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A new species of the hitherto monotypic genus Chalarodon is described from southern Madagascar and a lectotype (ZMB 4360) is designated for C. madagascariensis Peters, 1854. The new species of terrestrial iguana, Chalarodon steinkampi sp. nov., is defined by several morphological characters and by concordant differentiation in mitochondrial and nuclear DNA with >5% uncorrected pairwise genetic distance in the 16S rRNA gene. It can be most clearly recognized by the presence of smooth (vs. keeled) gular and ventral scales, a spotted pattern extending from flanks onto belly, and an unpigmented throat. The new species is known from only a small area between the villages of Amboasary Sud and Esomony, located west of the Andohahela Massif, while C. madagascariensis appears to be widespread over much of southern and western Madagascar. We highlight the need for further exploration of this unprotected region which might host several other microendemic species.
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Accepted by S. Carranza: 4 Mar. 2015; published: 9 Apr. 2015
ZOOTAXA
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Copyright © 2015 Magnolia Press
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http://dx.doi.org/10.11646/zootaxa.3946.2.3
http://zoobank.org/urn:lsid:zoobank.org:pub:09375347-C794-4890-9EDE-8518739B8B01
A likely microendemic new species of terrestrial iguana, genus Chalarodon,
from Madagascar
AURELIEN MIRALLES
1,2,5
, FRANK GLAW
3
, FANOMEZANA M. RATSOAVINA
2,4
& MIGUEL VENCES
2
1
CNRS-UMR5175 CEFE, Centre d’Ecologie Fonctionnelle et Evolutive, 119 route de Mende, 34293 Montpellier cedex 5, France
2
Division of Evolutionary Biology, Zoological Institute, Technical University of Braunschweig, Mendelssohnstr. 4, 38106
Braunschweig, Germany
3
Zoologische Staatssammlung München, Münchhausenstr. 21, 81247 München, Germany
4
Département de Biologie Animale, Université d’Antananarivo, BP 906. Antananarivo 101, Madagascar
5
Corresponding author. E-mail: miralles.skink@gmail.com
Abstract
A new species of the hitherto monotypic genus Chalarodon is described from southern Madagascar and a lectotype (ZMB
4360) is designated for C. madagascariensis Peters, 1854. The new species of terrestrial iguana, Chalarodon steinkampi
sp. nov., is defined by several morphological characters and by concordant differentiation in mitochondrial and nuclear
DNA with >5% uncorrected pairwise genetic distance in the 16S rRNA gene. It can be most clearly recognized by the
presence of smooth (vs. keeled) gular and ventral scales, a spotted pattern extending from flanks onto belly, and an unpig-
mented throat. The new species is known from only a small area between the villages of Amboasary Sud and Esomony,
located west of the Andohahela Massif, while C. madagascariensis appears to be widespread over much of southern and
western Madagascar. We highlight the need for further exploration of this unprotected region which might host several
other microendemic species.
Key words: Taxonomy, Chalarodon steinkampi sp. nov., Iguanidae, Oplurinae
Introduction
Iguanas (Iguanidae sensu lato) are a species-rich radiation of lizards with a peculiar disjunct geographic
distribution. They predominantly occur in the New World, with over 1000 Neotropical species distributed in 11
major clades that are recognized as either families or subfamilies. The Iguanidae also contains two relictual insular
clades: three species of Brachylophus occur in the Pacific Ocean on the Fiji archipelago and neighboring islands
(Keogh et al. 2008; Noonan & Sites 2010) and seven species of the subfamily Oplurinae (with the two genera,
Oplurus Cuvier and Chalarodon Peters) in the Western Indian Ocean region are endemic to Madagascar and the
Comoros archipelago (Blanc 1977).
Although recent time tree analyses provide different divergence times of the Malagasy iguana clade from its
South American sister group (Noonan & Chippindale 2006; Okajima & Kumazawa 2009; Crottini et al. 2012), all
these studies agree in placing the divergence into the Mesozoic. The ancestor of oplurines might have dispersed to
Madagascar via Antarctica and evolved in isolation after these land masses became completely isolated (Noonan &
Chippindale 2006; but see Ali & Aitchison 2009).
Among Malagasy iguanas, the genus Oplurus contains six rock-dwelling, terrestrial or arboreal species, and is
morphologically characterized by rows of enlarged and sometimes spiny scales encircling the tail (Blanc 1977;
Cadle 2003; Münchenberg et al. 2008). The genus Chalarodon is on the contrary monotypic, represented by a
single described species. Chalarodon madagascariensis is a terrestrial species commonly observed on the sandy
soils of forested or bushy areas of the sub-arid to semi-arid southern and western regions of Madagascar (Blanc
1969, 1970). It can be easily differentiated from Oplurus by its smaller size, longitudinal dorsal crest (especially
marked in sexually mature males) running from the occipital region to the mid-length of the tail, and absence of
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enlarged spiny scale rows on tail (Blanc 1977; Glaw & Vences 2007). Intensive earlier work by Petit (1928) and
especially by C. P. Blanc has dealt with the ecology, osteology and reproductive biology of Chalarodon (e.g., Blanc
1965, 1969, 1970, Blanc & Carpenter 1969), and more recently a comprehensive, popular account on the species
was published (Schlüter 2013), but a detailed assessment of its intraspecific variation is so far missing.
The phylogenetic relationships among Malagasy iguanas have been studied by Blanc et al. (1983), Titus &
Frost (1996), Münchenberg et al. (2008), and more recently by Pyron et al. (2013) in a molecular supermatrix
analysis of squamate phylogeny. The molecular phylogeny of Münchenberg et al. (2008) retrieved the two genera
Oplurus and Chalarodon as reciprocally monophyletic, with good support for Chalarodon but weak support for the
genus Oplurus, whereas Pyron et al. (2013) recovered Oplurus as paraphyletic with respect to Chalarodon. Taken
together this suggests the need of additional, comprehensive molecular datasets to test the relationships among
these iguanas, and such data are likely to become available in the near future through next-generation sequencing
methods.
The study of Münchenberg et al. (2008) also included preliminary assessments of phylogeographic pattern in a
number of widespread oplurids, among them C. madagascariensis. Most populations of this species were only
weakly differentiated from each other, but one lineage, exclusively found around the south-eastern locality of
Esomony, was genetically strongly divergent in mitochondrial (16S) as well as nuclear DNA (c-mos) sequences. A
subsequent DNA barcoding study of Malagasy reptiles confirmed strong genetic differentiation of the Esomony
population for the COI gene (Nagy et al. 2012). Because only a few juvenile specimens from this locality were
available for morphological examination, its status could not be further assessed.
To elucidate the status of this divergent lineage of Chalarodon we undertook additional fieldwork in 2010,
focusing on southern Madagascar. Newly generated molecular datasets were combined with those published by
Münchenberg et al. (2008), and a series of morphological characters were scored in the genotyped specimens. We
here follow a congruence approach to species delimitation (Padial et al. 2010) to increase reliability of species
hypotheses and minimize the risk of false positives (Carstens et al. 2013; Miralles & Vences 2013). We analyze
three independent lines of evidence separately (mitochondrial DNA, nuclear DNA and morphology), and based on
the congruent differentiation of the Esomony population we conclude it represents a new species, which is
described herein.
Material and methods
Samples and localities. A total of 27 new tissue samples of Chalarodon were collected in December 2010 in
western and southern Madagascar. For all of them, a piece of tissue was removed and stored in 96% ethanol.
Among these new specimens, 20 have been collected as representative voucher specimens, fixed in 5% formalin or
95% ethanol and stored in 70% ethanol. Twelve of them have been deposited at the Zoologische Staatssammlung
München (ZSM) and 6 at the Université d'Antananarivo, Département de Biologie Animale (UADBA, not
analyzed for morphology). External morphology was examined from a total of 15 preserved specimens; for the
majority of these, molecular data were available as well. All voucher specimens analyzed for morphology are
deposited in the ZSM and the Museum für Naturkunde at Berlin (ZMB). Lists of all voucher specimens used for
morphological and molecular study are included in Table 1. FGZC, FGMV, MV, MIR refer to Frank Glaw, Miguel
Vences and Aurélien Miralles field numbers, TM to Tobias Münchenberg lab numbers.
Morphology. Measurements of specimens were recorded to the nearest 0.1 mm using a dial caliper. Meristic,
mensural and qualitative characters examined include a number of scale counts, coloration characters, and
measurements used routinely in squamate taxonomy as well as several others that we defined after first exploratory
examination of Chalarodon specimens. Abbreviations used for morphological characters are as follows:
Snout-vent length (SVL); maximum head width (HW); head length (HL); forelimb length from axilla to tip of
longest finger (FOL); hindlimb length from groin to tip of longest toe (HIL); number of scales along thigh (THSC);
number of scales along shank (SHSC); number of scales along upper arm (UASC); number of scales along lower
arm (LASC); number of spiny scales in dorsal crest (DCSC); number of scales in contact with mental (NSCM);
distinctly enlarged flat scales behind cloaca (ESBC); number of supraocular scales (SOCU); number of interoculars
scales (INTOC); number of ventral scales from chin fold to cloaca (VEN); number of scales around midbody
(MID); number of scales from eye to snout tip (ESTSC); number of supralabial scales (total number, adding up left
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and right) (SLAB); number of infralabial scales (total number, adding up left and right) (ILAB); number of ventral
scales on longest finger (LFSC); number of ventral scales on longest toe (LTSC); number of dorsal cross bands
(DCB); number of black spots on hindlimb (HILBSP); number of white spots on hindlimb (HILWSP); number of
dark cross bars on throat (TCB). Statistical analysis was done in Statistica 7.1 (Statsoft Inc.). Morphometric
measurements HW, HL, FOL, and HIL were analyzed as size-corrected relative values to SVL (RELHW, RELHL,
RELFOL, RELHIL). Due to the low sample size, data of males and females were merged for analysis. We included
all variables in the analysis despite possible autocorrelation (e.g., between relative length of fore and hind limb)
because our goal, from a taxonomic point of view, was to identify all the variables showing differences between the
two species of Chalarodon.
DNA sequencing and phylogenetic analysis. DNA sequence data were collected for one mitochondrial DNA
(mtDNA) fragment of the 16S rRNA (16S) gene, and for one fragment of nuclear DNA (nDNA) of the oocyte
maturation factor (c-mos). Standard polymerase chain reactions were performed in a final volume of 12.5 μl
containing 0.3 μl each of 10 pmol primer, 0.25 μl of total dNTP 10 mM (Promega), 0.1 μl of 5 U/ml GoTaq, and 2.5
μl of GoTaq Reaction Buffer (Promega). Primers and PCR conditions were as in Münchenberg et al. (2008). The
successfully amplified products were purified using the ExoSAP-IT purification kit according to the
manufacturer’s instructions. Purified PCR templates were sequenced using dye-labeled dideoxy terminator cycle
sequencing on an ABI 3130 automated DNA sequencer. The data matrix was 100 % complete. Sequences were
checked and aligned by eye in CodonCode Aligner (CodonCode Corp.). Due to the low number of indels needed
for alignment of 16S sequences the alignment was unambiguous. All newly determined sequences were deposited
in GenBank under accession numbers KJ170247−KJ170300.
Phylogenetic analysis of the 16S mtDNA alignment by Bayesian inference (BI) was carried out using MrBayes
3.1.2 (Huelsenbeck & Ronquist 2001). The GTR+G+I model of evolution was determined by AIC in MrModeltest
2.3 (Posada & Buckley 2004; Posada & Crandall 1998). We performed two runs of 20 million generations each
(started on random trees) with four incrementally heated Markov chains each (using default heating values),
sampling the Markov chains at intervals of 1000 generations. The convergence of the Markov chains was checked
with Tracer v1.5 (Rambaut & Drummond 2009) and we verified the posterior likelihood values reached stability by
visual inspection and stationarity of the PRSF (Potential Scale Reduction Factor). The first 10 million generations
were conservatively discarded and 10000 trees were retained post burn-in and summed to generate a 50%-majority
rule consensus tree. We used one species of Oplurus (O. cyclurus) as an outgroup.
The PHASE algorithm (Stephens et al. 2001) implemented in DnaSP v5 (Librado & Rozas 2009) was used to
infer haplotypes from the c-mos nuclear DNA sequences. Haplotype networks were reconstructed using statistical
parsimony (Templeton et al. 1992), as implemented in the program TCS v1.21 (Clement et al. 2000) with a
connection limit of 95%. Networks were imported in Adobe Illustrator CS2 (Adobe Systems) to add colors and
connections between haplotypes co-occurring in heterozygote specimens (cf. Flot et al. 2010).
Results
Identity of Chalarodon madagascariensis Peters, 1854 and designation of a lectotype
The original description of Chalarodon madagascariensis by Peters (1854: 616) is extremely short, but in a later
publication the same author provided further details and a figure (Peters 1882: 32) and stated that the original
collection consisted of ten specimens from the Bay of St. Augustin. All of these were of small size although the
species was expected to grow significantly larger. Of the ten syntypes, a minimum of six have been traced by Bauer
et al. (1995: 58): “ZMB 4360 (2 specimens), 5617, 9214 (2 specimens), MNHN 270 (fide Brygoo [1989]);
BMNH?? (fide Boulenger [1885]).” In the meantime, one of the two specimens with the number ZMB 4360 was
renumbered to ZMB 69147, one of the two specimens with the number ZMB 9214 was renumbered to ZMB
69144, and the formerly overlooked skeleton ZMB 13569 was identified as an additional syntype, whereas ZMB
5617 is currently missing from the collection (F. Tillack, pers. comm. 27. Jan. 2015). As indicated by Bauer et al.
(1995) one syntype has been exchanged with the Muséum nationale d'Histoire naturelle at Paris and now bears the
number MNHN 270 (see Brygoo 1989). According to the VertNet database (accessed on 27 Jan. 2015), MNHN-
RA-0.270 was collected by Peters, but the locality of this specimen is given as “Sans localité précise” and its status
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as syntype is not indicated. It is apparently the only Chalarodon specimen present in the MNHN collected by
Peters and therefore should be considered as syntype. One of the syntypes was exchanged with the collection of the
Natural History Museum in London (Boulenger 1885: 128). Altogether, the list of syntypes according to current
knowledge includes eight specimens: (1) ZMB 4360; (2) ZMB 5617 (missing from collection); (3) ZMB 9214; (4)
ZMB 13569 (skeleton); (5) ZMB 69144 (ex 9214); (6) ZMB 69147 (ex ZMB 4360); (7) MNHN 0.270; (8) one
syntype in BMNH (number unknown).
Four of the syntypes (ZMB 4360, ZMB 9214, ZMB 69147, ZMB 69144) were available for our study, and as
was expected from the work of Peters (1882) are all juveniles. The following account refers exclusively to these
four ZMB syntypes examined by us. All are in relatively poor state of preservation, thus only basic data (not
included in table 3) are given. The four specimens are characterized by distinctly keeled ventral and gular scales
and in all of them six scales are in contact with the mental, thereby confirming the differences to the new species
described below. The four specimens are characterized as follows:
(1) ZMB 4360: SVL 42.9 mm, tail 68.3 mm (almost complete), two strongly enlarged flat scales ventrally on
tail base posteriorly to the cloaca, therefore probably a juvenile male.
(2) ZMB 9214: SVL 36.0 mm, tail 18.2 (mutilated), no strongly enlarged flat scales ventrally on tail base
posteriorly to the cloaca, therefore probably a juvenile female.
(3) ZMB 69144: SVL 37.0 mm, tail 34.8 (tail tip mutilated and remaining tail broken into two pieces), no
strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca, therefore probably a juvenile female.
(4) ZMB 69147: SVL 39.0 mm, tail ca. 68.7 mm (tail complete but almost broken at one point), belly ventrally
opened, two strongly enlarged flat scales ventrally on tail base posteriorly to the cloaca, therefore probably a
juvenile male.
Although the four syntypes examined are morphologically uniform, and the entire series was reported to
originate from the same locality, we cannot exclude that any of the six syntypes not examined by us might turn out
to belong to the new species below, or to yet another, still undiscovered taxon. Hence, considering the existence of
at least two Chalarodon species and the unknown fate of several syntypes it appears important for taxonomic
stability to define a single name-bearing type for C. madagascariensis. Following this rationale, we hereby
designate ZMB 4360 as lectotype (Fig. 1). This is the largest of the four studied syntypes, has a nearly complete
tail, a darkened throat, and is in the relatively best state of preservation. It therefore is taxonomically the most
informative specimen. See Figure 1 for a photograph of the lectotype.
Analysis of DNA sequences
The mitochondrial tree based on the 16S dataset retrieved results similar to those published by Münchenberg et al.
(2008), showing a strong differentiation between two well-supported sister clades within Chalarodon (Fig. 2): (i)
the Esomony clade, endemic to the South East of Madagascar (from the previously known locality Esomony, and a
nearby site located 30 km north to Amboasary, on the road towards Esomony); and (ii) a widespread clade
containing all the other populations of Chalarodon sampled over the island of Madagascar, including also a
population from 5 km north of Amboasary and thus just 25 km from a locality of the Esomony clade. Within-clade
variation is low within the widespread clade (uncorrected 16S p-distances of 0.0 to 1.4 %) and absent in the
Esomony clade. On the contrary, genetic distance values between the clades are relatively high (p-distances of 5.4
to 6.5%).
The haplotype network based on the c-mos sequences (Fig. 2) is congruent with the mtDNA analysis. Likewise
it supports two well-differentiated units corresponding to the Esomony clade and the widespread clade in the 16S
tree. These two groups are separated by six mutational steps and do not present any haplotype sharing. All
specimens of the Esomony clade are homozygote for one exclusive c-mos haplotype, whereas four closely related
haplotypes occur in the widespread clade, with adjacent haplotypes always separated by single mutational steps. Of
these, one predominant haplotype is widely distributed across Madagascar and three less common haplotypes were
all identified from the region of Toliara/Ifaty.
It should be noted that one of the c-mos sequences obtained by Münchenberg et al. (2008), GenBank accession
number EU099661 corresponding to specimen ZSM 140/2005 from Esomony (see Table 1), showed three
heterozygous positions suggesting possible hybridization between the Esomony and the widespread clade.
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However, a new DNA extraction, amplification and sequencing of this sample (realised on this sole sample several
months after the initial labwork) produced a strictly homozygote sequence (in both forward and reverse directions;
unambiguous and clean chromatograms) of the typical Esomony haplotype (GenBank accession number
KJ170300). This suggests a very likely PCR crossover contamination (or an editing error during alignment
cleanup, given that ambiguities edited by IUPAC codes might refer to heterozygous positions or simply to poor-
quality reads) in the previous study (Münchenberg et al. 2008), confirming the clear separation of the two clades.
FIGURE 1. Designated lectotype of Chalarodon madagascariensis (ZMB 4360) in (A) dorsal and (B) ventral view.
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FIGURE 2. Molecular results and geographical context: (A) localities of the specimens of Chalarodon madagascariensis and
C. steinkampi sp. nov. included in the molecular analyses. (B) Bayesian phylogenetic tree (see Table 1 for voucher numbers)
based on the 16S mtDNA dataset and (C) haplotype network of the nDNA marker c-mos. Circles represent haplotypes (size
proportional to the number of individuals), black lines represent mutational steps and black dots missing haplotypes, and brown
curves represent connections between haplotypes found co-occurring in heterozygous individuals.
Analysis of morphological data
Although at first glance specimens belonging to the Esomony clade are morphologically similar to those of the
widespread clade, a detailed analysis revealed numerous differences in scale counts, morphometrics, and
coloration.
First, regarding coloration in life or in preservative (Fig. 3), specimens of the Esomony clade can be
recognized by a more extended ventral pattern in the belly region. The pattern consists of a distinct dark-light
spotting and extends from the flanks widely onto the belly except for an unpatterned central band, while in
specimens of the widespread clade, these spots are restricted to the flanks. Furthermore, the throat is blackish in
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specimens of the widespread clade and whitish in the Esomony clade, and dark dorsal crossbands are more
distinctly expressed in most specimens of the Esomony clade.
Second, in body proportions, forelimbs and hindlimbs were relatively shorter in specimens of the Esomony
clade. The difference was statistically significant for relative hindlimb length in non-parametric Mann-Whitney U-
tests (Fig. 4; P=0.019), and for both fore- and hindlimb length it was significant in Tukey's post-hoc tests following
a multivariate ANCOVA (SVL used as covariable), with all variables compared simultaneously and thus corrected
for multiple testing (P=0.037 and 0.003, respectively).
Third, specimens of the two clades differed in a series of scale counts and scale morphologies. The gular and
ventral scales were smooth in specimens of the Esomony clade but always keeled (with 1–3 keels per scale) in
those of the widespread clade. The mental scale contacted four other scales in the Esomony clade, whereas it
contacted 5–8, usually 6 other scales in the widespread clade (variable NSCM, Table 3; significant in U-test,
P=0.0007). Specimens of the Esomony clade had a trend of a lower number of spines in their dorsal crest (Fig. 4D),
a difference significant in U-test (0.019) but not in Tukey's post-hoc test (P=0.098). In addition, a statistically
significant difference was revealed by both tests in the number of ventral scales (U-test, P=0.005; post-hoc,
P=0.015) and of mid-dorsal scale rows (P=0.045 and 0.046, respectively), with specimens of the Esomony clade
having lower scale numbers.
It also is possible that specimens of the Esomony clade reach smaller maximum body sizes than those of the
widespread clade. However, low sample sizes preclude final conclusions on this possible difference.
A further potentially important differential character was observed in the hemipenes (based on those specimens
who had hemipenes everted; see Tab. 2). A distinct, tube-like lobe was visible on the apical part of the hemipenis of
specimens of the Esomony clade, whereas this structure was not recognizable in specimens of the widespread
clade. However, in comparison with the hemipenes presented in Blanc (1977) those of our specimens appear to be
incompletely everted, thus the diagnostic value of this character needs to be re-evaluated when more material
becomes available.
Taxonomic description
Individuals of Chalarodon from the Esomony area show concordant differentiation in mtDNA and nDNA as
compared to specimens from all other sampled localities. These two groups of individuals also differ in
morphology, with some diagnostic differences and others that are statistically significant despite overlapping
values. Possible differences in genital morphology were also noted. Gene flow between the two groups was not
detected, despite their occurrence in close proximity (less than 30 km distance) and without any obvious
geographical barrier between their populations. In fact, vegetation along the road from Amboasary to Esomony is
almost continuous. Strong degradation of original vegetation occurs but according to our observations, Chalarodon
can dwell in partly degraded habitat, suggesting that a direct zone of contact and sympatry might occur in this area.
The congruent results of the three datasets (mtDNA, nDNA, morphology) strongly support the existence of an
independent evolutionary lineage within Chalarodon, meriting recognition as a separate species. The type locality
of Chalarodon madagascariensis Peters, 1854 is the Bay of St. Augustin (i.e., the mouth of the Onilahy river, 35
km to the south to Toliara). The designated lectotype of C. madagascariensis agrees in morphology with
genetically studied specimens of the widespread lineage to which also all our samples from the Toliara region
belong, suggesting that the name C. madagascariensis should be applied to this lineage.
The genus Chalarodon has always been considered as monotypic. It is easily diagnosable and has not been in
the center of attention of taxonomists. No additional binomen has been proposed for this genus after the original
description by Peters in 1854. Duméril & Bibron (1837: 330) described Tropidogaster Blainvillii, a species of
Iguanidae of unknown geographic origin, whose type specimen (MNHN 0.6869; see also Brygoo 1989) was
identified more than a century later as a Chalarodon madagascariensis by Etheridge (1969). This author proposed
to suppress this binomen for purposes of priority and homonymy, and it was subsequently placed on the Official
List of Rejected and Invalid Names in Zoology (Anomymous 1971). Hence, no possible earlier name is available
for the Esomony lineage which therefore is here formally described as new species.
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Chalarodon steinkampi sp. nov.
(Figs. 3, 5, 6)
Holotype. ZSM 835/2010 (field number ZCMV 12909), adult male, collected at a locality 30 km north of
Amboasary Sud along the road to Esomony (-24.7721, 46.4207, 160 m above sea level), on 17 December 2010, by
A. Miralles and F. M. Ratsoavina.
Paratypes. ZSM 836/2010 (ZCMV 12910), adult male, and ZSM 834/2010 (ZCMV 12908), female (possibly
subadult), same locality and collection data as holotype; ZSM 139/2005 (FGZC 2330), adult male, from a site
north of Tranomaro along the road to Esomony (-24.51792, 46.59895, 420 m above sea level), collected by F.
Glaw, M. Vences and P. Bora on 25 January 2005; ZSM 140/2005 (FGZC 2550), adult female, and two specimens
not examined morphologically and deposited in UADBA collection (FGZC 2549 and 2363), all from the
surroundings of Esomony village near crossing of Manambolo river (-24.51762, 46.60405, 420 m above sea level),
collected by F. Glaw, M. Vences and P. Bora on 1 February 2005.
Note. Although the two specimens in the UADBA collection (FGZC 2549 and 2363) have not yet been
catalogued and were not available for morphological examination, we still decided to assign them to the species
(and thereby define them as part of the type series, as paratypes, in agreement with article 72.4.1 of the
International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature1999))
because their identity was unambiguously determined through DNA sequences, and because their paratype status is
in line with general taxonomic practice of distributing types among several collections.
Diagnosis. The new species is a member of the genus Chalarodon, easily distinguished from Oplurus, the only
other genus of iguanids present in Madagascar, by (i) a rather small size, (ii) absence of enlarged or spiny scales
encircling the tail and (iii) the presence of a longitudinal dorsal crest (especially marked in sexually mature males)
running from the occipital region to the mid-length of the tail.
The new species is morphologically differentiated from Chalarodon madagascariensis, the only nominal
species in the genus, by (1) smooth (unkeeled) gular and ventral scales (vs. strongly keeled with 1–3 keels per
scale), (2) mental scale in contact with four scales (vs. with 5–8, usually 6 scales), (3) a spotted pattern extending
from flanks onto belly (vs. this pattern restricted to flanks), (4) an unpigmented throat (vs. mostly pigmented
throat), (5) a distinct, dark dorsal pattern (vs. absent or poorly delimited), (6) on average relatively shorter limbs
(FOL/SVL ratio 0.40−0.43 vs. 0.40−0.49; HIL/SVL ratio 0.68−0.79 vs. 0.74−0.87), (7) on average a lower number
of spines in dorsal crest (81−97 vs. 90−109), and (8) on average a lower number of scale rows around midbody
(144−170 vs. 165−204). Furthermore, C. steinkampi is distinguished from C. madagascariensis by a relevant
differentiation in mtDNA and nDNA, without indications for admixture despite occurrence in close geographical
proximity, without obvious geographical barriers between populations.
Description of the holotype. Adult male in good condition with apparently complete original tail and largely
everted hemipenes. See Table 2 for morphometric measurements and numerical coloration characters, Table 3 for
meristic characters, and Fig. 3 for coloration.
Snout-vent length (56.3 mm) distinctly shorter than tail length (87.4 mm). Head slightly wider than neck.
Snout pointed in dorsal view, rounded in lateral view. Canthus rostralis distinct. Ear opening (auditory meatus)
vertical (height 3.1 mm; width 1.2 mm). Relative finger length: 1≤5<2<4<3, relative toe length 1<5<2<3<4, with
toe 5 deeply separated from toes 1–4.
Rostral scale wider (2.7 mm) than tall (1.0 mm), wider as mental (1.7 mm). Nostrils directed laterally, not in
contact with rostral. Supralabials smooth, with numerous pits; scale rows above supralabials with distinct keels and
pits. Dorsal scales on head very heterogeneous in size, mostly with multiple, long, ornament-like keels. Parietal eye
round, lense-like with shiny blackish surface, 0.5 mm in diameter, surrounded by a single ring-like scales, 1.5 mm
in diameter, and a further ring consisting of 10 scales. Eyelids with numerous small conical tubercles in the
periphery, with one row of enlarged flat scales around eye opening which is surrounded by a row of elevated
enlarged tubercles. Eye opening 2.6 mm in diameter.
One heterogeneous row of distinctly enlarged vertebral scales each with one very distinct keel from the neck
along the back, fading towards the base of the tail. Scales on the back generally small, not imbricated, with one
distinct keel or a conical tubercle, with keels generally getting less distinct towards the flanks. Scales on dorsal
surfaces of forelimbs, hindlimbs and tail distinctly keeled, partly with slight imbrications. Two strongly enlarged
flat scales ventrally on tail base posteriorly to the cloaca. No precloacal pores. Tail base laterally enlarged
anteriorly, nearly round in cross section posteriorly, with sharply pointed tip, without distinct tubercles or spines.
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FIGURE 3. Photographic comparisons of specimens of Chalarodon steinkampi sp. nov. (A: probably ZSM 836/2010, C: ZSM
835/2010, E: ZSM 834/2010, 835/2010, 836/2010) collected 30 km north of Amboasary with specimens of C.
madagascariensis (B, D, F) collected at the closest known neighboring locality, 5 km north of Amboasary. (A–D) Specimens in
life, photographed in the field and (E, F) ventral views of freshly dead specimens; (E) ZSM 834–836/2010, two paratypes and
holotype of C. steinkampi; (F) ZSM 832/2010, ZSM 833/2010, and UADBA (ZCMV 12907).
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FIGURE 4. Boxplots comparing selected morphometric measurements and scale counts among Chalarodon madagascariensis
(left, A, dark brown) and C. steinkampi sp. nov. (right, B, light brown). Plots show relative forelimb lengths (ratios to snout-
vent length; RELFOL), number of dorsal crest scales (DCSC), ventral scales from chin fold to cloaca (VEN), scales around
midbody (MID), number of cross bars on throat (TCB), and number of scales in contact with mental (NSCM). Test statistics are
from non-parametric Kruskal-Wallis comparisons (equivalent to Mann-Whitney U-tests given only two groups are compared).
In addition to these scale counts, most of which are statistically different but show overlap in values between the two species,
diagnostic differences are found in the keeled vs. smooth state of ventrals and gulars (variables VK and GK in Tab. 3).
Ventrally, a distinct skin fold between throat and chest. Ventral scales of chest and abdomen largely flat and
mostly hexagonal without keels, gulars mostly small and rounded, but on the belly larger than scales on the flanks.
Ventral scales on thighs without keels, on shanks ventrally with distinct keels, on feet and tail strongly keeled,
except in the very anterior part of the tail, where scales are only slightly keeled. Subdigitals in single rows. Claws
curving downwards, claws of fourth toe 1.7 mm, claws of third finger 1.0 mm.
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FIGURE 5. Detailed views on mental region(A–B), belly scales (C–D) and hemipenes (E–F) of Chalarodon steinkampi sp.
nov. (ZSM 835/2010) in comparison with C. madagascariensis (ZSM 832/2010). The ulcer-like tubercles on the belly in (C)
were not studied but might be due to unidentified parasites. Note the long distal lobe of the hemipenis of C. steinkampi (F) and
the pair of distinctly enlarged scales posterior to the cloaca of the males (E–F). Photographs by M. Franzen.
Sublabial scales and ca. 3 adjacent scale rows towards the throat with distinct pit-like holes.
Mental triangular rounded, bordered by four scales (two infralabials and posteriorly by a pair of non-elongate,
irregular hexagonal postmentals). Postmentals contact mental, other postmental, first infralabial, two enlarged
lateral gulars, one smaller posterolateral gular and one tiny central gular. First infralabial taller than others. Gulars
small, granular. Hemipenes with distinct long horn-like lobe.
Va ri at io n . See Table 2 for morphometric measurements and numerical chromatic characters. See Table 3 for
meristic characters of the paratypes and Fig. 3 for ventral coloration. The general morphology and coloration of the
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paratypes are similar to the holotype, especially the scalation of the throat and the almost complete absence of
keeled scales on throat and belly.
Etymology. The specific name is dedicated to Martin Steinkamp, in recognition of his support of biodiversity
research and nature conservation through the BIOPAT initiative.
Distribution and natural history. The species is only known from a small area adjacent to the western slopes
of the Andohahela Massif. Three sites are known, all along the road Amboasary-Tranomaro-Esomony, including
the surroundings of Esomony village. All known specimens were active during the day on sandy soils among a
vegetation of xerophytic bushes.
FIGURE 6. Holotype of Chalarodon steinkampi sp. nov. (ZSM 835/2010): (A) head with parietal eye in dorsal view, (B)
dorsal view, (C) ventral view. Photographs by M. Franzen.
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TABLE 1. List of samples and specimens included in the present study, with their respective localities, voucher field numbers, institutional catalogue number (when available) and
GenBank accession numbers (16S and c-mos). Accessions of newly determined sequences are in bold. Where two voucher numbers are given, the second one (in parentheses)
refers to the field number. For collection acronyms, see Materials and Methods. Additional abbreviations: HT, holotype and PT, paratype of C. steinkampi. Specimens analyzed for
morphology are marked "Yes" in column "Morphology". NA, not applicable. An asterisk marks c-mos sequence of one C. steinkampi (ZSM 140/2005) which in the work of
Münchenberg et al. (2008) was determined as heterozygote (hybrid), formerly deposited in GenBank under the accession number EU099661, but was retrieved as homozygote in
this study.
Sample lab
number
Voucher number Locality GPS coordinates Morphology Accession 16S Accession c-mos
C. madagascariensis
Antsalova (TM64) UADBA/FGZC 954 Antsalova -18.6667, 44.6167 - EU099705 EU099654
Betioky 1 (TM75) UADBA/FGZC 335 Betioky -23.7167, 44.3833 - EU099713 EU099662
Betioky 2 (TM76) UADBA/FGZC 336 Betioky -23.7167, 44.3833 - EU099714 EU099663
Isalo (TM85) UADBA 21059 (FGMV 2002.1465) Analalava (Isalo) -21.6917, 44.7833 - EU099706 EU099655
Toliara 1 (TM86) ZSM 946/2003 (FGMV 2002.1551) Toliara -23.35, 43.67 Yes EU099707 EU099656
Toliara 2 (TM87) UADBA (FGMV 2002.2017) Toliara -23.35, 43.67 - EU099708 EU099657
Toliara 3 (TM88) UADBA 21038 (FGMV 2002.2018) Toliara -23.35, 43.67 - EU099709 EU099658
Ifaty 1 UADBA (ZCMV 12797) Ifaty site 1 -23.1266, 43.6340 - KJ170247 KJ170273
Ifaty 2 UADBA (ZCMV 12798) Ifaty site 1 -23.1266, 43.6340 - KJ170248 KJ170274
Ifaty 6 UADBA (MIR 160) Ifaty site 1 -23.1266, 43.6340 - KJ170252 KJ170278
Ifaty 7 UADBA (MIR 187) Ifaty site 2 -23.1241, 43.6496 - NA KJ170279
Ifaty 3 UADBA (ZCMV 12840) Ifaty site 2 -23.1241, 43.6496 - KJ170249 KJ170275
Ifaty 4 UADBA (ZCMV 12839) Ifaty site 2 -23.1241, 43.6496 - KJ170250 KJ170276
Ifaty 5 UADBA (ZCMV 12838) Ifaty site 2 -23.1241, 43.6496 - KJ170251 KJ170277
Tombohina ZSM 828/2010 (ZCMV 12861) Tombohina -23.8673, 44.0877 Yes KJ170253 KJ170280
Beloha ZSM 829/2010 (ZCMV 12865) Road Beloha –
Tsiombe
-25.1375, 45.1550 Yes KJ170254 KJ170281
Beloha UADBA (ZCMV 12866) Road Beloha –
Tsiombe
-25.1375, 45.1550 - KJ170255 KJ170282
Beloha UADBA (ZCMV 12867) Road Beloha –
Tsiombe
-25.1375, 45.1550 - KJ170256 KJ170283
Beloha UADBA (ZCMV 12868) Road Beloha –
Tsiombe
-25.1375, 45.1550 - KJ170257 KJ170284
Faux Cap ZSM 830/2010 (ZCMV 12898) 2 km N Faux Cap -25.55, 45.52 Yes KJ170258 KJ170285
Amboasary 1 ZSM 832/2010 (ZCMV 12905) 5 km N Amboasary -24.9998, 46.3969 Yes KJ170259 KJ170286
Amboasary 2 ZSM 833/2010 (ZCMV 12906) 5 km N Amboasary -24.9998, 46.3969 Yes KJ170260 KJ170287
……continued on the next page
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TABLE 1. (Continued)
Sample lab
number
Voucher number Locality GPS coordinates Morphology Accession 16S Accession c-mos
Amboasary 3 UADBA (ZCMV 12907) 5 km N Amboasary -24.9998, 46.3969 - KJ170261 KJ170288
Petriky 1 UADBA (MIR 223) Petriky forest -25.0484, 46.8629 - KJ170262 KJ170289
Petriky 2 UADBA (MIR 230) Petriky forest -25.0484, 46.8629 - KJ170263 KJ170290
Petriky 3 UADBA (MIR 234) Petriky forest -25.0484, 46.8629 - KJ170264 KJ170291
Petriky 4 UADBA (MIR 233) Petriky forest -25.0484, 46.8629 - KJ170265 KJ170292
Petriky 5 UADBA (MIR 239) Petriky forest -25.0484, 46.8629 - KJ170266 KJ170293
Petriky 6 UADBA (ZCMV 12911) Petriky forest -25.0484, 46.8629 - KJ170267 KJ170294
Petriky 7 ZSM 831/2010 (ZCMV 12912) Petriky forest -25.0484, 46.8629 Yes KJ170268 KJ170295
Petriky 8 UADBA (MIR 227) Petriky forest -25.0484, 46.8629 - KJ170269 KJ170296
NA ZSM 60/1907 Tsimanampetsotsa (N
Mahafaly)
-24.11, 43.84 Yes NA NA
NA ZSM 616/2000 (FGMV 2000.505) Ifaty -23.15, 43.62 Yes NA NA
NA ZSM 422/2002 Toliara -23.35, 43.67 Yes NA NA
C. steinkampi sp.
nov.
Esomony 1
(TM71)
ZSM 139/2005 (FGZC 2330) (PT) N Tranomaro -24.51792, 46.59895 Yes EU099710 EU099659
Esomony 2
(TM72)
UADBA (FGZC 2363) (PT) Esomony -24.51762, 46.60405 - EU099711 EU099660
Esomony 3
(TM73)
UADBA (FGZC 2549) (PT) Esomony -24.51762, 46.60405 - EU099704 EU099653
Esomony 4
(TM74)
ZSM 140/2005 (FGZC 2550) (PT) Esomony -24.51762, 46.60405 Yes EU099712 KJ170300*
Amboasary 4 ZSM 834/2010 (ZCMV 12908) (PT) 30 km N Amboasary -24.7721, 46.4207 Yes KJ170270 KJ170297
Amboasary 5 ZSM 835/2010 (ZCMV 12909) (HT) 30 km N Amboasary -24.7721, 46.4207 Yes KJ170271 KJ170298
Amboasary 6 ZSM 836/2010 (ZCMV 12910) (PT) 30 km N Amboasary -24.7721, 46.4207 Yes KJ170272 KJ170299
Oplurus cyclurus
(outgroup)
TM100 UADBA (FGMV 2002.2093) Ifaty -23.15, 43.62 - EU099749 EU099697
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TABLE 2. Morphometric measurements (in mm) and chromatic characters in specimens of Chalarodon. See Table 1 for locality information and Materials and Methods for
abbreviations. Additional abbreviations NM, not measured; M, male; F, female; SA, subadult, JU, juvenile. HP in second column (Sex) marks males of confirmed sex based on
everted hemipenes.
Voucher
Sex
SVL
HW
HL
FOL
HIL
DCB
HILBSP
HILWSP
TCB
C. madagascariensis
ZSM 60/1907
JU
27.0
6.2
8.0
13.2
23.4
8
NM
8
4
ZSM 828/2010
M
65.1
13.6
15.7
27.0
52.7
10
NM
19
NM
ZSM 829/2010
M
68.3
13.8
17.6
27.4
50.7
8
15
12
NM
ZSM 830/2010
M
63.8
12.6
16.0
30.8
51.6
NM
15
18
3
ZSM 831/2010
M (HP)
59.4
12.7
14.1
27.4
51.4
13
NM
18
NM
ZSM 616/2000
F
57.7
11.6
14.8
NM
49.0
8
9
NM
4
ZSM 946/2003
F (eggs)
54.0
10.9
13.9
24.4
44.8
8
NM
7
4
ZSM 832/2010
M
67.6
13.0
15.8
29.2
51.2
15
8
13
NM
C. steinkampi sp. nov.
ZSM 139/2005
M
59.6
11.7
14.1
25.8
46.4
7
7
13
7
ZSM 140/2005
F
52.4
10.2
13.2
21.9
35.7
8
NM
6
6
ZSM 834/2010
F
46.8
9.3
10.5
19.0
37.2
9
12
9
4
ZSM 835/2010 (holotype)
M (HP)
56.3
11.3
14.2
22.7
41.5
8
26
12
7
ZSM 836/2010
M (HP)
57.0
24.0
14.9
24.1
44.0
9
14
18
5
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TABLE 3. Meristic characters in specimens of Chalarodon. See Table 1 for locality information, Table 2 for sex of specimens, and Materials and Methods for abbreviations.
Additional abbreviations: NM, not measured. Presence of very obvious enlarged scales behind cloaca in column ESBC is emphasized by an exclamation mark. The last two
columns indicate keeled (k) vs. smooth (s) state of ventrals (VK) and gulars (GK).
Voucher NSCM SOCU INTOC DCSC VEN MID ESTSC LTSC LFSC SLAB ILAB THSC SHSC UASC LASC ESBC VK GK
C. madagascariensis
ZSM 60/1907 8 7 20 97 NM NM 12 NM NM 33 33 28 25 24 24 2! k k
ZSM 828/2010 6 9 18 107 88 198 11 32 21 23 23 21 23 24 31 2 k k
ZSM 829/2010 6 9 19 90 88 196 10 36 19 35 37 NM NM NM NM 3 k k
ZSM 830/2010 6 7 18 100 85 191 10 36 19 33 37 28 21 22 23 2 k k
ZSM 831/2010 6 6 25 98 88 180 10 34 20 27 29 20 22 24 17 3 k k
ZSM 616/2000 5 9 14 109 114 200 10 36 18 27 31 26 23 24 18 0 k (k)
ZSM 422/2002 6 9 15 109 102 204 10 36 17 25 35 36 27 42 26 2! k k
ZSM 946/2003 6 7 18 93 NM NM 9 32 21 15 27 NM NM NM NM 0 k k
ZSM 832/2010 6 10 21 100 81 NM 10 28 15 22 33 30 22 22 24 3 k k
ZSM 833/2010 6 8 24 95 76 165 9 34 18 23 27 24 25 25 25 (2) k k
C. steinkampi sp. nov.
ZSM 139/2005 4 10 20 86 82 145 10 36 20 27 33 26 26 25 21 2 s s
ZSM 140/2005 4 8 22 97 85 162 9 28 17 13 17 26 23 25 21 0 s s
ZSM 834/2010 4 8 19 87 75 144 11 32 17 23 35 22 22 28 20 (2) s s
ZSM 835/2010
(holotype)
4 8 23 81 78 170 10 32 20 25 41 24 23 19 19 2 s s
ZSM 836/2010 4 8 17 95 80 164 10 34 22 8 17 26 22 26 21 2 s s
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Discussion
Although oplurine iguanids received substantial scientific attention in recent years (e. g. Randriamahazo & Mori
2001, 2005; Vieira et al. 2007; Okajima & Kumazawa 2009; Dujsebajewa et al. 2009; Hawlitschek et al. 2011;
Chan et al. 2012), the taxonomy of this group remained unchanged for many years. Our description adds a second
species to the previously monotypic genus Chalarodon, and an eighth species to the enigmatic Malagasy clade of
oplurine iguanas. Considering that the last description of an oplurine was that of Oplurus grandidieri in 1900 (but
see Gardner et al. 2011), this discovery is remarkable and highlights that some of the most prominent components
of the Malagasy herpetofauna require taxonomic revision. A similar situation occurs in the giant snakes of the
genera Sanzinia and Acrantophis where a recently described western subspecies of Sanzinia has been elevated to
species rank (Reynolds et al. 2014), and a divergent mitochondrial lineage of Acrantophis of unclarified taxonomy
occurs in arid southern Madagascar (Orozco-terWengel et al. 2008).
Superficially, Chalarodon steinkampi could be considered as a cryptic and poorly differentiated species, given
the overlap of several morphometric and pholidotic characters. However, there are distinct differences in ventral
scalation (being strongly keeled in all C. madagascariensis and unkeeled in C. steinkampi) and the configuration of
the scales on the throat also differ consistently. Both species can therefore be easily distinguished by morphology
alone, and additional distinguishing characters might be detected when more material becomes available for study.
One interesting character, the presence of mostly two distinctly enlarged flat scales behind the cloaca ventrally on
the tail base, is not useful to distinguish between both species, but their presence seems to be a reliable character to
identify a specimen as male, even if it is still juvenile as in ZSM 60/1907. In females these scales are only slightly
enlarged or even not recognizable.
In addition to the distinct morphological characters, the strong genetic differentiation, with absence of any
haplotype sharing in a nuclear gene despite occurrence in close geographical proximity, leaves no doubt about the
species status of these two lineages. In fact, haplotype sharing in c-mos is common among well-established
squamate species, such as various species of day geckos in the genus Phelsuma (Gehring et al. 2013). Its absence
in Chalarodon, despite a rather limited sampling, suggests that no admixture between the two lineages occurs, and
we have no doubt that C. madagascariensis and C. steinkampi will qualify as species also under a biological
species criterion.
Understanding the phylogeography of Chalarodon is difficult with the limited data available. Of the three
localities of C. steinkampi, two are at elevations above 400 m and one is at 160 m above sea level, while C.
madagascariensis is a typical coastal species occurring mostly at very low elevations. Several inland localities are
known, however (Blanc 1977; Glaw & Vences 2007), and it will be important to include these in further molecular
studies. In any case, no obvious autecological difference between the two species of Chalarodon is known. A more
detailed study of the contact zone between them might reveal subtle differences in ecology and behavior.
The distribution area of C. steinkampi is within a poorly explored area of Madagascar. While intensive
herpetological inventories have taken place in the area around Tolagnaro and within the Andohahela reserve (e.g.,
Andreone & Randriamahazo 997; Nussbaum et al. 1999), few studies have targeted the hilly lowland area adjacent
to the western slopes of the Andohahela massif in which the village of Esomony is located. Several anecdotal data
suggest that this area might qualify as a regional center of endemism warranting further exploration: The small
Angavo Massif, not far from this area, is the type locality of an enigmatic species of dwarf gecko, Lygodactylus
decaryi, which is completely unstudied in terms of distribution and natural history (Puente et al. 2009). Besides
Chalarodon, Münchenberg et al. (2008) also found evidence for a limited amount of genetic differentiation of
populations of a second iguanid, Oplurus saxicola, from this area. The populations of the skink Trachylepis vato
from the Esomony area seem to be genetically distinct as well (Lima et al. 2013), and Crottini et al. (2015) have
identified a new gecko species of the genus Paragehyra from the mountain slopes immediately adjacent to
Esomony. Common to all these taxa is that they live in rather dry habitat, and that the divergent species or lineages
are superficially cryptic, recognizable only after molecular analysis or thorough morphological comparison.
Furthermore, all of the taxa with apparently endemic lineages in this area were found in at least somewhat
degraded habitats, suggesting that they are tolerant of a certain degree of habitat alteration. Nevertheless, given that
none of the possibly microendemic species and lineages from this area are known from within any protected area
and the rate of loss of original spiny bush habitat in south-western and southern Madagascar is high, further survey
work is warranted to better understand habitat requirements, taxonomy, and conservation status of these reptiles.
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The new species C. steinkampi likely qualifies for one of the Red List threat categories according to the
classification of the International Union for the Conservation of Nature (IUCN 2001), but given the limited
available data, we here propose to consider its status as Data Deficient (DD).
Acknowledgements
Samples used in this study have been collected in the framework of collaboration agreements among the
Département de Biologie Animale, Université d'Antananarvo, the Direction des Eaux et Forêts, Madagascar, the
Zoologische Staatssammlung München, and the Zoological Institute, TU Braunschweig. We are grateful to the
Malagasy institutions for granting research and export permits. Meike Kondermann and Gabriele Keunecke
assisted with labwork. Morphological data were partly scored by participants of the BD08 Vertebrate Morphology
course at TU Braunschweig, supervised by MV. We are grateful to these students for their help: Lisa Bewersdorf,
Bianca Bobowski, Benjamin Eichhofer, Laura Japke, Christina Kubik, Jutta Pruß, Kim Eileen Rennhack, Hendrik
Reuper, Leonie Salzburger, Felix Scholübbers, Charlotte Tacke, Fanni Thrum, Kathrin Tichy, and Catherine
Tindall. We are indebted to Michael Franzen for providing photographs as well as to Mark-Oliver Rödel and Frank
Tillack (ZMB) for the loan of the Chalarodon specimens under their care and information on the missing syntypes.
Field work in Madagascar was supported by the Volkswagen Foundation, and by a postdoctoral fellowship of the
Humboldt Foundation to AM.
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... E-mail: lukas.kratochvil@natur.cuni.cz Glaw, Ratsoavina & Vences, 2015, was recently described (Miralles et al., 2015). The genus Oplurus includes six species forming two lineages differing in ecology (Blanc, 1977): a rock-dwelling group containing species Oplurus quadrimaculatus Dum eril & Bibron, 1851, Oplurus saxicola Grandidier, 1869, Oplurus fierinensis Grandidier, 1869 and Oplurus grandidieri (Mocquard, 1900), and the more arboreal Oplurus cyclurus (Merrem, 1820) and Oplurus cuvieri Gray, 1831. ...
... E-mail: lukas.kratochvil@natur.cuni.cz Glaw, Ratsoavina & Vences, 2015, was recently described (Miralles et al., 2015). The genus Oplurus includes six species forming two lineages differing in ecology (Blanc, 1977): a rock-dwelling group containing species Oplurus quadrimaculatus Dum eril & Bibron, 1851, Oplurus saxicola Grandidier, 1869, Oplurus fierinensis Grandidier, 1869 and Oplurus grandidieri (Mocquard, 1900), and the more arboreal Oplurus cyclurus (Merrem, 1820) and Oplurus cuvieri Gray, 1831. ...
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