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Zootaxa 3404: 1-52 (2012)
ISSN 1175-5326 (print edition)
Copyright (c) 2012 * Magnolia Press
ISSN 1175-5334 (online edition)
Revision of the giant geckos of New Caledonia (Reptilia: Diplodactylidae:
AARON M. BAUER1,4, TODD R. JACKMAN1, ROSS A. SADLIER2 & ANTHONY H. WHITAKER3
1Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA.
E-mail: firstname.lastname@example.org; email@example.com
2Department of Herpetology, Australian Museum, 6 College Street, Sydney 2010, New South Wales, Australia.
3Whitaker Consultants, 270 Thorpe-Orinoco Road, Orinoco, R.D. 1, Motueka 7196, New Zealand.
We employed a molecular phylogenetic approach using the mitochondrial ND2 gene and five associated tRNAs
(tryptophan, alanine, asparagine, cysteine, tyrosine) and the nuclear RAG1 gene to investigate relationships within the
diplodactylid geckos of New Caledonia and particularly among the giant geckos, Rhacodactylus, a charismatic group of
lizards that are extremely popular among herpetoculturalists. The current generic allocation of species within New
Caledonian diplodactylids does not adequately reflect their phylogenetic relationships. Bavayia madjo, a high-elevation
endemic is not closely related to other Bavayia or to members of any other genus and is placed in a new genus, Paniegekko
gen. nov. Rhacodactylus is not monophyletic. The small-bodied and highly autapomorphic genus Eurydactylodes is
embedded within Rhacodactylus as sister to R. chahoua. Rhacodactylus ciliatus and R. sarasinorum are sister taxa but are
not part of the same clade as other giant geckos and the generic name Correlophus Guichenot is resurrected for them.
Remaining New Caledonian giant geckos (R. leachianus, R. trachrhynchus, R. auriculatus) receive weak support as a
monophyletic group. Although the monophyly of Rhacodactylus (including Eurydactylodes) exclusive of Correlophus
cannot be rejected, our results support the recognition of a R. chahoua + Eurydactylodes clade separate from
Rhacodactylus sensu stricto. Because of the distinctiveness of Eurydactylodes from R. chahoua (and other New
Caledonian `giant geckos'), we retain this name for the four species to which it has been consistently applied and erect a
new genus, Mniarogekko gen. nov. to accommodate R. chahoua. There is little genetic differentiation within the narrowly
distributed Corrrelophis sarasinorum, but C. ciliatus from southern New Caledonia are both genetically and
morphologically differentiated from a recently discovered Correlophus from the Iles Belep, north of the Grande Terre,
which is here described as C. belepensis sp. nov. Although only subtley different morphologically, the populations of
Mniarogekko from the far northwest of the Grande Terre and from the Iles Belep are strongly differentiated genetically
from M. chahoua populations in the central part of the Grande Terre and are described as M. jalu sp. nov. Rhacodactylus
auriculatus exhibits some genetic substructure across its nearly island-wide range in New Caledonia, but overall
divergence is minimal. Rhacodactylus leachianus exhibits low levels of divergence across its range and southern insular
forms previously assigned to R. l. henkeli are not divergent from southern Grande Terre populations. The few populations
of R. trachyrhynchus sampled are strongly divergent from one another and a specimen from Ilot Moro near the Ile des Pins
is especially distinctive. This specimen and others examined from Ilot Moro are morphologically assignable to the species
described by Boulenger in 1878 as Chameleonurus trachycephalus and is recognized here as a full species. New diagnoses
are provided for each of the eight genera of endemic New Caledonian diplodactylid geckos now recognized. The results
of our study necessitate determinations of the conservation status of the new species described or recognized.
Mniarogekko jalu sp. nov. is considered Endangered, but is locally abundant. Correlophus belepensis sp. nov. is
considered Critically Endangered and is restricted to the ultramafic plateaux of Ile Art. Although described from the Ile
des Pins, we have only been able to confirm the existence of Rhacodactylus trachycephalus on the tiny satellite island Ilot
Moro and consider it to be Critically Endangered. If indeed restricted to this islet, R. trachycephalus may well have the
smallest range and perhaps the smallest population of any gecko in the world.
Key words: Squamata, Rhacodactylus, Correlophus, Mniarogekko gen. nov., Paniegekko gen. nov., Correlophus
belepensis sp. nov., Mniarogekko jalu sp. nov., New Caledonia, molecular phylogenetics, conservation
Accepted by S. Carranza: 28 May 2012; published: 31 Jul. 2012
The biota of New Caledonia is both phylogenetically and ecologically diverse and is noted for its high level of
endemism (Holloway 1979; Chazeau 1993), and the New Caledonian region has been identified as one of the
world's hotspots of tropical biodiversity (Myers 1988, 1990; Mittermeier et al. 1996; Myers et al. 2000; Lowry et
al. 2004). Among terrestrial vertebrates, lizards constitute the most diverse and highly-endemic component of the
fauna (Bauer 1989, 1999; Bauer & Sadlier 2000; Smith et al. 2007). The indigenous lizard fauna is dominated by
lygosomatine skinks and diplodactylid geckos. The best known and perhaps the most distinctive of the New
Caledonian geckos, and among the most noteworthy of all geckos, are the members of the genus Rhacodactylus
Fitzinger, 1843. The genus includes the two largest living species of geckos (Russell & Bauer 1986), the only
viviparous geckos outside of New Zealand (Bartmann & Minuth 1979), and perhaps the most saurophagous of all
geckos (Snyder et al. 2010). While biological data on members of the genus remains limited (Bauer & Sadlier
2000; Henkel 2009; Snyder et al. 2010), all six recognized species are regularly kept in captivity and there exists a
voluminous literature associated with their captive care and breeding (Tytle 1992; Seipp & Henkel 2000, 2011;
Troger 2001; de Vosjoli et al. 2003; Henkel & Schmidt 2007; Cemelli 2009; Schonecker & Schonecker 2009a,
inter alia). On the one hand, the success of these species in captivity and the ease with which at least some species
can be kept and bred has probably decreased demand for wild caught individuals in the pet trade and brought a
global awareness to the uniqueness of these geckos. On the other hand, popular awareness of attractive color
morphs and `varieties' may drive illegal collection of Rhacodactylus, particularly those species that have proven
more difficult to breed in captivity.
Despite being represented by only six species, the genus has had a relatively complex and convoluted
taxonomic history. Perhaps more than most geckos, individual species of Rhacodactylus are highly distinctive and
early workers placed the few species into four genera: Rhacodactylus Fitzinger, 1843, Correlophus Guichenot,
1866, Ceratolophus Bocage, 1873, and Chameleonurus Boulenger, 1878. The generic revision of Boulenger
(1883) stabilized the nomenclature of the group, synonymizing the known forms into five species in a single genus.
In 1913 a sixth species, R. sarasinorum, was described by Roux. Composition of the genus has remained relatively
stable, although two non-nominate subspecies of R. leachianus (Cuvier, 1829), R. l. aubrianus Bocage, 1873 and R.
l. henkeli Seipp & Obst, 1994, and one of the live-bearing R. trachyrhynchus Bocage, 1873, R. t. trachycephalus
(Boulenger, 1878), have been variously recognized by some authors (e.g., Kluge 2001; Seipp & Henkel 2011).
Bauer (1990; Bauer & Henle 1994) recognized the three species of Pseudothecadactylus Brongersma, 1936, a
northern Australian genus, as subgenerically distinct within Rhacodactylus, based on a morphologically-derived
phylogeny. However, subsequent molecular evidence has confirmed that this group is outside the New Caledonian
diplodactylid radiation (Bauer & Jackman 2006) and is probably its immediate sister group (Nielsen et al. 2011); as
such it will not be discussed further here.
Although representative Rhacodactylus have been included in a number of molecular phylogenetic analyses
(e.g., Donnellan et al. 1999; Oliver & Sanders 2009), phylogenetic analyses of the genus as a whole have been
limited. Bauer (1990) and Bauer et al. (1993) using morphological data only, recovered R. auriculatus (Bavay,
1869) as the sister to all remaining species and R. chahoua (Bavay, 1869) and R. ciliatus (Guichenot, 1866) as
sister taxa. The more recent of these analyses placed R. sarasinorum as sister to the chahoua + ciliatus pair, with
leachianus + trachyrhynchus as sister to this clade. Both Bauer (1990) and Good et al. (1997; see also Bauer &
Sadlier 2000), using allozyme data plus morphology, found sarasinorum and trachyrhynchus as sister taxa and
placed leachianus as the sister to the chahoua + ciliatus pair.
In the first analysis based on DNA sequence data, Vences et al. (2001) used a 513 bp fragment of the 16S
mitochondrial gene to elucidate relationships. They found low support for the monophyly of the genus and the only
supraspecific clusters receiving ML bootstrap support of greater than 70% were R. ciliatus + R. sarasinorum (85%)
and this clade + R. chahoua. They also found quite deep divergence between southern mainland + insular
populations of R. leachianus and those on the northern mainland, but little divergence between insular and
mainland R. trachyrhynchus. Patterns of implied species' relationships differed in each of their analyses (neighbor-
joining, maximum parsimony, maximum likelihood). Bauer et al. (2004, 2009) and Bauer and Jackman (2006)
presented preliminary data on relationships of New Caledonian diplodactylids and indicated that data from a
combination of nuclear and mitochondrial genes strongly suggested that Rhacodactylus was made paraphyletic by
Eurydactylodes (not included in the study of Vences et al. 2001), which was found to be the sister to R. chahoua.
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Bauer et al. (2004) also noted that R. sarasinorum and R. ciliatus were strongly supported as sister taxa and that
they found no support for the genetic distinctiveness of R. l. henkeli. Bauer and colleagues, however, did not
publish their explicit trees for New Caledonian diplodactylids at that time.
Thus, each of the previous studies of Rhacodactylus has supported a different pattern of interspecific
relationships, and there has been no agreement even upon the monophyly of the group. We employed a taxon
complete, multi-gene approach with representative intra-specific sampling to evaluate phylogenetic patterns within
Rhacodactylus. Specifically, we investigated 1) the monophyly of Rhacodactylus, 2) the pattern of species-level
relationships, 3) the validity of the subspecies R. l. henkeli and R. t. trachycephalus, and 4) the relationship of
recently discovered disjunct populations resembling R. chahoua, R. ciliatus, and R. auriculatus (Whitaker et al.
2004; Bauer et al. 2006a,b). Of necessity, these objectives required us to reevaluate phylogenetic relationships
among all new Caledonian diplodactylids and our findings have led us to propose a new generic level classification
for this clade.
Materials and methods
Specimens. The majority of specimens examined (Appendix), as well as those from which DNA sequences were
obtained (Table 1), are housed in the collections of the Australian Museum, Sydney (AMS) and the California
Academy of Sciences, San Francisco (CAS and CAS-SU). Additional Rhacodactylus and outgroup specimens
were cited or examined (and in some cases sequenced) from the following collections and institutions: Aaron M.
Bauer collection, Villanova (AMB), American Museum of Natural History, New York (AMNH), The Natural
History Museum, London (BMNH), Monty L. Bean Museum, Brigham Young University, Provo (BYUH), Musee
de l'Ecole de Medecine Navale, Brest [no longer in existence] (EMNB), Field Museum of Natural History,
Chicago (FMNH), Institut Royal des Sciences Naturelles de Belgique, Brussels (IRSNB), Museum of Comparative
Zoology, Harvard University, Cambridge, MA (MCZ), Museum d'Histoire Naturelle, Geneve (MHNG), Museu de
Lisboa, Lisbon [destroyed by fire] (MLI), Musee d'Histoire Naturelle, Marseille (MMNH), Museum National
d'Histoire Naturelle, Paris (MNHN), Museum fur Tierkunde, Senckenberg Naturhistorische Sammlungen, Dresden
(MTKD), Museum of Vertebrate Zoology, University of California, Berkeley (MVZ), Naturhistoriska Riksmuseet,
Goteborg (NHMG), Naturhistorisches Museum Basel (NMBA), Naturhistorisches Museum, Wien (NMW),
Naturalis-Nationaal Natuurhistorisch Museum, Leiden (RMNH), Royal Ontario Museum, Toronto (ROM),
Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main (SMF), University of Michigan Museum of
Zoology, Ann Arbor (UMMZ), United States National Museum of Natural History, Washington, DC (USNM), Yale
Peabody Museum of Natural History, New Haven (YPM), Zoologisches Forschungsmuseum Alexander Koenig,
Bonn (ZFMK), Zoological Institute, Russian Academy of Sciences, St. Petersburg [formerly ZIL] (ZIN),
Zoological Museum Hamburg (ZMH), and Zoologische Sammlung der Bayerischen Staates, Munchen (ZSM).
Morphology. Specimens were examined under a Nikon SMZ 1000 binocular microscope and photographs
were taken with a Canon G11 Powershot digital camera. The following measurements were taken with digital
calipers (to the nearest 0.1 mm): snout-vent length (SVL; from tip of snout to vent), trunk length (TrunkL; distance
from axilla to groin measured from posterior edge of forelimb insertion to anterior edge of hindlimb insertion, with
limbs at right angles to the body axis), forearm length (ForeaL; from base of palm to elbow, with limb partially
flexed); crus length (CrusL; from base of heel to knee, with limb partially flexed); tail length (TailL; from vent to
tip of tail), tail width (TailW; measured at widest point of tail); head length (HeadL; distance between posterior
margin of retroarticular process of jaw and snout-tip), head width (HeadW; maximum width of head), ear length
(EarL; longest dimension of ear); orbital diameter (OrbD; greatest diameter of orbit), naris to eye distance (NarEye;
distance between anteriormost point of eye and posteriormost point of nostril), snout to eye distance (SnEye;
distance between anteriormost point of eye and tip of snout), eye to ear distance (EyeEar; distance from anterior
edge of ear opening to posterior corner of eye), internarial distance (Internar; distance between nares), and
interorbital distance (Interorb; shortest distance between left and right supraciliary scale rows). Unless otherwise
stated, measurements were made on right side of specimens. Number of supralabials (and number to midpoint of
eye) (SupraL), infralabials (InfraL), and lamellae under digits of the manus (LamManus) and pes (LamPes) were
recorded bilaterally. Digital X-ray images of specimens were obtained using a Faxitron closed cabinet X-ray (LX-
60, Faxitron Corp.) with a Varian flat-panel digital X-ray detector.
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Molecular methods. Nucleotide sequences from the mitochondrial ND2 and five flanking tRNAs (tryptophan,
alanine, asparagine, cysteine, tyrosine), and from the nuclear RAG1 genes were obtained from representatives of
all described genera and species of New Caledonian diplodactylid geckos (except the recently described Bavayia
nubila Bauer, Sadlier, Jackman & Shea, 2012, which is the sister species to B. goroensis Bauer, Jackman, Sadlier,
Shea & Whitaker, 2008. In addition, representative New Zealand and Australian diplodactylids, including two
species of Pseudothecadactylus -- the immediate sister group to the New Caledonian clade -- and representatives
of the Carphodactylidae and Pygopodidae were included as outgroup taxa. In total 2286 bp of sequence were
generated for 144 pygopodoid gecko samples including 25 outgroup taxa and 34 taxa of New Caledonian
diplodactylids (Table 1). Genomic DNA was extracted using the Qiagen QIAmp tissue kit and PCR amplification
was conducted under a variety of thermocyler parameters using a diversity of primers (see Nielsen et al. 2011 for
detailed primer information and PCR conditions). Products were visualized via 1.5% agarose gel electrophoresis.
Amplified products were purified either using an AmPure magnetic bead PCR purification kit or reamplified
products were purified on 2.5% acrylamide gels (Maniatis et al., 1982) after being reamplified from 2.5% low-melt
agarose plugs. DNA from acrylamide gels was eluted from the acrylamide passively over two days with Maniatis
elution buffer (Maniatis et al. 1982). Cycle-sequencing reactions were performed using the Applied Biosystems
BigDyeTM primer cycle sequencing ready reaction kit. The resulting products were purified using SeqClean
magnetic bead purification kit. Purified sequencing reactions were analyzed on an ABI 373A stretch gel sequencer
or an ABI 3700 automated sequencer. To ensure accuracy, negative controls were included in every reaction,
complementary strands were sequenced, and sequences were manually aligned using the original chromatograph
data in the program SeqMan II. Sequences have been deposited in GenBank (Table 1).
Phylogenetic methods. Phylogenetic trees were estimated using maximum parsimony (MP), maximum
likelihood (ML) and Bayesian inference (BI). PAUP* 4.0b10a (Swofford 2002) was used to estimate parsimony
trees. Parsimony searches were conducted with 100 heuristic searches using random addition of sequences. Non-
parametric bootstrap resampling was used to assess support for individual nodes using 1000 bootstrap replicates
with ten random addition searches. For maximum likelihood analyses, a partitioned RAXML 7.2.6 (Stamatakis
2006) analysis with the General Time Reversible plus Gamma model was used with a 7 partition analysis (3
mitochondrial codon positions, 3 nuclear codon positions, and 1 tRNA partition). The best tree was estimated from
100 runs and 1000 bootstrap replicates were performed with bootstrap percentages associated with the best
maximum likelihood tree and the branch lengths associated with the best tree. To estimate a phylogenetic tree with
a Bayesian framework MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003) was used with the model chosen using
ModelTest 3.7. (Posada & Crandall 1998) using the seven partition analysis described above. The Bayesian
analyses were initiated from random starting trees and run for 10,000,000 generations with four incrementally-
heated Markov chains. Likelihood parameter values were estimated from the data and initiated using flat priors.
Trees were sampled every 1000 generations. The first 25% of saved trees were discarded as `burn-in' samples.
Stationarity was confirmed both by the standard deviation of split frequencies being below 0.01, and by comparing
the two tree files using Tracer (Rambaut & Drummond 2007) and AWTY (Nylander et al. 2008).
The concatenated tree using all genes, with seven data partitions (codon positions for each gene plus tRNAs) had a
likelihood of -ln 42164.83. An SH test in RAXML that compared the best tree with a monophyly constraint for the
genus Rhacodactylus was significantly different from the optimal tree at p <0.05. The difference in likelihoods was
-ln 44.26, exceeding the standard deviation of the RELL bootstrapped tress by greater than a factor of 2. There
were 1034 variable and 882 parsimony-informative characters for the ND2 analysis (henceforth referring to ND2
plus the five flanking tRNAs) and 1440 variable and 1122 parsimony informative characters in the combined ND2
and RAG1 analysis.
All analyses (ND2 only, RAG1, ND2 + RAG1; MP, ML and BI) found strong support for the monophyly of the
New Caledonian diplodactylids as a group (Figs. 1-2). RAG1 only analyses (not shown) yielded no significant
support for most internal nodes and not all species were recovered with support. All other analyses, however,
retrieved monophyletic Eurydactylodes Wermuth, 1965 and Dierogekko Bauer, Jackman, Sadlier & Whitaker, 2006
with strong support, the latter as sister to the monotypic Oedodera Bauer, Jackman, Sadlier & Whitaker, 2006,
although only with strong support in the Bayesian analyses. Bavayia madjo Bauer, Jones & Sadlier, 2000 was
recovered as the sister to Rhacodactylus sensu lato (exclusive of the species assigned to Correlophus--see below)
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and Eurydactylodes + all remaining Bavayia Roux, 1913 (Fig. 1, as Paniegekko madjo). As such, strong support
for a monophyletic Bavayia was only obtained if B. madjo was excluded. Remaining Bavayia were strongly
supported in the Bayesian analyses (pP = 0.98), but only weakly so under likelihood (69% bootstrap for combined
tree, 53% for ND2 only). Within Bavayia the morphologically well-defined B. cyclura, B. sauvagii, and B. ornata/
septuiclavis groups were retrieved with varying levels of support. None of the analyses found a monophyletic
Rhacodactylus. In all cases Eurydactylodes was embedded inside part of Rhacodactylus as the sister to R. chahoua.
This relationship has posterior probabilities of > 0.98 in the Bayesian analyses and bootstrap support of > 96% in
the ML analyses. In addition, the strongly-supported sister species pair of R. ciliatus and R. sarasinorum were
consistently outside the clade that included their remaining congeners plus Eurydactylodes as sister to a clade
comprising all New Caledonian taxa exclusive of Oedodera + Dierogekko, although with low support. One of the
only conflicts between the ND2 and combined trees is seen in Rhacodactylus. In the ND2 tree Rhacodactylus
trachyrhynchus (including R. trachycephalus) is the sister of R. auriculatus, but with poor support, and was sister
to R. leachianus in the combined analysis, again with poor support. In the combined tree R. auriculatus was sister
to (R. trachyrhynchus + R. leachianus) + (Eurydactylodes + R. chahoua). This pattern received strong support in
the Bayesian analysis, but only moderate bootstrap support under ML and MP. No higher order groupings of
Rhacodactylus species receive support except that the clade including Eurydactylodes plus all Rhacodactylus
exclusive of R. ciliatus and R. sarasinorum is strongly supported under BI (pP = 0.96-1.00) and weakly so in the
likelihood analyses (66-68% bootstraps).
All Rhacodactylus species are monophyletic and levels of intraspecific variation are generally much lower
than interspecific differences. There is virtually no variation across the 10 specimens of R. sarasinorum sampled
and divergences across R. leachianus samples are also relatively small. Rhacodactylus auriculatus exhibits near
uniformity across its continuous range in southern New Caledonia, whereas northern populations are modestly
divergent from one another. Deeper divergences characterize R. trachyrhynchus, R. chahoua, and especially R.
Systematics. Our dataset is dominated by the mitochondrial ND2 gene. Although RAG1 did not recover well-
supported relationships within Rhacodactylus or other New Caledonian genera, its combination with ND2 (Fig. 2)
resulted in topologies that differed somewhat from the ND2 tree (Fig. 1) only with respect to the placement of R.
auriculatus and several species of Dierogekko. We believe that the relatively rapid diversification of the New
Caledonian gecko radiation has not been captured by the slowly evolving nuclear locus. Further, whereas the ND2
topology is strongly supported, the conflicting RAG1 topology is not. We therefore accept the ND2 topology as the
current best estimation of relationships and reevaluate the taxonomy of Rhacodactylus accordingly. Interestingly,
however, some relationships in the combined tree, for example, the monophyly of both Bavayia (exclusive of B.
madjo) and Eurydactylodes, receive substantially higher ML bootstrap support than in the ND2 tree only. The
effect of additional nuclear genes on clade support has been considered by Skipwith (2011).
Taxonomic Implications at the Generic Level. We reject the monophyly of Rhacodactylus both on the
grounds that it is made paraphyletic by its inclusion of Eurydactylodes, and because of the apparent polyphyletic
origin of the six recognized species.
The type species of Rhacodactylus Fitzinger, 1843 by original designation is Ascalabotes leachianus Cuvier,
1829 and the name is therefore linked to this species. That R. ciliatus and R. sarasinorum are sister taxa is
unambiguous and consistent with the findings of Vences et al. (2001) and Bauer et al. (2004). That this clade is also
not part of Rhacodactylus sensu stricto is likewise strongly supported by our analyses. Correlophus Guichenot,
1866, with C. ciliatus Guichenot, 1866 as its type species by monotypy, is the available generic name for this clade
which we here resurrect from the synonymy of Rhacodactylus.
The sister relationship between Eurydactylodes and Rhacodactylus chahoua has been previously noted (Bauer
et al. 2009), although the taxonomic implications of this finding have not yet been addressed. To maintain the
monophyly of Rhacodactylus (exclusive of Correlophus) would require that Eurydactylodes be synonymized with
it. Alternatively, if Eurydactylodes were to be retained this would necessitate the recognition of one or more
additional genera for the giant geckos, depending upon the topology of the reference phylogeny. Either solution
requires some degree of disruption to the existing usage of names, which has been relatively stable for more than a
century (Boulenger 1883; Roux 1913). Although we are opposed to the arbitrary proliferation of generic names,
particularly when monotypic taxa are involved, in this instance we believe that the maintenance of Eurydactylodes
as a separate genus is warranted for this clade of four species that is defined by an extensive suite of morphological
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