes

BEV.11127

16.56

-24.28

yes

BEV.11128

16.56

-24.28

yes

DB-ULPGC-GG-8

yes

MNHN 2011.0209

16.64

-24.32

yes

BEV.6120

Tgb

16.66

-24.67

yes

BEV.9190

Tgg

16.62

-24.59

yes

BEV.9191

Tgb

16.66

-24.67

yes

BEV.11129

14.91

-23.51

yes

BEV.11130

14.91

-23.51

yes

BEV.11131

14.91

-23.51

yes

BEV.11132

14.91

-23.51

yes

BEV.11133

14.91

-23.51

yes

BEV.11134

14.91

-23.51

yes

BEV.11135

14.91

-23.51

yes

BEV.11136

14.91

-23.51

yes

DB-ULPGC-GG-11

14.91

-23.51

yes

AF185013

MNHN 2011.021

14.91

-23.51

yes

BEV.11137

Tph

14.88

-24.70

yes

BEV.11138

Tph

14.88

-24.69

yes

BEV.11139

Tph

14.84

-24.72

yes

BEV.11140

Tph

14.84

-24.72

yes

BEV.11141

Tph

14.81

-24.71

yes

BEV.11142

Tph

14.83

-24.70

yes

BEV.11143

Tph

14.83

-24.70

yes

BEV.11144

Tph

14.83

-24.70

yes

DB-ULPGC-GG-15

Tph

yes

AF185025

DB-ULPGC-GG-16

Tph

yes

AF185028

MNHN 2011.0211

Tph

14.83

-24.70

yes

BEV.11145

Tph

yes

BEV.11146

Tph

yes

DB-ULPGC-GG-13

Tph

yes

AF185020

DB-ULPGC-GG-14

Tph

yes

AF185021

BEV.11147

15.27

-23.20

yes

BEV.11148

15.15

-23.13

yes

BEV.11149

15.31

-23.15

yes

BEV.11150

15.27

-23.12

yes

MNHN 2011.0212

15.32

-23.12

yes

CDFA refers to Canonical Discriminant Function Analysis. Individuals marked with * have

introgressed mtDNA. Individuals marked with † † are holotypes and with † are paratypes.

Tv, T. boavistensis; Tb, T. bocagei; Tf, T. fogoensis; Td, T. darwini; Ts, T. substituta; Tz, T. raziana;

Tc, T. caboverdiana; Tn, T. nicolauensis; Tg, T. gigas; Tgg, T. gigas gigas; Tr, T. rudis; Tpp, T.

protogigas protogigas; Tph, T. protogigas hartogi; Tm, T. maioensis.

SV, S. Vicente; SL, Sta. Luzia; ra, Raso; br, Branco; SA, Santo Antão; SN, S. Nicolau; ST, Santiago;

sm, Santa Maria; F, Fogo; B, Brava; ro, Rombos; M, Maio; BV, Boavista.

Appendix 2. Descriptive statistics for all the linear measurements and meristic variables of adult specimens of the different Tarentola taxa included in

this study.

T. boavistensis

T. nicolauensis

Boavista

São Nicolau

Males (N = 6)

Females (N = 5)

All (N = 11)

Males (N = 18)

Females (N = 18)

All (N = 36)

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

SVL

69.67±7.09

60.00–79.00

59.9±4.01

53.50–64.00

65.23±7.59

53.50–79.00

58.97±5.16

51.50–71.00

58.31±4.51

50.00–65.50

58.64±4.79

50.00–71.00

TrL

27.94±3.26

23.28–31.09

24.79±2.01

22.07–27.73

26.51±3.1

22.07–31.09

22.70±2.67

19.61–30.33

23.46±2.08

20.04–27.49

23.08±2.39

19.61–30.33

6.73±0.74

5.68–7.52

5.56±0.57

4.93–6.42

6.20±0.88

4.93–7.52

6.75±0.94

5.45–9.11

6.43±0.90

5.07–8.30

6.59±0.92

5.07–9.11

FLL

26.56±2.92

23.37–31.81

23.22±1.11

21.25–23.86

25.04±2.79

21.25–31.81

20.33±1.83

17.42–23.93

19.70±1.68

16.80–22.88

20.02±1.76

16.80–23.93

CFL

16.94±2.31

13.06–19.66

14.12±0.86

13.19–15.24

15.65±2.27

13.06–19.66

12.98±1.26

10.52–15.83

12.48±1.26

10.63–14.65

12.73±1.27

10.52–15.83

HLL

32.60±3.65

27.04–36.83

27.82±2.35

24.55–30.66

30.43±3.89

24.55–36.83

25.09±2.07

22.36–29.39

24.61±1.97

20.69–28.17

24.86±2.01

20.69–29.39

FFL

19.02±1.93

16.28–21.75

16.21±1.01

15.22–17.45

17.74±2.10

15.22–21.75

14.77±1.19

12.79–16.64

14.64±0.98

12.90–16.21

14.71±1.08

12.79–16.64

15.94±1.02

14.75–17.13

14.02±0.54

13.33–14.76

15.07±1.28

13.33–17.13

13.54±1.39

11.40–16.99

12.91±1.05

11.29–14.78

13.23±1.26

11.29–16.99

9.63±0.92

8.45–10.80

8.58±0.34

8.10–8.99

9.16±0.88

8.10–10.80

8.18±0.75

7.25–10.27

7.92±0.75

6.89–9.17

8.05±0.75

6.89–10.27

4.09±0.34

3.62–4.54

3.79±0.33

3.36–4.20

3.96±0.35

3.36–4.54

3.68±0.21

3.32–4.19

3.76±0.28

3.35–4.21

3.72±0.25

3.32–4.21

2.70±0.38

2.28–3.31

2.32±0.36

1.88–2.82

2.53±0.40

1.88–3.31

2.44±0.33

1.93–3.19

2.49±0.29

1.78–2.94

2.46±0.31

1.78–3.19

NED

6.75±0.55

6.26–7.74

6.11±0.38

5.70–6.54

6.46±0.57

5.70–7.74

5.51±0.45

4.80–6.45

5.44±0.45

4.39–6.07

5.47±0.45

4.39–6.45

SED

8.43±0.68

7.70–9.53

7.69±0.52

7.23–8.32

8.09±0.69

7.23–9.53

6.74±0.87

5.24–8.93

6.44±0.89

5.06–8.02

6.59±0.88

5.06–8.93

EED

7.47±0.83

6.55–8.68

5.91±0.43

5.36–6.52

6.76±1.04

5.36–8.68

5.23±0.47

4.55–6.20

5.24±0.50

4.35–6.08

5.23±0.48

4.35–6.20

SLS

9.67±1.03

8–11

9.40±1.14

8–11

9.55±1.04

8–11

11.00±0.84

9–12

10.22±0.65

9–12

10.61±0.84

9–12

ILS

7.50±0.55

7–8

7.80±0.84

7–9

7.64±0.67

7–9

9.11±0.58

8–10

8.67±0.69

8–10

8.89±0.67

8–10

Lam

9.67±0.52

9–10

9.67±0.58

9–10

9.67±0.50

9–10

9.11±0.68

8–10

9.35±0.70

8–11

9.23±0.69

8–11

Trow

15.33±1.03

14–16

16.40±1.14

15–18

15.82±1.17

14–18

15.67±1.03

14–17

15.67±1.19

14–18

15.67±1.10

14–18

Tline

22.17±1.33

20–24

23.40±1.34

21–24

22.73±1.42

20–24

20.56±1.85

18–24

21.61±2.33

18–26

21.08±2.14

18–26

Srow

2.17±0.26

2.00–2.50

2.20±0.84

1.00–3.00

2.18±0.56

1.00–3.00

2.56±0.57

2.00–4.00

2.42±0.46

2.00–3.00

2.49±0.51

2.00–4.00

medS

1.25±0.32

0.75–1.50

1.15±0.38

0.75–1.75

1.20±0.33

0.75–1.75

2.10±0.38

1.25–2.75

2.14±0.50

1.50–3.00

2.12±0.44

1.25–3.00

For each variable mean ± standard deviation (SD), range, and sample size (N) is given.

T. substituta (n=167): Mean SVL = 51.60 ± 3.64; Range= 46.00–65.50 (Vasconcelos, Santos & Carretero, submitted);

T. raziana (n=10): Mean SVL = 49.30 ± 4.00; Range= 44.00–55.50;

T. caboverdiana (n=11): Mean SVL = 56.70 ± 3.70; Range= 51.50–64

For each variable mean ± standard deviation (SD), range, and sample size (N) is given.

T. maioensis (N = 16): Mean SVL = 60.80 ± 3.70; Range= 52.00–71.00

T. protogigas from Brava (N = 11 vouchers): 135–157 scales around midbody; 13–16 and 21–24 lamellas under the 1st toe and 5th toe, respectively; 38–47 gular scales

T. protogigas from Rombos (N = 4 vouchers): 143–153 scales around midbody; 13–17 and 19–24 lamellas under the 1st toe and 5th toe, respectively; 38–46 gular scales

T. rudis

T. protogigas

Santiago

Fogo

Brava

Males ( N = 9)

Females ( N = 16)

All ( N = 25)

Males N = 1)

Females (N = 1)

All ( N = 2)

Males (N = 16)

Females (N = 11)

All (N = 27)

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

Mean ± SD

Range

SVL

71.78±9.49

56.00–83.00

66.91±5.19

60.00–76.00

68.66±7.25

56.00–83.00

83.00

77.00

80.00 ± 4.24

77.00 – 83.00

65.25 ± 6.49

56.00 – 77.00

61.73 ± 4.00

57.00 – 69.00

63.81 ± 5.79

56.00 – 77.00

TrL

30.18±4.65

23.05–36.12

28.64±4.16

21.32–37.70

29.19±4.31

21.32–37.70

28.39

34.84

31.62 ± 4.56

28.39 – 34.84

28.28 ± 3.28

24.64 – 34.77

27.07 ± 4.40

21.34 – 36.57

27.79 ± 3.74

21.34 – 36.57

8.41±1.10

6.47–10.11

7.14±0.96

5.62–08.85

7.60±1.17

5.62–10.11

8.53

7.77

8.15 ± 0.54

7.77 – 8.53

6.70 ± 1.02

4.97 – 8.57

5.66 ± 0.41

5.18 – 6.48

6.28 ± 0.97

4.97 – 8.57

FLL

25.08±3.17

20.11–28.95

24.23±2.46

18.67–29.14

24.54±2.70

18.67–29.14

28.88

29.54

29.21 ± 0.47

28.88 – 29.54

23.92 ± 2.39

20.44 – 28.14

22.90 ± 2.28

20.74 – 26.54

23.51 ± 2.36

20.44 – 28.14

CFL

16.59±2.26

13.12–19.17

16.03±1.82

11.8–19.54

16.23±1.96

11.80–19.54

18.34

18.97

18.66 ± 0.45

18.34 – 18.97

15.31 ± 1.69

12.32 – 17.78

14.47 ± 2.2

10.54 – 17.85

14.97 ± 1.92

10.54 – 17.85

HLL

30.52±4.67

23.15–38.61

28.99±2.34

24.48–32.11

29.54±3.36

23.15–38.61

41.02

33.86

37.44 ± 5.06

33.86 – 41.02

30.13 ± 2.60

26.6 – 35.73

29.17 ± 1.93

26.31 – 32.82

29.74 ± 2.36

26.31 – 35.73

FFL

17.58±2.36

13.63–21.24

16.59±1.46

13.66–18.69

16.94±1.85

13.63–21.24

22.93

20.54

21.74 ± 1.69

20.54 – 22.93

17.37 ± 1.67

14.68 – 19.99

16.95 ± 1.32

15.53 – 18.96

17.20 ± 1.52

14.68 – 19.99

16.35±1.93

12.68–18.84

15.66±1.31

14.04–18.31

15.91±1.56

12.68–18.84

19.66

17.55

18.61 ± 1.49

17.55 – 19.66

14.98 ± 1.44

12.42 – 17.50

13.90 ± 0.90

12.65 – 15.47

14.54 ± 1.34

12.42 – 17.50

10.14±1.47

7.57–12.35

9.43±0.83

7.90–11.23

9.69±1.13

7.57–12.35

12.52

10.46

11.49 ± 1.46

10.46 – 12.52

9.25 ± 0.87

7.45 – 10.51

8.55 ± 0.45

7.89 – 9.2

8.96 ± 0.8

7.45 – 10.51

4.04±0.32

3.55–4.40

3.87±0.29

3.36–4.28

3.93±0.31

3.36–4.40

3.83

4.15

3.99 ± 0.23

3.83 – 4.15

3.68 ± 0.32

3.14 – 4.19

3.7 ± 0.34

3.23 – 4.12

3.69 ± 0.32

3.14 – 4.19

2.20±0.31

1.91–2.80

2.01±0.22

1.65–2.42

2.08±0.27

1.65–2.80

2.97

2.75

2.86 ± 0.16

2.75 – 2.97

2.23 ± 0.29

1.85 – 2.84

2.12 ± 0.28

1.58 – 2.54

2.19 ± 0.29

1.58 – 2.84

NED

7.17±1.09

5.08–8.59

7.03±0.56

6.24–8.04

7.08±0.77

5.08–8.59

9.09

8.55

8.82 ± 0.38

8.55 – 9.09

6.62 ± 0.7

5.57 – 7.84

6.37 ± 0.44

5.75 – 7.27

6.52 ± 0.61

5.57 – 7.84

SED

8.97±1.33

6.17–10.53

8.85±0.72

7.81–9.98

8.89±0.96

6.17–10.53

11.14

10.77

10.96 ± 0.26

10.77 – 11.14

8.25 ± 0.74

6.81 – 9.51

7.99 ± 0.51

7.12 – 8.88

8.15 ± 0.66

6.81 – 9.51

EED

7.22±1.10

5.19–8.44

6.75±0.59

5.90–7.70

6.92±0.82

5.19–8.44

9.32

7.96

8.64 ± 0.96

7.96 – 9.32

6.23 ± 0.66

5.3 – 7.45

5.79 ± 0.7

4.43 – 7.26

6.05 ± 0.7

4.43 – 7.45

SLS

10.22±0.83

9–12

10.69±0.87

9–12

10.52±0.87

9–12

10.50 ± 0.71

10 – 11

10.06 ± 0.85

9 – 12

9.64 ± 0.92

8 – 11

9.89 ± 0.89

8 – 12

ILS

8.78±0.83

8–10

9.31±0.79

8–11

9.12±0.83

8–11

9.00 ± 0.00

8.19 ± 0.66

7 – 9

7.73 ± 0.65

7 – 9

8 ± 0.68

7 – 9

Lam

11.11±0.92

10–13

10.81±1.17

9–13

10.92±1.08

9–13

10.50 ± 0.71

10 – 11

11.86 ± 0.86

11 – 13

11.82 ± 0.87

11 – 13

11.84 ± 0.85

11 – 13

Trow

14.44±1.33

12-16

14.75±0.93

13–16

14.64±1.08

12–16

12.50 ± 0.71

12 – 13

14.38 ± 0.62

13 –15

14.00 ± 0.77

13 –15

14.22 ± 0.70

13 –15

Tline

18.67±1.87

16–22

18.81±1.60

16–21

18.76±1.67

16–22

20.06 ± 1.57

15 – 21

20.27 ± 0.65

19 – 21

20.15 ± 1.26

15 – 21

Srow

2.58±0.34

1.75–3

2.50±0.45

1.5–3.5

2.53±0.42

1.5–3.5

4.00

3.50

3.75 ± 0.35

3.50 – 4.00

3.78 ± 0.48

3.00 – 4.50

3.95 ± 0.27

3.50 – 4.50

3.85 ± 0.41

3.00 – 4.50

medS

2.64±0.22

2.25–3

2.61±0.36

2–3.25

2.62±0.32

2–3.25

2.50

2.00

2.25 ± 0.35

2.00 – 2.50

2.95 ± 0.44

2.25 – 3.75

2.86 ± 0.41

2.25 – 3.75

2.92 ± 0.42

2.00 – 3.75

Appendix 3. Networks corresponding to cytochrome b sequence variation in endemic Cape

Verde Tarentola geckos (modified from Vasconcelos et al., 2010).

Lines represent a mutational step, dots missing haplotypes and circles haplotypes. The circle

area is proportional to the number of individuals. Dotted circles represent probable ancestral

haplotypes. For correspondences of sample and location codes, see Vasconcelos et al. (2010).

Appendix 4. Marginal probabilities of migration rates (m1 and m2) and time of

divergence (t) between T. bocagei and T. nicolauensis, present in S. Nicolau Island,

obtained by fitting the IM model to the three-locus data (PDC, ACM4, MC1R) set.

Appendix 5. Estimates of genetic differentiation of the PDC, ACM4 and MC1R between ESUs

using Snn test values.

PDC

ACM4

MC1R

Taxa 1

ESU 1

Taxa 2

ESU 2

Snn

P-value

Snn

P-value

Snn

P-value

0.9375

0.0000 ***

0.9306

0.0000 ***

1.00000

0.0000 ***

0.9412

0.0000 ***

0.9524

0.0000 ***

1.00000

0.0000 ***

0.6957

0.0000 ***

0.9635

0.0000 ***

1.00000

0.0000 ***

1.0000

0.0000 ***

0.9167

0.0010 **

1.00000

0.0000 ***

0.6683

0.0470 *

0.9167

0.0000 ***

1.00000

0.0000 ***

0.9643

0.0000 ***

0.9286

0.0000 ***

1.00000

0.0000 ***

0.6487

0.0790 NS

0.9167

0.0000 ***

1.00000

0.0000 ***

Tgg

0.6487

0.2390 NS

0.9231

0.0000 ***

1.00000

0.0000 ***

0.5573

0.0630 NS

0.9333

0.0000 ***

1.00000

0.0000 ***

Tpp

1.0000

0.0000 ***

0.9167

0.0010 **

1.00000

0.0000 ***

Tph

0.9286

0.0000 ***

0.9286

0.0000 ***

1.00000

0.0000 ***

0.9167

0.0010 **

1.00000

0.0000 ***

0.9412

0.0000 ***

0.6334

0.0070 **

0.93750

0.0000 ***

0.9600

0.0000 ***

0.7082

0.0090 **

0.99605

0.0000 ***

0.9833

0.0000 ***

0.5947

0.5860 NS

1.00000

0.0000 ***

0.8667

0.0000 ***

0.6055

0.1410 NS

1.00000

0.0000 ***

0.9306

0.0000 ***

0.5233

0.5240 NS

1.00000

0.0000 ***

0.7778

0.0030 **

0.5527

0.4360 NS

0.57792

0.4860 NS

Tgg

0.8796

0.0020 **

0.6397

0.1200 NS

0.57792

0.3950 NS

0.8990

0.0000 ***

0.5217

0.3480 NS

1.00000

0.0000 ***

Tpp

1.0000

0.0000 ***

0.5947

0.5730 NS

0.57792

0.5230 NS

Tph

0.9000

0.0000 ***

0.5233

0.5110 NS

0.47778

0.6940 NS

0.8750

0.0120 *

0.5947

0.5750 NS

1.00000

0.0000 ***

0.7692

0.0000 ***

0.6590

0.0000 ***

1.00000

0.0000 ***

1.0000

0.0000 ***

0.7409

0.3660 NS

1.00000

0.0000 ***

0.7179

0.0110 *

0.7322

0.0160 *

1.00000

0.0000 ***

0.9667

0.0000 ***

0.6290

0.0860 NS

1.00000

0.0000 ***

0.7069

0.0330 *

0.6871

0.0740 NS

1.00000

0.0000 ***

Tgg

0.7069

0.0240 *

0.7594

0.0120 *

1.00000

0.0000 ***

0.6467

0.0020 **

0.6000

0.1550 NS

1.00000

0.0000 ***

Tpp

1.0000

0.0000 ***

0.7409

0.3400 NS

1.00000

0.0000 ***

Tph

0.9333

0.0000 ***

0.6290

0.0680 NS

1.00000

0.0000 ***

0.7539

0.1230 NS

0.7409

0.3600 NS

1.00000

0.0000 ***

1.0000

0.0000 ***

0.8194

0.4090 NS

1.00000

0.0000 ***

0.8126

0.0010 **

0.8084

0.0300 *

1.00000

0.0000 ***

0.9783

0.0000 ***

0.7154

0.1010 NS

1.00000

0.0000 ***

0.8035

0.0030 **

0.7734

0.1360 NS

1.00000

0.0000 ***

Tgg

0.8035

0.0030 **

0.8280

0.0100 *

1.00000

0.0000 ***

0.7456

0.0010 **

0.6809

0.1200 NS

1.00000

0.0000 ***

Tpp

1.0000

0.0000 ***

0.8194

0.3700 NS

1.00000

0.0000 ***

Tph

0.9565

0.0000 ***

0.7154

0.1170 NS

1.00000

0.0000 ***

0.8413

0.0290 *

0.8194

0.3680 NS

1.00000

0.0000 ***

PDC

ACM4

MC1R!

Taxa 1

ESU 1

Taxa 2

ESU 2

Snn

P-value

Snn

P-value

Snn

P-value

1.0000

0.0040 **

0.4984

0.3720 NS

1.00000

0.0000 ***

0.5887

0.1840 NS

0.97893

0.0000 ***

0.9286

0.0000 ***

0.5405

0.1680 NS

0.99864

0.0000 ***

1.0000

0.0050 **

0.4556

1.0000 NS

1.00000

0.0000 ***

0.4537

1.0000 NS

0.5046

0.4410 NS

1.00000

0.0000 ***

0.9286

0.0000 ***

0.4799

0.3970 NS

0.90346

0.0020 **

Tgg

1.0000

0.0050 **

0.5429

0.4270 NS

1.00000

0.0240 *

1.0000

0.0010 **

1.00000

0.0010 **

Tpp

1.0000

0.0330 *

1.00000

0.0220 *

Tph

0.9167

0.0010 **

1.00000

0.0000 ***

1.0000

0.0230 *

1.00000

0.0210 *

Tgg

0.4537

1.0000 NS

0.5820

0.2360 NS

1.00000

0.0270 *

0.5114

0.4510 NS

0.5737

0.1390 NS

1.00000

0.0000 ***

Tpp

1.0000

0.0040 **

0.4984

0.3770 NS

1.00000

0.0310 *

Tph

0.8286

0.0000 ***

0.5405

0.1730 NS

1.00000

0.0020 **

0.4429

1.0000 NS

0.4984

0.3510 NS

1.00000

0.0250 *

Tgg

0.9286

0.0010 **

0.5844

0.1640 NS

1.00000

0.0270 *

0.9383

0.0010 **

1.00000

0.0020 **

Tpp

0.8333

0.0260 *

1.00000

0.0370 *

Tph

0.8125

0.0000 ***

1.00000

0.0020 **

0.9167

0.0110 *

1.00000

0.0330 *

Tgg

0.4537

1.0000 NS

0.5463

0.4680 NS

0.42500

1.0000 NS

0.4968

0.8070 NS

0.5119

0.3720 NS

1.00000

0.0000 ***

Tpp

1.0000

0.0070 **

0.4556

1.0000 NS

0.42500

1.0000 NS

Tph

0.8452

0.0040 **

0.4799

0.4360 NS

0.52780

0.6000 NS

0.4556

1.0000 NS

0.4556

1.0000 NS

1.00000

0.0300 *

Tgg

0.4968

0.7740 NS

0.6154

0.0420 *

1.00000

0.0000 ***

Tgg

Tpp

1.0000

0.0060 **

0.5429

0.4910 NS

0.42500

1.0000 NS

Tgg

Tph

0.8452

0.0080 **

0.5844

0.1440 NS

0.52780

0.5740 NS

Tgg

0.4556

1.0000 NS

0.5429

0.4310 NS

1.00000

0.0270 *

Tpp

0.9762

0.0010 **

1.00000

0.0000 ***

Tph

0.8549

0.0010 **

1.00000

0.0000 ***

0.5368

1.0000 NS

0.57792

0.5400 NS

Tpp

Tph

0.5030

1.0000 NS

0.44440

1.0000 NS

Tpp

1.0000

0.0260 *

1.00000

0.0310 *

Tph

0.8333

0.0090 **

1.00000

0.0010 **

All results are based on 1,000 permutation tests of 148, 146 and 136 sequences (homozygotes

duplicated), respectively. Analyses were conducted in DNAsp (NS, not significant, P > 0.05; *,

0.01<P<0.05; ** P <0.01). All positions containing missing data were eliminated from the

dataset. There were a total of 392, 431 and 668 positions in each final dataset, respectively.

Tv, T. boavistensis; Tb, T. bocagei; Tf, T. fogoensis; Td, T. darwini; Ts, T. substituta; Tz, T.

raziana; Tc, T. caboverdiana; Tn, T. nicolauensis; Tg, T. gigas; Tr, T. rudis; Tpp, T. protogigas

protogigas; Tph, T. protogigas hartogi; Tm, T. maioensis.

Cabo Verde giant gecko: how many units for conservation?

Article

Full-text available

Jun 2021
AMPHIBIA-REPTILIA

Tarentola gigas (Bocage, 1875) is the largest gecko living in the Cabo Verde Archipelago. It is subdivided into two subspecies, one confined to the Branco Islet, Tarentola gigas brancoensis (Schleich, 1984), and another to the Raso Islet, Tarentola gigas gigas (Bocage, 1875). These islets were classified as Integral Natural Reserves and further studies on the species are needed to outline more assertive conservation measures. Thus, this study aims to integrate for the first time genetic, morphometric and meristic data to test if there are significant differences between these two taxonomical groups that would support the subspecific designation. The results indicated that they are two closely related subspecies, with some visible differences in size and shape, possibly related to diet, habitat conditions or drift. Given the conservation status of the species, this should be further investigated, aiming an adequate management of these two evolutionarily significant units.

The relevance of Evolutionary Significant Units for the conservation of island-restricted reptiles: Tarentola boettgeri bischoffi as a case study

Article

Full-text available

Jun 2024
AMPHIBIA-REPTILIA

Within vertebrates, reptiles are good island colonisers, often leading to considerable levels of intraspecific diversity among populations inhabiting different islands/archipelagos. This study explores the mitochondrial phylogeographic structure of Tarentola boettgeri, a gecko species endemic to the Macaronesian archipelagos of Selvagens and Canary Islands. Our research introduces a novel monophyletic group, comprising the populations from the islands of Selvagem Pequena and Ilhéu de Fora. Furthermore, we confirm the previously identified genetic clusters associated with Selvagem Grande, Gran Canaria and El Hierro. We estimate that the origin of T. boettgeri dates to the upper Miocene (ca. 6.4 Mya), and that the separation of T. boettgeri bischoffi on Selvagem Grande, Selvagem Pequena, and Ilhéu de Fora, occurred ca. 0.5 Mya. The absence of genetic differences between Selvagem Pequena and Ilhéu de Fora suggests recent gene flow or founder events, possibly facilitated by land connections during major glaciations. Conversely, the geographic barriers between Selvagem Grande and Selvagem Pequena likely persisted, preventing genetic admixing. The significant genetic distances observed among all populations underscore the necessity of an integrative taxonomic revision for T. boettgeri. In light of our findings, and with particular consideration of the small population sizes of T. boettgeri bischoffi on Selvagem Pequena and Ilhéu de Fora, we propose that the identified monophyletic groups should be managed as Evolutionarily Significant Units (ESUs). Accordingly, our study highlights the importance of recognizing ESUs in island-restricted reptile populations for targeted conservation efforts, especially given their unique intraspecific diversity and the vulnerability of their habitats.

More cryptic diversity among spiny lizards of the Sceloporus torquatus complex discovered through a multilocus approach

Article

Dec 2023

A phylogenetic reassessment of the Sceloporus torquatus species complex is presented using a multilocus approach. Topological conflicts related to a clade belonging to disjunct southern populations of S. madrensis are resolved, confirming the identity of such populations as distinct from the nominal populations of S. madrensis located 175 kilometres to the north. We describe this southern clade as a new species distributed in the Sierra Madre Oriental based on its phylogenetic distinctiveness and a combination of morphological characters and measurements, increasing the number of recognised species in the S. torquatus complex to seven.

On the identity of west Saharan geckos of the Tarentola ephippiata complex (Squamata: Phyllodactylidae), with comments on an extreme case of syntopy with their close relative T. annularis

Article

Full-text available

Apr 2022

Tarentola geckos have a widespread geographic distribution and occur both in the Palearctic and Afrotropical realms, as well as the Neotropical region. Particularly, across North Africa phenotypically similar and cryptic species can be found, like the west Saharan members of the T. ephippiata complex. However, the taxonomic relationships and phylogeographic patterns of these geckos are not fully understood. Here we show that some specimens of Tarentola geckos from Mauritania and southern Morocco previously identified as T. hoggarensis can actually be assigned to T. panousei, a taxon treated as a synonym until now. Because the corresponding type specimen has apparently been lost, we designate a neotype for T. panousei referring to a suitable specimen from the type locality. Based on a morphological examination of the neotype and comparative material we provide a detailed redescription of T. panousei and evaluation of its diagnostic characters. Moreover, we report on a syntopic occurrence of T. annularis and T. panousei on an isolated acacia tree in the Western Sahara and the (micro) habitat use of both species.

Green matters: Dietary assessment of a reptile community using DNA metabarcoding

Article

Full-text available

Oct 2023

DNA metabarcoding is widely used for diet characterization and is becoming increasingly important for biodiversity conservation, allowing the understanding of trophic networks and community assemblies. However, to our knowledge, few studies have used this approach to investigate trophic interactions for whole communities and none for reptiles. In particular, few studies have examined the diet composition of Saudi Arabian reptiles, and all have used classical methods only. Therefore, in this work, a non-invasive approach using DNA metabarcoding of faecal pellets was implemented to investigate the diet composition of the reptile community of Wadi Ashar, in AlUla County, north-western Saudi Arabia Kingdom. In the overall diet composition of the community, arthropods were present in 90% of the samples, and plants were present in 63%, revealing the unforeseen importance of plants to this community as a secondary, but also a primary dietary item. For some species, this is the first time that plants have been reported in their diet. A significant effect of reptile body size on diet composition was also demonstrated, indicating its strong influence on prey selection and resource partitioning in the community. This study highlights the importance of community assessments and the power of combining these with non-invasive DNA metabarcoding to accurately assess biodiversity and feeding habits, revealing unknown ecological interactions of often neglected groups. This revolutionary tool for conservation and management provided rapid and holistic information at relatively low costs, allowing to inform local authorities about which elements are central to the sustainable management of the Wadi Ashar community.

Reptile monitoring on the natural reserve of Santa Luzia Island

Article

Full-text available

Dec 2022

Herpetological notes from the islands of São Vicente and Santo Antão, Cabo Verde

Article

Full-text available

Aug 2023

Jirí Moravec

RESUMO Este estudo resume a informação faunística e de história natural de um anfíbio e seis espécies de répteis registadas nas ilhas de São Vicente e Santo Antão entre 3-22 de Outubro de 2022. Foi observada uma forte predação de ovos e juvenis de Caretta caretta por cães selvagens na costa nordeste de São Vicente. A osga sinantrópica Hemidactylus mabouia é considerada uma espécie invasora que pode estar a afectar a distribuição da rara espécie endémica do mesmo género, pois ocupa agora uma grande variedade de habitats antropogénicos. A omnívoria foi documentada em Chioninia stangeri. ABSTRACT This study summarizes the faunistic and natural history information for one amphibian and six reptile species recorded on the islands of São Vicente and Santo Antão from 3-22 October 2022. Strong predation of Caretta caretta nests by feral dogs was observed on the northeastern coast of São Vicente. The synanthropic gecko Hemidactylus mabouia is considered an invasive species that may be affecting the distribution of the rare endemic species of the same genus, as it is now occupying widely different types of anthropogenic habitats. Omnivory was documented in Chioninia stangeri.

Macaronesia as a Fruitful Arena for Ecology, Evolution, and Conservation Biology

Article

Full-text available

Nov 2021

Research in Macaronesia has led to substantial advances in ecology, evolution and conservation biology. We review the scientific developments achieved in this region, and outline promising research avenues enhancing conservation. Some of these discoveries indicate that the Macaronesian flora and fauna are composed of rather young lineages, not Tertiary relicts, predominantly of European origin. Macaronesia also seems to be an important source region for back-colonisation of continental fringe regions on both sides of the Atlantic. This group of archipelagos (Azores, Madeira, Selvagens, Canary Islands, and Cabo Verde) has been crucial to learn about the particularities of macroecological patterns and interaction networks on islands, providing evidence for the development of the General Dynamic Model of oceanic island biogeography and subsequent updates. However, in addition to exceptionally high richness of endemic species, Macaronesia is also home to a growing number of threatened species, along with invasive alien plants and animals. Several innovative conservation and management actions are in place to protect its biodiversity from these and other drivers of global change. The Macaronesian Islands are a well-suited field of study for island ecology and evolution research, mostly due to its special geological layout with 40 islands grouped within five archipelagos differing in geological age, climate and isolation. A large amount of data is now available for several groups of organisms on and around many of these islands. However, continued efforts should be made toward compiling new information on their biodiversity, to pursue various fruitful research avenues and develop appropriate conservation management tools.

Between sand, rocks and branches: An integrative taxonomic revision of Angolan Hemidactylus Goldfuss, 1820, with description of four new species

Article

Full-text available

Aug 2021

The taxonomy of Angolan Hemidactylus has recently been revised. However, the lack of fresh material for some groups and regions, has led to the misidentification of some taxa and an underestimation of actual diversity in others. To shed light on the evolutionary history and systematics of Angolan Hemidactylus , we generated a new phylogenetic hypothesis for the group, and updated the taxonomy following an integrative approach. This resulted in the description of four new species ( H. pfindaensis sp. nov. , H. faustus sp. nov. , H. carivoensis sp. nov. and H. cinganji sp. nov. ), the reevaluation of two recently described species ( H. vernayi and H. paivae ) and the synonymization of a recently described species ( H. hannahsabinnae ). We estimate divergence times for these lineages, providing a preliminary interpretation of their speciation process. Moreover, we suggest and outline 13 Angolan Main Biogeographic Units (AMBUs) in the area, defining a new biogeographic context for future works on Angolan herpetofauna. We now recognize eleven Hemidactylus species in Angola, and we provide here a new morphological key for Angolan Hemidactylus to help with identifications and species assignments in this group.

Una parada en la Macaronesia. Sobre la biota de la isla de Sal (Cabo Verde).

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Full-text available

Jan 2016

Lista vermelha para os repteis (Reptilia)

Article

Full-text available

Jan 1996

Herman Schleich

Les types de Gekkonidés (Reptiles, Sauriens) du Muséum national d'Histoire naturelle Catalogue critique

Article

Jan 1990

E. R. Brygoo

A New Statistic for Detecting Genetic Differentiation

Article

Aug 2000
GENETICS

Richard R. Hudson

A new statistic for detecting genetic differentiation of subpopulations is described. The statistic can be calculated when genetic data are collected on individuals sampled from two or more localities. It is assumed that haplotypic data are obtained, either in the form of DNA sequences or data on many tightly linked markers. Using a symmetric island model, and assuming an infinite-sites model of mutation, it is found that the new statistic is as powerful or more powerful than previously proposed statistics for a wide range of parameter values.

Die radiation der gattung Tarentola in makaronesien (Reptilia: Sauria: Gekkonidae)

Article