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A molecular phylogeny of the "Madascincus polleni species complex", with description of a new species of scincid lizard from the coastal dune area of northern Madagascar

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The present paper constitutes a study on a taxonomically confusing group of closely related species belonging to the Malagasy skink genus Madascincus, currently encompassing the nominal species M. polleni, M. intermedius and M. stumpffi. Based on combined analyses of mitochondrial and nuclear DNA sequences (ND1 and RAG2 genes, respectively), and morphological examination, we provide evidence for the existence of at least four distinct evolutionary lineages within this complex: Madascincus stumpffi; Madascincus arenicola sp. nov. from northern Madagascar; and two cryptic species morphologically similar to the name-bearing types of M. polleni and M. intermedius. The two latter species, although genetically distinct, appear to be morphologically indistinguishable and their taxonomic status cannot be resolved until a better sampling will be available.
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Accepted by S. Carranza: 25 Feb. 2011; published: 12 May 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2876: 116 (2011)
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1
A molecular phylogeny of the “Madascincus polleni species complex”,
with description of a new species of scincid lizard from the coastal dune
area of northern Madagascar
AURÉLIEN MIRALLES1, JÖRN KÖHLER2, FRANK GLAW3 & MIGUEL VENCES1
1Division of Evolutionary Biology, Zoological Institute, Technical University of Braunschweig, Spielmannstr. 8, 38106 Braunschweig,
Germany. E-mails: miralles.skink@gmail.com, m.vences@tu-bs.de
2Department of Natural History - Zoology, Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany.
E-mail: joern.koehler@hlmd.de
3Zoologische Staatssammlung München, Münchhausenstr. 21, 81247 München, Germany. E-mail: Frank.Glaw@zsm.mwn.de
Abstract
The present paper constitutes a study on a taxonomically confusing group of closely related species belonging to the Ma-
lagasy skink genus Madascincus, currently encompassing the nominal species M. polleni, M. intermedius and M. stumpffi.
Based on combined analyses of mitochondrial and nuclear DNA sequences (ND1 and RAG2 genes, respectively), and
morphological examination, we provide evidence for the existence of at least four distinct evolutionary lineages within
this complex: Madascincus stumpffi; Madascincus arenicola sp. nov. from northern Madagascar; and two cryptic species
morphologically similar to the name-bearing types of M. polleni and M. intermedius. The two latter species, although ge-
netically distinct, appear to be morphologically indistinguishable and their taxonomic status cannot be resolved until a bet-
ter sampling will be available.
Key words: Scincinae, Madascincus, M. arenicola sp. nov., M. intermedius, M. polleni, M. stumpffi, Madagascar, phy-
logeny, morphology, species complex
Introduction
With a number of 1455 known species (Uetz & Hallermann 2010), the family Scincidae (skinks) forms the most
diverse radiation of lizards. Given the remarkable richness of this group and its relatively conserved morphology,
the taxonomy of many speciose (and frequently non-monophyletic) genera is still controversial (e.g. Mausfeld et
al. 2002; Reeder 2003; Carranza et al. 2008; Donnellan et al. 2009; Grismer et al. 2009). Following Australia,
Madagascar constitutes the second most important area of diversification for this family with almost 80 recognized
species, nearly all of them endemic to the island (Glaw & Vences 2007). Three independent radiations of skinks
have reached and diversified in Madagascar: two distinct lineages of the subfamily Lygosominae (genera Trachyle-
pis and Cryptoblepharus), and a third monophyletic group including all Malagasy genera in the Scincinae, namely
Amphiglossus” (paraphyletic), Androngo (most probably a synonym of Pygomeles), Madascincus, Paracontias
(including Cryptoposcincus as synonym), Pseudoacontias, Pygomeles, Sirenoscincus and Voeltzkowia (Whiting et
al. 2004; Schmitz et al. 2005; Crottini et al. 2009). The Scincinae lineage is by far the most speciose and morpho-
logically diverse radiation of Malagasy skinks, with 58 described species currently recognized (Glaw & Vences
2007; Köhler et al. 2009, 2010; Miralles et al., 2011). Notably, during the last ten years (2001-2010) twice more
species have been described in this group than during any other decade (Fig. 1), strongly suggesting that the actual
specific diversity in Malagasy scincines is far from being satisfactorily depicted (see also Andreone & Greer 2002).
Several molecular studies have recently been published on the Malagasy Scincinae (Whiting et al. 2004; Schmitz et
al. 2005; Crottini et al. 2009), but none of these has focused on alpha-taxonomic questions, i.e., molecular species
delimitations and monophyly of species-level lineages. Another difficulty involved in the taxonomic study of this
MIRALLES ET AL.
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group is the often low number of voucher specimens available in collections, particularly of forms with fossorial
and secretive habits.
The present study aims to clarify the confusing taxonomy of a small group (and subclade, according to our
unpublished, more comprehensive molecular surveys) within the genus Madascincus, encompassing until now
only three described species—M. polleni (Grandidier, 1869), M. stumpffi (Boettger, 1882) and M. intermedius
(Boettger, 1913). Challenges for this task are (1) the very high and confusing morphological similarity of the name
bearing types of M. polleni and M. intermedius, (2) the uncertain taxonomic status of M. stumpffi (which has been
considered as subspecies of M. polleni, as distinct species, or as synonym of M. polleni) and (3) the existence of
several populations that are not easily assignable to one of these described species.
The present paper constitutes a first step toward the taxonomic clarification of the taxa of this species complex
by providing (1) an exploratory molecular phylogeny of representatives of numerous populations, and (2) the
description of a recently discovered new species from northern Madagascar, which is morphologically well-differ-
entiated from all described taxa.
FIGURE 1. Number of scincine skink species described per decade in Madagascar. The dark line represents the cumulated
data. Dataset based on the species recognised by Glaw & Vences (2007), with addition of two species subsequently described
by Köhler et al. (2009, 2010).
Material and methods
Taxonomic framework. In the present study, the content of the genus Madascincus is defined following Glaw &
Vences (2007), encompassing M. ankodabensis (Angel, 1930), M. igneocaudatus (Grandidier, 1867), M. macrole-
pis (Boulenger, 1888), M. mouroundavae (Grandidier, 1872), M. melanopleura (Günther, 1877), M. minutus
(Raxworthy & Nussbaum, 1993), M. nanus (Andreone & Greer, 2002), the “Madascincus polleni species com-
plex”, and two candidate species referred to by Glaw & Vences (2007) as Madascincus sp. “vitreus” and Madascin-
cus sp. “baeus”. The monophyly of this genus is supported by previous molecular phylogenetic studies (Whiting et
al. 2004; Schmitz et al. 2005; Crottini et al. 2009), but several species (M. ankodabensis, M. macrolepis, M. minu-
tus, M. nanus) have not yet been included in any molecular analysis. Geographic regions in Madagascar are named
according to Boumans et al. (2007).
Morphological study. The specimens examined (all preserved in 70% ethanol) are deposited in the Muséum
National d’Histoire Naturelle, Paris (MNHN); Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am
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SYSTEMATICS OF THE MADASCINCUS POLLENI COMPLEX
Main (SMF), Université d'Antananarivo, Département de Biologie Animale (UADBA), and Zoologische
Staatssammlung München (ZSM). FGZC, FG/MV, MV, MgF and ZCMV refer to Frank Glaw, Miguel Vences and
Madagascar Frontiers field numbers. Measurements of specimens were recorded to the nearest 0.1 mm using dial
caliper. Meristic, mensural and qualitative characters examined here are routinely used in the taxonomy of Scinci-
dae, such as scale counts, presence or absence of homologous scale fusions or the variability in color patterns.
Scale nomenclature, scale counts, and measurements used in the morphological analyses essentially follow
Andreone & Greer (2002). The frontal scale is considered hourglass-shaped when constricted by first supraocular,
bell-shaped when it is not the case (Greer & Shea 2000). Nuchal scales are defined as enlarged scales of the nape,
occupying transversally the place of two or more rows of dorsal cycloid scale (Miralles 2006). Ranges are given for
each meristic and mensural character, followed by the mean ± the standard deviation, with sample size in parenthe-
ses. For some bilateral characters, the sample size has been noted as the number of sides rather than specimens,
then indicated after the sample size. Drawings were made using Adobe Illustrator CS2 and a WACOM graphic tab-
let CTE-640, after photographs taken through a ZEISS stereomicroscope SteREO Discovery V12.
Molecular procedures. Total genomic DNA was extracted using proteinase K (10 mg/ml) digestion followed
by a standard salt-extraction protocol (Bruford et al. 1992). From the mitochondrial DNA (mtDNA), we amplified
fragments of the ND1 and tRNAMet genes, plus the entire mitochondrial tRNAGln and tRNAIle genes. Addition-
ally, nuclear DNA (nuDNA) has been amplified, the recombination activating gene 2 (RAG2). Standard Poly-
merase chain reactions were performed in a final volume of 12.5 l containing 0.3 l each of 10 pmol primer, 0.25
l of total dNTP 10 mM (Promega), 0.1 l of 5 U/ml GoTaq, and 2.5 l of GoTaq Reaction Buffer (Promega). The
primers ND1 intf2 (5’– AAY CGV GCV CCW TTY GAC CTW ACA GA–3’) and ND1 tmet (5’– TCG GGG TAT
GGG CCC RAR AGC TT–3’) of Schmitz et al. (2005) and Leaché & Reeder (2002) were used to amplify a section
of the ND1 and associated tRNAs genes, with the following PCR cycling procedure: 95°C (2:00); 95°C (0:30),
50°C (0:30), 72°C (1:00) for 40 cycles; 72° (10:00).
TABLE 1. List of specimens, collection and accession numbers of the sequences with their references, and localities (with
elevation above sea level).
Species Voucher Locality Sequences
ND1 RAG2
M. arenicola ZSM 2076/2007 (FGZC 1031) Baie des Dunes (E Ramena), 14 m HQ993074 HQ913902
ZSM 1564/2008 (FGZC 1922) Ampombofofo region (frontier base
camp), 28 m HQ993075 HQ913903
ZSM 1565/2008 (FGZC 1703*) Baie des Sakalava (ca. 5km SE
Ramena), 25 m HQ993076 HQ913904
«M. polleni » ZSM 1570/2008 (FGZC 1917) Ampombofofo region (trapsite 5), 28 m HQ993077 HQ913905
ZSM 242/2004 (FGZC 0474) Montagne des Français, 334 m HQ993078 HQ913906
ZSM 1572/2008 (FGZC 1844) Baie des Sakalava (ca. 5 km SE
Ramena), 28 m HQ993079 HQ913907
ZSM 522/2001
(MV2001-313 /2001-C19) Ankarafantsika (Ampijoroa) HQ993080 HQ913908
ZSM 1563/2008 (FGZC 1827) Ankarana, near Petit Tsingy, 90 m HQ993081 HQ913909
M. stumpffi ZSM 1561/2008 (FGZC 1823) Montagne des Français. HQ993082 HQ913910
ZSM 0032/2005 (ZCMV 2031) Marojejy, "Camp Mantella"
(14°26.260´S, 49°46.533`E), 481 m HQ993083 HQ913911
ZSM 206/2003 Antanambao (Maevatanana, on way to
Manongarivo) HQ993084 HQ913912
Outgroup:
M. melanopleura UADBA collection (ZCMV 2257) Andasibe HQ993073 HQ913901
MIRALLES ET AL.
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FIGURE 2. Schematic drawings of the holotype of Madascincus arenicola sp. nov. (ZSM 1565/2008); (A) dorsal, (B) ventral
and (C) lateral view of the right side of the head, (D) detail of right eye. Scale bars = 1 mm.
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SYSTEMATICS OF THE MADASCINCUS POLLENI COMPLEX
FIGURE 3. Schematic drawings of dorsal and lateral views of the type specimens of (A, B) Madascincus polleni (holotype
MNHN 1895.210); (C, D) M. intermedius (lectotype SMF 16027), and (E, F) M. stumpffi (holotype SMF 16019). Scale bars =
1 mm.
MIRALLES ET AL.
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The RAG2 gene was amplified using the primers PY1-F (5’– CCC TGA GTT TGG ATG CTG TAC TT–3’) and
PY1-R (5’– AAC TGC CTR TTG TCC CCT GGT AT–3’) of Gamble et al. (2008) [(94°C (5:00); 94°C (0:30),
55°C (0:45), 72°C (1:00) for 32 cycles; 72° (5:00)]. The successfully amplified products were purified using Exo-
SAP-IT purification kit according to the manufacturer’s instruction (USB corporation). Purified PCR templates
were sequenced using dye-labeled dideoxy terminator cycle sequencing on an ABI 3130 automated DNA
sequencer. Chromatographs were checked and sequences were edited using CodonCode Aligner (v. 2.0.6, Codon
Code Corporation). All new sequences have been deposited in GenBank (HQ913901 to HQ913912, HQ993073 to
HQ993084, details in Table 1).
Phylogenetic analyses. We conducted maximum parsimony (MP), and partitioned Bayesian inference
searches based on the full dataset, as well as Bayesian inferences on single-gene datasets. We used PAUP* 4.0b10
(Swofford 2002) to perform MP analyses with 100 random addition sequence replicates, equal character weighting,
tree bisection and reconnection (TBR) branch swapping, and gaps coded as missing data. Nodal support was
obtained by bootstrap analyses, with 10,000 replicates, 10 random addition sequences replicates and TBR branch
swapping. For the Bayesian analyses, we determined the models of evolution by AIC in MrModeltest 2.3
(Nylander 2004) which selected the GTR+G model for the ND1 and the GTR+I model for RAG2. We performed
one run of 4 million generations (started on random trees) and four incrementally heated Markov chains (using
default heating values) each, sampling the Markov chains at intervals of 100 generations. The first 2 million gener-
ations were conservatively discarded and the trees corresponding to 2 million generations were retained post burn-
in and summed to generate a majority rule consensus tree.
Results
Morphology
Based on morphological characters, the specimens examined and attributable to the “Madascincus polleni species
complex” can be classified into three main distinct phenotypes, morphologically well differentiated from each
other (see Figures 2–5, Table 2, and the Appendix for the complete list of specimens examined):
1. The "polleni" phenotype (Fig. 3A–D, 5A). This phenotype is characterized by: (a) the presence of postnasal
scales, (b) a relatively small size (maximum snout-vent length (SVL) 76.0 mm), (c) a low number of scales
rows around midbody (24–26), and (d) a bell-shaped frontal scale. Additionally, a pair of dark lateral lines
extends from snout to hindlimbs, relatively large and well defined anteriorly, then becoming one (or two paral-
lel) very thin dashed line posteriorly to forelimbs. Specimens assigned to this phenotype are morphologically
homogeneous, and the name-bearing types of both M. polleni and M. intermedius perfectly fit with all their
characters.
2. The "stumpffi" phenotype (Fig. 3E, F, 5B). This phenotype is characterized by: (a) the presence of postnasal
scales, (b) a relative large size (maximum SVL 114.0 mm), (c) a high number of scales rows around midbody
(30–32), and (d) almost always a hourglass-shaped frontal (except in the population from Marojejy where fron-
tals are bell-shaped). This phenotype can be subdivided into two distinct color morphs: the 'lined morph', with
a two-scale wide pair of dark lateral lines extending from snout to hindlimbs, and the 'uniform morph', only
with a very short dark streak extending from the snout to the ear-opening, progressively fading and disappear-
ing on the neck. The coloration of the M. stumpffi holotype perfectly fits with the 'lined morph'.
3. The "new" phenotype (Fig. 2, 4A–B). This phenotype is characterized by: (a) the absence of postnasal scales,
(b) a medium size (maximum SVL 81.7 mm), (c) a low number of scale rows around midbody (26), and (d) a
bell-shaped frontal. The coloration is characterized by the presence of a pair of two-scale wide dark lateral
lines extending from snout to hindlimbs, well defined all along. This phenotype is very homogeneous, and can-
not be attributed to any described taxon. We therefore consider it as a new species, both on the basis of mor-
phological and molecular results (see below).
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SYSTEMATICS OF THE MADASCINCUS POLLENI COMPLEX
TABLE 2. Comparisons of some characteristics distinguishing the different phenotypes/species within the “Madascincus polleni species complex”. For each character, range,
mean ± standard deviation (SD) and sample size (n; inside parentheses) are given. For some bilateral characters, the sample size has been noted as the number of sides rather
than specimens.
(1)
single asymmetrical enlarged nuchal scale, only present on one side.
Phenotype polleni Madascincus stumpffi Madascincus
arenicola
total Type
M. intermedius
Type
M. polleni
total Type
M. stumpffi
total
N lamellae
under 4th finger
min–max:
mean ± SD:
n sides:
6–9
7.62 ± 0.62
(42)
8 – 9
(2)
7
(2)
6–9
7.34 ± 0.75
(32)
8 – 9
(2)
6–7
6.36 ± 0.50
(11)
N lamellae
under 4th toe
min–max:
mean ± SD:
n sides:
18–23
19.89 ±1.49
(36)
19
(1)
18
(1)
15–20
18.00 ± 1.19
(32)
18
(2)
16–19
17.54 ± 0.77
(13)
N ventral
scale rows
min–max:
mean ± SD:
n:
65–78
71.58 ± 3.90
(24)
76
78
70–88
80.67 ± 3.96
(18)
88
75–80
77.86 ± 1.57
(7)
N paravertebral
scale rows
min–max:
mean ± SD:
n:
65–81
72.65 ± 5.72
(23)
78
80
76–88
82.76 ± 2.97
(17)
85
74–81
79.00 ± 2.31
(7)
N longitudinal
scale
rows at mid–body
min–max:
mean ± SD:
n:
24–26
24.83 ± 1.01
(24)
26
-
-
26
-
-
30-32
31.67 ± 0.77
(18)
32
26
26.0 ± 0
(7)
Enlarged nuchal
scales
one pair:
half a pair
1
:
none:
n:
1 (4.16%)
4 (16.66%)
19 (79.81%)
(24)
x
x
1 (5.55%)
2 (11.11%)
15 (83.33%)
(18)
x
3 (42.9%)
2 (28.6%)
2 (28.6%)
(7)
Postnasal presence (%):
n sides:
93.75%
(48)
yes
yes
94.44%
(36)
yes
0%
(14)
Frontal bell shaped:
hour–glass shaped:
n:
100%
0%
(24)
x
x
47.22%
52.77%
(18)
x
100%
0%
(7)
SVL
min–max:
mean ± SD:
n:
49.0–76.0
60.0 ± 8.80
(26)
66.2
74.0
67.0–114.0
89.57 ± 10.76
(14)
91.0
61.5–81,7
72.30 ± 6.05
(7)
MIRALLES ET AL.
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Remark. Two specimens (MNHN 2862, 2862A) cannot be reliably assigned to any of the above mentioned
phenotypes as they share a mosaic of characters of the “polleni” and the “stumpffi” phenotypes. Both specimens are
highly similar to the “polleni” phenotype in terms of size, shape, coloration and presence of a bell-shaped frontal,
whereas they share two scalation characteristics with the "stumpffi" phenotype, namely a high number of paraverte-
bral scale rows (respectively 84 and 82, see Table 2) and a high number of scale rows at mid-body (32 in both spe-
cimens). Both specimens have been collected by the same collector (Charles Coquerel) and likely at the same
locality, which however, is only imprecisely given as “Madagascar”. For these reasons, we discarded these two
enigmatic specimens from the rest of the study.
Molecular phylogeny
The combined analysis of the mitochondrial (ND1) and nuclear (RAG2) genes constitutes a matrix of 885 charac-
ters (503 and 382 for each gene, respectively); 151 sites were variable (129 and 22, respectively) and 84 of them
were parsimony-informative (76 and 8, respectively). The results of the phylogenetic analyses are summarized in
Fig. 7. The heuristic search using MP analysis produced six equally most-parsimonious trees (n tax = 12; tree
length = 254; CI = 0.709; RI = 0.643; RC = 0.455). The two approaches (MP and Bayesian combined analyses; Fig
7C) and the two single gene data-set (mtDNA versus nDNA genes, Figs. 7A, 7B) support the same overall phylo-
genetic topology, without any incongruence. Both MP and Bayesian approaches support (1) the existence of two
distinct and not directly related groups (clades 1 and 2) sharing the same "polleni" phenotype, (2) the monophyly of
the "new" phenotype which we describe below as Madascincus arenicola, and (3) the branching of M. arenicola as
sister species of the clade 1 of the "polleni" phenotype. The monophyly of specimens of the "stumpffi" phenotype is
weakly supported by the Bayesian inference, but not supported nor rejected by the MP approach, and the basalmost
relationships are not resolved.
Description of a new species
Madascincus arenicola sp. nov.
Holotype. ZSM 1565/2008 (field number FGZC 1703), adult male, collected at Baie des Sakalava (ca. 5 km SE
Ramena), 12°16'34'' S, 49°23'24'' E, 25 m above sea level, Antsiranana Province, northern Madagascar, collected
on 20 February 2008 by S. Megson.
Paratypes (8 specimens). ZSM 1564/2008 (FGZC 1922), Ampombofofo region (Frontier base camp),
12°05'53'' S, 49°19'49'' E, 28 m a.s.l., collected on 10 June 2006 by S. Megson; ZSM 2076/2007 (FGZC 1031),
UADBA uncatalogued (FGZC 1029, 1030), Baie des Dunes (E Ramena), 12°14'43'' S, 49°22'53'' E, 14 m a.s.l.,
collected on 23 February 2007 by P. Bora, H. Enting, F. Glaw, A. Knoll and J. Köhler; ZSM 1566–1569/2008
(FGZC 1743, 1744, 1767, 1797), Baie des Sakalava (ca. 5 km SE Ramena), 12°16'34'' S, 49°23'24'' E, 25 m a.s.l.,
collected from 21 to 23 February 2008 by S. Megson. All localities within Antsiranana Province, northern Mada-
gascar.
Diagnosis. As a member of the Malagasy scincine lineage, Madascincus arenicola differs from the other
Malagasy lygosomine genera (Cryptoblepharus and Trachylepis) by the presence of entirely scaly movable eyelids
(versus fused immovable eyelids forming a spectacle over the eye in Cryptoblepharus; or movable eyelids with a
translucent disk or window in the lower eyelid in Trachylepis), absence of prefrontal scales (present in both Cryp-
toblepharus and Trachylepis), and absence of frontoparietal scales (present in Trachylepis).
Within the Malagasy Scincinae, Madascincus arenicola is easily distinguishable from all other species by the
combination of the following characters: presence of two pairs of pentadactyl legs, light bronze dorsal coloration
with two dark well-defined lateral stripes, and absence of postnasals. The new species is assigned to the genus
Madascincus based on its molecular phylogenetic relationships.
Within the genus Madascincus, the absence of postnasal scales is exclusively associated to M. arenicola. In
addition to this highly discriminant character, M. arenicola is distinguished from all its congeners by the following
combination of characters: SVL ranging from 61.5 to 81.7 mm (versus a maximum SVL of 29 mm in M. nanus,
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SYSTEMATICS OF THE MADASCINCUS POLLENI COMPLEX
33.5 mm in M. macrolepis, 39 mm in M. minutus, 50 mm in M. ankodabensis, 57 mm in M. melanopleura); 74–81
rows of paravertebral scales (versus 51–64 in M. melanopleura, 60–68 in M. mouroundavae, and 48–57 in M.
nanus); 75–80 rows of ventral scales (versus 48–54 in M. minutus, 56–63 in M. melanopleura, 59–63 in M. nanus,
60 in M. macrolepis, 61 in M. ankodabensis, 63–67 in M. mouroundavae; relatively long toes, with 16–19 (most
often 17 or 18) subdigital lamellae under the fourth (versus 5–6 in M. macrolepis, 6–9 in M. nanus, 10–11 in M.
minutus, 13 in M. ankodabensis, 9–16 in M. melanopleura); 26 rows of scales around midbody (versus 18 in M.
macrolepis, 20 in M. nanus, 20–22 in M. minutus, 22 in M. ankodabensis, 28–32 in M. mouroundavae, 30–32 in
specimens of the "stumpffi" phenotype); presence of two well defined two-scales wide dark longitudinal stripes on
the flank (versus six dark stripes in M. igneocaudatus, dorsum uniformly dark, without lateral stripes in M. minutus
and M. nanus; pentadactyl forelimbs (versus 4–5 digits in M. nanus and 3–4 in M. sp. "baeus"); and more generally
by the very characteristic snout shape and its slender body.
FIGURE 4. Paratype specimen of Madascincus arenicola sp. nov. (ZSM 2076/2007) in life. (A) dorsolateral view, (B) detail
of anterior body and head.
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FIGURE 5. Comparison of the (A) Madascincus "polleni" phenotype (Ankarana, clade 2) and (B) Madascincus stumpffi (uni-
form color morph from Forêt dAmbre).
Additionally, M. arenicola is also distinguishable from the other species of the “polleni” species complex by its
contrasted coloration, characterized by the presence of a pair of two-scale wide dark lateral lines extending from
snout to hindlimbs, well defined all along the body (versus lateral lines well defined anteriorly, becoming one – or
two parallel – very thin dashed line posteriorly to forelimbs in M. polleni and in the lined morph of M. stumpffi, and
lateral lines reduced to very short dark streak extending from the snout to the ear-opening, progressively fading and
disappearing on the neck in the uniform morph of M. stumpffi), and by the characteristic shape of its snout, being
relatively long and acute in lateral aspect (shorter snout in M. polleni, and in lateral aspect more rounded both in M.
polleni and M. stumpffi).
Description of the holotype. ZSM 1565/2008 (Fig. 2). Good state of preservation, with exception of a little
circular sampling incision on the right flank (± 4 mm of diameter). Adult male, a mature white testis being recog-
nizable inside the abdominal cavity, via the incision). SVL (71.0 mm) 8.2 times head length (8.7 mm), almost as
long as tail (69.9 mm, apparently entire and not regenerated). Limbs remarkably short: SVL 9.4–11.2 times front
limb length (6.4–7.6 mm) and 4.7–4.8 times hind limb lenght (15.1–14.9 mm). Snout relatively long and acute on
lateral aspect, with a rostral tip bluntly rounded in dorsal aspect. Rostral wider than high / long, contacting first
supralabials, nasals, and supranasals. Paired supranasals in median contact, contacting loreals. Frontonasal roughly
triangular, wider than long, contacting loreals, first supraciliaries and first suproculars. Prefrontals absent. Frontal
approximately as wide as long, wider posteriorly, in contact with frontonasal, supraoculars, parietals and interpari-
etal. Supraoculars four, all of them in contact with frontal; subequal in size, except the posteriormost pair signifi-
cantly smaller. Frontoparietals absent. Interparietal longer than wide, well separated from supraoculars; parietal
eyespot present with parietal eye evident. Parietals contact posterior to interparietal. Absence of enlarged nuchals.
Zootaxa 2876 © 2011 Magnolia Press · 11
SYSTEMATICS OF THE MADASCINCUS POLLENI COMPLEX
Nasals just slightly larger than nostrils; contacting rostral, first supralabials and supranasals. Postnasals absent,
seemingly fused with the first supralabials. Loreal single, about as high as long. Preocular trapezoidal, longer than
high, single. Presubocular roughly square, single. Six supraciliaries on the right sight, seven on the left side, in con-
tinuous row; first and last pairs significantly larger and longer than the intermediate ones; last pair projecting onto
supraocular shelf. Upper palpebrals small except for last which projects dorsomedially. Pretemporals two, both
contacted by parietal. Postsuboculars two; upper contacting lower pretemporal; both contacting penultimate
supralabial. Lower eyelid moveable, scaly; lower palpebrals small, longer than high, interdigitating with large
polygonal scales of central eyelid. Contact between upper palpebrals and supraciliaries seemingly direct but flexi-
ble, i.e. palpebral cleft narrow. Primary temporal single. Secondary temporals two; upper long, contacting lower
pretemporal anteriorly and overlapping lower secondary temporal ventrally. Two tertiary temporals bordering
lower secondary temporal. Supralabials six, with the fourth being the enlarged subocular contacting scales of lower
eyelid. Postsupralabial single. External ear opening relatively small, without lobules. Mental wider than long, pos-
terior margin convex. Postmental wider than long, contacting first two pairs of infralabials. Infralabials six. Three
pairs of large chin scales, both members of first and second pairs separated by the same single median scale, and
members of third pair separated by three scale rows. No scales extending between infralabials and large chin
scales; two asymmetrical postgenials posterolaterally in contact with the third pair of chin scales. Gulars similar in
size and outline to ventrals. All scales, except head shields and scales on palms, soles, and digits, cycloid, smooth,
and imbricate; longitudinal scale rows at mid-body 26; paravertebrals 80, similar in size to adjacent scales; ventrals
79. Inner preanals larger than outer. Both pairs of limbs pentadactyl; fingers and toes very short, clawed; relative
length of toes in the following order: I<II=V<III<IV. Subdigital lamellae smooth, single, with 7 under fourth finger
(on each side), 17 under right fourth toe, and 18 under left fourth toe.
In preservative, a pair of lateral dark brown stripes (about two scales wide at midbody) separating a bronze
dorsal field (about eight scales wide at midbody) from the whitish ventral field (about 14 scales wide at midbody),
extending from the loreals, around eye (except the eyelids that are whitish), above ear opening, above forelimb and
hindlimbs, then progressively breaking up into a succession of little dots along tail. Dorsal sides of forelimbs,
hindlimbs and tail dark chocolate brown. Light bronze dorsal field extending from dorsal side of head and neck to
dorsum; laterally lighter on two scale wide along the lateral dark brown stripes. The bronze dorsal field is covered
by numerous little dark dots, each one of them in the middle of a dorsal scale, in contact with its posterior edge;
resulting in four (anteriorly) to six (posteriorly) thin dash lines, darker and more contrasted in the posterior part of
the dorsum, becoming progressively indistinguishable from the dark chocolate-brown background coloration of
tail. Immaculate whitish ventral field extending from lower side of head, throat, lower side of limbs and venter, to
the ventral side of tail. Palms and soles darker than venter, slightly pigmented. The life coloration of the holotype is
unknown. According to the photograph of the paratype specimen ZSM 2075/2007 (Fig. 4A, B), the coloration is
warmer and brighter than after fixation: the dorsal field tends to be orange; under the black lateral stripes, the
lateral side of the ventral field are pale orange colored; and most of the dark parts of the body, especially the dorsal
sides of the tail and limbs, as well as the edges of the black lateral stripes, show a subtle purplish tint. A pinkish tint
is recognizable on the cream colored throat. The two median rows of dorsal scales are slightly lighter than the
adjacent scales. Venter uniformly cream. Eyes totally black.
Va ri at io n . The variation among the ZSM type specimens is as follows: number of lamellae under fourth fin-
ger: 6–7 (6.36 ± 0.50; 11 sides); number of lamellae under fourth toe: 16–19 (17.54 ± 0.77; 13 sides). Number of
ventral scale rows: 75–80 (77.86 ± 1.57; 7); number of paravertebrals scale rows: 74–81 (79.00 ± 2.31; 7); number
of longitudinal scale rows at mid-body: 26 (100%, n = 7). Number of supraciliaries (n = 14 sides): six (92.9%),
seven (7.1%). Three specimens have a pair of enlarged nuchal scales (42.9%), two specimens have a single nuchal
(one on the left side, the other on the right side; 28.6%), two specimens have no nuchal scale (28.6%). All studied
specimen have six supralabials on each sides, the fourth being the enlarged subocular (n = 14 sides); all have supra-
nasals in median contact, parietals in median contact, and are devoid of postnasals (n = 7). Head length: 8.7–9.6
mm (9.15 ± 0.29; 7); snout-vent length: 61.5–81.7 mm (72.30 ± 6.05; 7); tail length: 58.0–71.0 mm (65.90 ± 5.23;
7); forelimb length: 5.32–7.59 mm (6.72 ± 0.63; 10 sides); hindlimb length: 14.9–17.8 mm (15.76 ± 0.93; 13 sides).
SVL/tail length ratio: 6.81–8.47; SVL/head length ratio: 0.94–1.36; SVL/forelimb length ratio: 8.86–13.28; SVL/
hindlimb length ratio: 3.94–4.92. There is no noteworthy variation of coloration within the material examined.
Etymology. The specific epithet is a composite of the Latin arena (sand) and incola (inhabitant). It refers to the
sand-dwelling habits of the species and is used as an invariable noun in apposition.
MIRALLES ET AL.
12 · Zootaxa 2876 © 2011 Magnolia Press
Natural history. All specimens of the type series were captured with pitfall traps and drift fences over night on
sandy soils within disturbed secondary forest or shrub. We were not able to observe active specimens during the
day and therefore suspect a mainly fossorial or sub-fossorial habit during daytime, whereas the species must be
active on the surface at least at night as demonstrated by the pitfall captures. Captured individuals had an excellent
ability to quickly entrench themselves into the sand. Madascincus arenicola occurs in sympatry with other sand-
dwelling fossorial squamates, namely Paracontias minimus, P. rothschildi, P. fasi ka and Xenotyphlops grandidieri
(see Megson et al. 2009). Furthermore, it occurs together with another species of Madascincus, here called pheno-
type “polleni" (corresponding to clade 1, see above).
Distribution. Only known from the Baie des Sakalava and Baie des Dunes in the Forêt d’Orangea area, and
north of the Antsiranana bay in the Ampombofofo region, northern Madagascar (Fig. 6C).
FIGURE 6. Distribution maps of the different taxa forming the "Madascincus polleni species complex". Colored circles repre-
sent localities with voucher specimens and DNA sequences included in the present phylogenetic analyses, whereas black dots
represent collection vouchers specimens that have been only used for the morphological study (see appendix for the exact local-
ities).
Discussion
Our results provide conclusive evidence that the systematics of the “polleni” species complex is more complicated
than expected. Molecular results support the existence of four main evolutionary units within this group, and we
here consider these as four distinct species. Their specific distinctiveness is strongly supported by significant mor-
phological divergence, most of the forms being easily distinguishable from the others. The one exception is the pair
of apparently unrelated clades 1 and 2 that share the same “polleni” phenotype and that we consider as cryptic spe-
cies given that we were unable to distinguish them on the basis of morphology. In addition to the molecular and
morphological results, the specific distinctiveness of the four clades is also confirmed by several cases of sympatry
observed between specimens of the "stumpffi" phenotype and the clade 1 of the "polleni" phenotype (Montagne des
Francais) and between M. arenicola and the clade 1 of the "polleni" phenotype (both in Baie de Sakalava and
Ampombofofo).
Type locality
of M. polleni
Type locality
of M. intermedius
Type locality
Madascincus stumpffi Phenotype “polleniM. arenicola sp. n.
5 km Type
locality
2
1
lined ( ) and ( )uniform
12
clades and
A B C
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SYSTEMATICS OF THE MADASCINCUS POLLENI COMPLEX
FIGURE 7. Phylogenetic trees of the “Madascincus polleni species complex”: Bayesian analyses inferred from single gene
data-set of (A) nuclear DNA (RAG2 gene) and (B) mitochondrial DNA (ND1 gene), and (C) Bayesian tree inferred from the
combined analysis of both ND1 and RAG2, with posterior probabilities followed by the bootstrap support values >50% from
Maximum Parsimony analysis.
MIRALLES ET AL.
14 · Zootaxa 2876 © 2011 Magnolia Press
The fact that two distinct evolutionary species share the same phenotype (here called "polleni" phenotype) pre-
vent us from clarifying their taxonomic status at this time. Indeed, clade 1 is present in the extreme North region of
Madagascar, whereas clade 2 is located in the North West of the island, relatively close to the type locality of M.
intermedius. Moreover, the type locality of M. polleni is located in the Central West of Madagascar, but unfortu-
nately no sample from this area has been available to us (Fig. 6B). As a conclusion, two sound taxonomic hypothe-
ses may be formulated: (I) If clade 2 is conspecific with the nominal M. polleni from the south, then M. intermedius
is probably a junior synonym of M. polleni, and the clade 2 should be called M. polleni. Additionally, the northern
clade (number 1) should be considered as a distinct undescribed species. (II) If clade 2 and the nominal M. polleni
from the south represent two distinct species, then clade 2 should be named M. intermedius, and the northern clade
(1) should be (again) considered as a new species. Moreover, a third hypothesis according to which the nominal M.
polleni would be conspecific with the clade 1 has to be mentioned as generally possible. Nevertheless, we consider
this third hypothesis as highly improbable, given that the second species (clade 2) occupies a range geographically
intermediate between these two distant populations.
The individuals sharing the "stumpffi" phenotype presently constitute a monophyletic unit. Even if the basal
node of this clade is not strongly supported (Fig. 7), its morphological distinctiveness prompt us to follow
Andreone & Greer (2002) in considering it as a distinct species, rather than as a subspecies (Brygoo 1980) or a syn-
onym (Glaw & Vences 2007) of M. polleni. Two different color morphs (lined versus uniform) are present within
this taxon, but there is no certainty whether these patterns reliably identify two separate lineages or not. Indeed, in
the present molecular study, the uniform morph is represented by only two specimens (that form a well supported
clade) whereas the lined morph is represented by a single specimen only, branching with poor support as a sister
taxon to the previous clade (of uniformly colored specimens).
As a conclusion, given the current evidence, we consider within the "Madascincus polleni species complex"
three valid species: Madascincus polleni, M. stumpffi, and M. arenicola, but the exact definition of M. polleni and
M. stumpffi requires further revisionary work. Our first attempt to resolve the taxonomy of this species complex has
revealed the existence of Madascincus arenicola, a new species probably endemic to the north of Madagascar. It
also provided evidence for the need to focus future sampling efforts on the relatively widespread “polleni” pheno-
type, as this nominal taxon is composed by at least two distinct species that cannot be distinguished morphologi-
cally, despite that they are not sister species. Additionally, a study focusing on the chromatic variation observed
within Madascincus stumpffi (lined versus uniform morphs) may be of interest, in order to determine, if both
morphs represent two different evolutionary units, or if they only reflect very local or seasonal patterns of color-
ation, without phylogenetic meaning.
Acknowledgments
We are grateful to Annemarie Ohler and Ivan Ineich (MNHN) and Gunther Köhler (SMF) for providing us access
to specimens under their care. Parfait Bora, Neil D'Cruze, Steven Megson, Angelika Knoll, Hildegard Enting,
Michael Franzen, Zoltan Nagy, Angelin and Angeluc Razafimanantsoa were of invaluable help in the field. The
work was carried out in collaboration with the Département de Biologie Animale, Université d’Antananarivo, and
the Association Nationale pour la Gestion des Aires Protégées (ANGAP). We are grateful to the Malagasy authori-
ties, in particular the Ministère de l’Environnement et des Eaux et Forêts, for research and export permits. Field-
work was made possible by financial support of the European Association of Zoos and Aquaria (EAZA). AM was
supported by a postdoctoral Research Fellowship from the Alexander von Humboldt Foundation and by a SYN-
THESYS grant (FR–TAF–842).
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Appendix. Additional specimens examined
Phenotype "polleni" (including the type specimens of Madascincus polleni and M. intermedius)
Madagascar: Antsiranana province: ZSM 242/2004 (FGZC 474), 245/2004 (FGZC 480), Montagne des Français, 334 m a.s.l.,
coll. on 23-24.2.2004 by Glaw, Puente and Randrianiana; ZSM 1571/2008 (FGZC 1766), 1572/2008 (FGZC 1844), Baie
des Sakalava (ca. 5 km SE Ramena), 28 m a.s.l., coll. on 22./26.2.2008 by Megson; ZSM 1573–1577/2008 (FGZC 1680,
1678, 1836, 1838, 1687), Montagne des Français (pitfall lines 1 & 5), coll. on 19./25.02.2008 by D'Cruze and native col-
lectors; ZSM 259/2004, Montagne des Français, coll. on 18.-28.2.2004 by Glaw, Puente, Randrianiaina and Razafimanant-
soa; ZSM 1570/2008 (FGZC 1917), Ampombofofo region (trapsite 5), 28 m a.s.l., coll. on 23.02.2007 by Mason; ZSM
1562–1563/2008 (FGZC 1658, 1827), Ankarana, near Petit Tsingy, 90 m a.s.l., coll. on 16./24.02.2008 by Franzen, Glaw,
Köhler and Nagy; MNHN 1897.31, Diego Suarez; MNHN 1980.1169, Bemanevika, Plateau Bealanana. Mahajanga prov-
ince: SMF 16027 (lectotype of Scelotes intermedius Boettger, 1913), NW Madagascar, “Majunga” (= Mahajanga), coll. in
1897 by Voeltzkow; ZSM 522/2001 (MV 2001-313), Ankarafantsika (Ampijoroa), coll. on 24.02.2001 by Vences, Vieites,
Garcia, Raherisoa and Rasoamamonjinirina; ZSM 523–524/2001, Ankarafantsika, coll. 2.2001 by Vences, Vieites, Garcia,
Raherisoa and Rasoamamonjinirina. Toliara province: MNHN 1895.210 (holotype of Gongylus polleni Grandidier, 1869),
Moroundava” (= Morondava), coll. by Grandidier; MNHN 1980.1201, coll. by Domergue; MNHN 1984.168, Beroboka
MNHN 1984.169, Tuléar (= Toliara); MNHN 1905.132, plain of the Fiherena, coll. 1869 by Geay.
Phenotype "stumpffi" - uniform morph
Madagascar: Antsiranana province: ZSM 2162/2007 (FGZC 1228), Forêt d'Ambre, ca. 4 km WSW Sakaramy, 470 m a.s.l.,
coll. on 12.3.2007 by Bora, Glaw and Köhler; ZSM 1559/2008 (MgF 061), Montagne des Français, ca. 1.8 km SW Anda-
vakoera ("trapsite 1"), 200 m a.s.l., coll. on 01.12.2006 by Randriamalala; ZSM 1558/2008 (FGZC 3124), Forêt d'Ambre,
ca. 5 km SW Sakaramy (trapsite), 479 m a.s.l., coll. on 19.02.2008 by D'Cruze; ZSM 1560–1561/2008 (FGZC 1772,
1823), Montagne des Français (pitfalls), coll. on 22.-24.02.2008 by native collectors; ZSM 0041/2005 (FGZC 2785),
0032/2005 (ZCMV 2031), 0033/2005 (ZCMV 2032), 0040/2005 (FGZC 2784), 0405/2005 (ZCMV 2034), 0404/2005
(ZCMV 2033), 0043/2005 (FGZC 2787), Marojejy, "Camp Mantella" (14°26.260´S, 49°46.533`E), 481 m a.s.l., coll on
16.02.2005 by Glaw, Vences and Randrianiaina.
Phenotype "stumpffi" - lined morph (including the holotype)
Madagascar: Antsiranana province: SMF 16019 (holotype of Gonglylus stumpffi Boettger, 1882 and lectotype of Scelotes
astrolabi boettgeri Angel, 1941), “Nossibé”, coll. in 1881 by Reuter and Stumpff; MNHN 1884.587, Nossi-Bé, coll. by
Deyrolles. Mahajanga province: ZSM 206/2003 (FG/MV 2002-2268), 5 km from Antambao, close to Maevatanana, road
to Manongarivo, coll. on 31.01.2003, by Glaw, Randrianiaina, and Vences. Toamasina province: MNHN 1980.1200,
Mananara, coll. in 1963 by Peyrieras.
Undetermined material
Madagascar: MNHN 2862, 2862A, coll. by Coquerel in “Madagascar” (no precise locality).
... Of these, fewer than 20 species are grouped within the subfamily Lygosominae (Lima et al., 2013), and these are all largely terrestrial, and generalized in morphology. In contrast, the evolution of a fossorial lifestyle and other ecomorphological adaptations is more prominent in a second clade of skinks, endemic to Madagascar, the Comoros and Glorioso Island, and containing about 60 species; this clade belongs into the poorly defined ''Scincinae" subfamily (Schmitz et al., 2005;Glaw and Vences, 2007;Miralles et al., 2011a). ...
... Their wide-ranging morphological variation makes Malagasy scincines an interesting model system to study macroevolutionary transitions in body form, but due to their often secretive fossorial lifestyle and highly confined distribution range they remain one of Madagascar's most poorly studied vertebrate clades. Nevertheless, as a result of intensive surveys conducted during recent years, new species continue to be discovered (Andreone and Greer, 2002;Sakata and Hikida, 2003a,b;Köhler et al., 2009Köhler et al., , 2010Miralles et al., 2011aMiralles et al., -c, 2012Miralles et al., , 2016a, and further insights in their biology appear on a steady basis (e.g. Andreone, 2004;Rakotoarison et al., 2015). ...
Article
Among the endemic biota of Madagascar, skinks are a diverse radiation of lizards that exhibit a striking ecomorphological variation, and could provide an interesting system to study body-form evolution in squamate reptiles. We provide a new phylogenetic hypothesis for Malagasy skinks of the subfamily Scincinae based on an extended molecular dataset comprising 8060 bp from three mitochondrial and nine nuclear loci. Our analysis also maximizes taxon sampling of the genus Amphiglossus by including 16 out of 25 nominal species. Additionally, we examined whether the molecular phylogenetic patterns coincide with morphological differentiation in the species currently assigned to this genus. Various methods of inference recover a mostly strongly supported phylogeny with three main clades of Amphiglossus. However, relationships among these three clades and the limb-reduced genera Grandidierina, Voeltzkowia and Pygomeles remain uncertain, mainly based on maximum likelihood and maximum parsimony estimates. Supported by a variety of morphological differences (predominantly related to the degree of body elongation), but considering the remaining phylogenetic uncertainty, we propose a redefinition of Amphiglossus into three different genera (Amphiglossus sensu stricto, Flexiseps new genus, and Brachyseps new genus) to remove the non-monophyly of Amphiglossus sensu lato and to facilitate future studies on this fascinating group of lizards.
... Subsequently, diversification led to species colonizing the dry lowlands of the western, southern and northern parts of the island, where the other quadrupedal Malagasy scincine skinks (genus Amphiglossus sensu stricto, and two new genera which will be described by Erens et al. in press) are absent or significantly under-represented (Glaw and Vences 2007). More specifically, the northern part of the island represents a center of diversification for the M. polleni species complex (sensu Miralles et al. 2011a) with a diversification trend oriented toward the north. Similar biogeographical patterns are also observed in the sister lineage of Madascincus (genus Paracontias, Miralles et al. 2016) and in several other squamates (Brown et al. 2014(Brown et al. , 2016. ...
... For some bilateral characters, the sample size was noted as the number of sides rather than specimens. Data fromAndreone and Greer 2002, Glaw and Vences 2007, Miralles et al. 2011a, Miralles and Vences 2013 ...
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In a previous study, Miralles & Vences (2013) compared seven different methods of species delimitation applied to the genus Madascincus. While focusing on methodological aspects their study involved an extensive data set of multilocus DNA sequences and of comparative morphology. On this basis they emphasized the need of revising the taxonomy of Madascincus, and revealed the existence of at least two well-supported candidate species. The present paper provides formal descriptions of these two taxa: (1) M. miafina sp. n., a species from dry areas of northern Madagascar, morphologically very similar to M. polleni (although both species are not retrieved as sister taxa), and (2) M. pyrurus sp. n., a montane species occurring >1500 m above sea level, endemic to the central highlands of Madagascar (Ibity and Itremo Massifs). Phylogenetically, M. pyrurus is the sister species of M. igneocaudatus, a taxon restricted to the dry littoral regions of the south and south-west of Madagascar in lowlands <500 m above sea level. To facilitate future taxonomic work, we furthermore elaborated an identification key for species of Madascincus. Finally, some aspects of the biogeographic patterns characterising the different main clades within the genus Madascincus are provided and discussed for the first time in the light of a robust phylogenetic framework.
... Here we focus on the diversification of the species-rich Malagasy scincine lizards (subfamily Scincinae). This radiation endemic to Madagascar and neighbouring islands (see 2.1 Samples and dataset) has recently been the object of intensive systematic and taxonomic research, with several species described over the last three decades (e.g., Raxworthy and Nussbaum, 1993;Andreone and Greer, 2002;Köhler et al., 2009;Miralles et al., 2011aMiralles et al., , 2011bMiralles et al., , 2011cMiralles et al., , 2012Miralles et al., , 2016aMiralles et al., , 2016b, and the knowledge of their phylogenetic relationships that has greatly improved (e.g., Crottini et al., 2009;Erens et al., 2017;Köhler et al., 2010;Miralles et al., 2015;Miralles and Vences, 2013;Schmitz et al., 2005;Whiting et al., 2004). Their richness and variety emphasise Madagascar's status as a global hotspot for skink diversity (Chapple et al., 2021). ...
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Most of the unique and diverse vertebrate fauna that inhabits Madagascar derives from in situ diversification from colonisers that reached this continental island through overseas dispersal. The endemic Malagasy Scincinae lizards are amongst the most species-rich squamate groups on the island. They colonised all bioclimatic zones and display many ecomorphological adaptations to a fossorial (burrowing) lifestyle. Here we propose a new phylogenetic hypothesis for their diversification based on the largest taxon sampling so far compiled for this group. We estimated divergence times and investigated several aspects of their diversification (diversification rate, body size and fossorial lifestyle evolution, and biogeography). We found that diversification rate was constant throughout most of the evolutionary history of the group, but decreased over the last 6–4 million years and independently from body size and fossorial lifestyle evolution. Fossoriality has evolved from fully quadrupedal ancestors at least five times independently, which demonstrates that even complex morphological syndromes – in this case involving traits such as limb regression, body elongation, modification of cephalic scalation, depigmentation, and eyes and ear-opening regression – can evolve repeatedly and independently given enough time and eco-evolutionary advantages. Initial diversification of the group likely occurred in forests, and the divergence of sand-swimmer genera around 20 Ma appears linked to a period of aridification. Our results show that the large phenotypic variability of Malagasy Scincinae has not influenced diversification rate and that their rich species diversity results from a constant accumulation of lineages through time. By compiling large geographic and trait-related datasets together with the computation of a new time tree for the group, our study contributes important insights on the diversification of Malagasy vertebrates.
... This resulted in several taxonomic revisions (mostly at the genus level) and in a remarkable number of new or resurrected amphibian and reptile species [e.g. Aglyptodactylus (Köhler et al., 2015), Boophis , Blommersia , Gephyromantis , Guibemantis (Lehtinen et al., 2011), Mantidactylus (Bora et al., 2011), Scaphiophryne (Raselimanana et al., 2014), Anodontyla (Vences et al., 2010a), Cophyla (Rakotoarison et al., 2015), Platypelis , Rhombophryne (Scherz et al., 2016); Stumpffia (Rakotoarison et al., 2017), Brookesia , Furcifer (Florio et al., 2012), Calumma , Chalarodon , Zonosaurus (Raselimanana et al., 2006), Madascincus (Miralles et al., 2011), Paracontias (Miralles et al., 2016), Paragehyra , Uroplatus , Phelsuma , Liopholidophis ]. ...
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Prior herpetological surveys in 1996 and 2000 identified 14 species of amphibians and 32 species of reptiles from the Sahamalaza Peninsula. This work increases the total number of amphibian and reptile species known from this area to 20 and 43 respectively. To maximise our chances of species detection, survey effort covered the entire wet season and part of the dry season, and utilised a combination of opportunistic searching, transect searching, pitfall trapping, and acoustic recording. We identified species through an integrative taxonomic approach, combining morphological, bio-acoustic and molecular taxonomy. Together, this enabled the detection of cryptic and seasonally inactive species that were missed in the shorter prior surveys that relied on morphological identification alone. The taxonomic identification of amphibians utilised a fragment of the mitochondrial 16S rRNA gene; taxonomic identification of reptiles utilised a fragment of the mitochondrial COI gene, and when necessary, also mitochondrial fragments of the 16S rRNA ND1, ND2, ND4 genes. All sequences were deposited in Genbank and COI sequences were also deposited in the BOLD database to foster taxonomic identification of malagasy reptiles. We report two new taxa: a species of Boophis, since described as B. ankarafensis, and a candidate new species of microhylid (ge-nus: Stumpffia). We document range expansions of Boophis tsilo-maro, Cophyla berara, Blaesodactylus ambonihazo beyond their type localities. Along with significant range expansions across a range of taxa, including Blommersia sp. Ca05, Boophys brachy-chir, Brookesia minima, Ebenavia inunguis, Geckolepis humblo-ti, Madascincus stumpffi, Pelomedus subrufa and Phelsuma ko-chi. Forest in the peninsula is under extreme pressure from human exploitation. Unless unsustainable agricultural and pastoral practices encroaching on these habitats halt immediately, both forest and the species that occur there, several of which appear to be local endemics, may be irreversibly lost.
... Interestingly, one partly fossorial Madascincus (M. arenicola (ii)), has a slightly simplified cephalic scalation (Miralles et al. 2011b) with absent postnasals (apparently fused with the first loreal (1+PN), based on size and position homology). We here propose that a similar event, likely convergently, might have led to the absence of postnasals in Paracontias. ...
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In this study, we provide an extended multilocus phylogenetic analysis combining mitochondrial and nuclear DNA of a group of fossorial and miniaturized legless lizards (genus Paracontias) from Madagascar, including the description of two species new to science, P. ampijoroensis sp. nov. and P. mahamavo sp. nov. Our analyses revealed the existence of two distinct, parapatric and diagnosable clades within the genus: (i) the ‘kankana clade’ (including P. kankana and the two newly described species), located in the north (but absent from the extreme northern tip) of the island and characterized by a pattern of cephalic scales very unusual for Malagasy Scincinae, with large loreal scales extending to and meeting each other at dorsal midline, and (ii) the ‘brocchii clade’ (including all other studied species), endemic to the north of Madagascar and characterized by small loreal scales separated from each other by the rostral and the frontonasal scale. By combining phylogenetic results with morphological traits observed among species, we develop novel hypotheses on the simplification of the cephalic scalation pattern within this genus, a trend frequently encountered among various lineages of legless squamates that convergently adapted to a burrowing lifestyle. Additionally, a user-friendly graphical identification key for species of Paracontias is provided and made available as supplementary information.
... Meristic, mensural and qualitative characters examined here are those routinely used in the taxonomy of Scincidae [24]. See Miralles et al. [7], [8], [25], [26] for details on the scale nomenclature used. ...
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Scincine lizards in Madagascar form an endemic clade of about 60 species exhibiting a variety of ecomorphological adaptations. Several subclades have adapted to burrowing and convergently regressed their limbs and eyes, resulting in a variety of partial and completely limbless morphologies among extant taxa. However, patterns of limb regression in these taxa have not been studied in detail. Here we fill this gap in knowledge by providing a phylo-genetic analysis of DNA sequences of three mitochondrial and four nuclear gene fragments in an extended sampling of Malagasy skinks, and microtomographic analyses of osteology of various burrowing taxa adapted to sand substrate. Based on our data we propose to (i) consider Sirenoscincus Sakata & Hikida, 2003, as junior synonym of Voeltzkowia Boettger, 1893; (ii) resurrect the genus name Grandidierina Mocquard, 1894, for four species previously included in Voeltzkowia; and (iii) consider Androngo Brygoo, 1982, as junior synonym of Pygomeles Grandidier, 1867. By supporting the clade consisting of the limbless Voeltzko-wia mira and the forelimb-only taxa V. mobydickand V. yamagishii, our data indicate that full regression of limbs and eyes occurred in parallel twice in the genus Voeltzkowia (as hitherto defined) that we consider as a sand-swimming ecomorph: in the Voeltzkowia clade sensu stricto the regression first affected the hindlimbs and subsequently the forelimbs, whereas the Grandidierina clade first regressed the forelimbs and subsequently the hindlimbs following the pattern prevalent in squamates. Timetree reconstructions for the Malagasy Scinci-dae contain a substantial amount of uncertainty due to the absence of suitable primary fossil calibrations. However, our preliminary reconstructions suggest rapid limb regression in Malagasy scincids with an estimated maximal duration of 6 MYr for a complete regression in Paracontias, and 4 and 8 MYr respectively for complete regression of forelimbs in Grandi-dierina and hindlimbs in Voeltzkowia.
... Regrettably, most of these phenomena, such as regression of the eyes, closure of the ear opening, miniaturisation, loss of pigmentation and high degree of cephalic scale reduction (cf. Gans 1974Gans , 1975Lee 1998;Sakata & Hikida 2003a;Miralles et al. 2011a) remain under-studied in comparison to works dealing with limb regression. It would nevertheless be essential to know more about these features and the communication system of fossorial skinks. ...
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The “forelimbs only” bauplan, characterised by the combined presence of well-developed fingered forelimbs and the complete absence of hindlimbs, is rare among terrestrial tetrapods. It is restricted to three lineages of squamates with elongated worm-like bodies, the amphisbaenian genus Bipes Lacépède, 1788 and the scincid genera Sirenoscincus Sakata & Hikida, 2003 and Jarujinia Chan-ard, Makchai & Cota, 2011. In the present study, we describe a new species of Sirenoscincus from Marosely, Port Bergé region, northwest Madagascar, which presents a remarkable variation of this bauplan. The forelimbs of S. mobydick n. sp. differ from S. yamagishii Sakata & Hikida, 2003 — the only other known species in the genus — by the complete absence of any fingers or claws, therefore superficially resembling flippers, a combination of characters unique among terrestrial tetrapods. Sirenoscincus mobydick n. sp. is also differentiated from S. yamagishii by several apomorphic cephalic scalation characters, such as: 1) the absence of the frontonasal, likely fused with the frontal (versus presence of both scales); 2) the absence of the preocular, likely fused with the loreal (versus presence of both scales); and 3) the absence of the postsubocular, likely fused with the pretemporal (versus presence of both scales). Additionally, we provide detailed data on the appendicular skeleton of this new species of “mermaid skink” based on X-ray computed tomography that reveal several significant regressions of skeletal elements: 1) autopodial bones highly reduced in size and number; 2) highly reduced pelvic girdle and complete absence of hindlimbs, with the notable exception of two faintly distinguishable bony corpuscles probably representing rudiments of ancestral hindlimb bones; and 3) regressed sclerotic ring with five ossicles only, therefore representing the lowest value ever observed among lizards. Our study highlights the importance of the rare “forelimbs only bauplan” for investigating macroevolutionary questions dealing with complete limb loss in vertebrates, a convergent phenomenon that has repeatedly occurred 16 to 20 times within Scincidae Gray, 1825.
... Scincid lizards have radiated extensively on the island of Madagascar, with almost 80 recognized species and many more candidate species yet to be described (Glaw & Vences 2007; Crottini et al. 2009; Köhler et al. 2009, 2010; Miralles et al., 2011 a, b). According to a recent molecular study, seven endemic genera of Scincinae skinks plus one genus present also at Comoros and Glorieuse islands are currently recognized in Madagascar (Amphiglossus Duméril & Bibron, Androngo Brygoo, Madascincus Brygoo, Paracontias Mocquard, Pseudoacontias Bocage, Pygomeles Grandidier, Sirenoscincus Sakata & Hikida and Voeltzkowia Boettger) although their phylogenetic relationships and taxonomy have not been completely clarified (Crottini et al. 2009). ...
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We describe a new species of Amphiglossus skink from the western edge of the Central Highlands of Madagascar in the Reserve of Makira, and also found in the Réserve Spéciale of Ambohijanahary and in the Réserve Spéciale of Marotandra-no. Amphiglossus meva n. sp. is characterized and differentiated from other species of the genus by a combination of mor-phological, chromatic and molecular characters: 1) a relatively large size (SVL of adults from 126 to 150 mm); 2) a characteristic pattern of coloration, Amphiglossus meva being the only skink in Madagascar together with Amphiglossus crenni with dark grey dorsum contrasting with orange flanks and ventrum; 3) the absence of a postnasal scale; 4) the pre-subocular frequently absent, 5) the presence of single elongated tertiary temporal bordering lower secondary temporal and 6) pentadactyl limbs. In addition to the morphological approach, a multi-locus genetic analysis based on eight mitochon-drial and nuclear genes clearly supports the distinctiveness of A. meva. This new species was found in areas of rainforest, sometimes containing transitional deciduous forest elements. It was typically observed under large rotten logs associated with dense layers of decomposed wood retaining certain humidity and providing habitat for invertebrate larvae and ter-mites.
... The geological and climatic diversity of this area is reflected by a high species diversity and a high degree of microendemism (e. g. Andreone 2004, Wilmé et al. 2006 Montagne des Français and adjacent littoral habitats (Ramanamanjato et al. 1999, Raselimanana et al. 2000, Glaw et al. 2001, 2005, Köhler et al. 2010a, 2010b, Miralles et al. 2011. These major habitat types are separated from each other by rather steep ecotones in northern Madagascar and thus in part constitute "habitat islands" for several species, possibly allowing allopatric speciation. ...
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Paroedura hordiesi sp. n. is described from Montagne des Français, a karstic limestone massif in the far north of Madagascar recently established as nature reserve. The new species has the nostril in contact with the rostral scale and shares many characters with P. karstophila and especially with P. homalorhina which are also restricted to karstic habitats. Paroedura hordiesi differs from P. karstophila by a smoother skin on dorsum and legs, by original and regenerated tails being both entirely smooth, by colouration, and by larger snout-vent length. Morphologically the new species is most similar to P. homalorhina from the Ankarana reserve from which it can be distinguished by shorter limbs and a less slender habitus. Published molecular data place the new species as close relative of P. homalorhina and another undescribed species from Nosy Hara Island, while newly determined data of the cox1 gene for P. karstophila confirm the distinctness of the new species from this taxon. Integrating the information from published and novel molecular data, the new species differs from all nominal Paroedura (except P. vahiny for which no molecular data are available to date) by strong genetic divergences. P. hordiesi might be another microendemic species of the Montagne des Français region. We suggest its IUCN Red List classification as “Critically Endangered” on the basis that it has an extent of occurrence of at most 50 km², it is known from a single location, and there is a continuing decline in the extent and quality of its habitat. Key Words Squamata, Gekkonidae, Paroedura, new species, Madagascar, Montagne des Français conservation
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(Abstract in english) Halfway through a process of modernization undertaken some twenty years ago, the challenges facing contemporary α-taxonomy are multiple, and are as much quantitative (accelerating the inventory of living organisms) as qualitative (having reliable species hypotheses). They consist, among others, of: appropriation of concepts and techniques developed in evolutionary biology; standardization / formalization of decision-making processes (when to describe a species hypothesis?); integration of new types of data (genome, transcriptome, Ctscan); and development of adapted digital tools (interoperable and specimen-centered databases, tools dedicated to species delimitation, assessment of confidence in species hypotheses, etc.). The extent of these changes and those to come also highlights the need to revise and strengthen the epistemological basis on which this multi-secular branch of biology rests. Beyond my works on squamates systematics (mainly phylogenies and integrative revisions of tropical skinks), this dissertation will focus on my generalist contributions, i.e. those addressed to all practitioners regardless of their field of specialization. After a presentation of the different notions related to the α-taxonomy and a historical overview intended to better measure the extent of the paradigm shift this discipline is going through, a detailed synthesis of my contributions to methodological or technical questions will be presented. They are addressing, among others, issues related to taxonomic data (characterization, management, sharing and integration) and to the objective comparison of partitions resulting from different species delimitation analyses.
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A new species of diminutive, upland, forest floor skink, Sphenomorphus temengorensis sp. nov. is described from the Belum-Temengor forest complex in northern Peninsular Malaysia. This species is differentiated from all other 36 Sundaland species of Sphenomorphus on the basis of a unique suite of morphological and color pattern characteristics. This is the first reptile known to be endemic to the Belum-Temengor forest complex and underscores the need for additional field research in this area that is actively being logged.
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A new species of Mabuya lizard from the isolated Caribbean island of San Andrés is described. This species is closely related to Mabuya pergravis Barbour, 1921, another poorly known species from Providencia Island, 87 km NNE of San Andrés. Unfortunately this new species, known from a single specimen, is now probably extinct. It differs from M. pergravis in many morphological characters such as a smaller size and very different patterns of coloration, but most importantly in the presence of a very high number of nuchal scales. A new definition of this last character, which is of systematic importance in the genus Mabuya, is also given and discussed.
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Using a combination of mitochondrial and nuclear genetic markers, karyotypes and morphology, we examine the taxonomy of the Australo-papuan scincid lizard Carlia 'fusca' complex in northern Australia, all of which had been assigned previously to C. longipes. Carlia longipes, shows substantial variation in Y chromosome morphology between populations, indeed more than is seen between other species of Carlia. Analyses of the molecular genetic data and morphology demonstrate that populations with different Y chromosomes are two different species and also lead to the recognition of a third species from the Torres Strait. We herein define each of these species, for which previously described names can be applied.
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.