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A new large and colorful skink of the genus Amphiglossus from Madagascar revealed by morphology and multilocus molecular study


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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.
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Accepted by S. Carranza: 3 May 2011; published: 15 Jun. 2011
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2918: 4767 (2011)
A new large and colorful skink of the genus Amphiglossus from Madagascar
revealed by morphology and multilocus molecular study
1Technical University of Braunschweig, Zoological Institute, Mendelssohnstr. 4, 38106 Braunschweig, Germany.
2Département de Biologie Animale, Université d’Antananarivo, B.P 906, Antananarivo (101), Madagascar, and Association Vahatra,
BP 3972, Antananarivo 101, Madagascar
3WWF Madagascar and West Indian Ocean Programme Office, BP. 738, Antananarivo (101), Madagascar
4Department of Biodiversity and Evolutionary Biology, Museo Nacional de Ciencias Naturales–CSIC, C/ José Gutiérrez Abascal 2,
28006 Madrid, Spain
5Corresponding author. E-mail:
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-
Key words: Amphiglossus, conservation, Madagascar, molecular phylogeny, rainforest, Squamata: Scincomorpha: Scin-
Une nouvelle espèce de scinque de genre Amphiglossus est décrite de la bordure ouest des hauts plateaux de Madagascar,
dans la Réserve de Makira, et dans les Réserves Spéciales de Marotandrano et d’Ambohijanahary. Amphiglossus meva sp.
nov. se distingue des autres espèces du genre par la combinaison des caractères suivants: 1) une taille relativement impor-
tante (distance tête-cloaque comprise entre 126 et 150 mm chez les adultes); 2) un modèle de coloration caractéristique,
puisqu’il s’agit avec Amphiglossus crenni des deux seuls scinques malgaches dotés d’une face dorsal gris foncée con-
trastant avec des flancs et une face ventrale orange; 3) l’absence d’écaille postnasale; 4) l’écaille presuboculaire qui est
fréquemment absente, 5) la présence d’une unique temporale tertiaire, allongée et bordant la temporale secondaire in-
férieure, et 6) des membres pentadactyles. En plus de l’approche morphologique, une analyse génétique basé sur sept
gènes mitochondriaux et nucléaires soutient également la validité taxinomique de A. meva. Cette espèce n’est actuellement
connue que par quelques spécimens récoltés dans des secteurs de forêt pluviale contenant parfois des éléments de forêt
décidue transitionnel. Son biotope préférentiel semble être constitué par les larges souches et troncs d’arbres en décom-
position retenant l’humidité et hébergant des larves d’invertébrés et des termites.
48 · Zootaxa 2918 © 2011 Magnolia Press
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 rela-
tionships and taxonomy have not been completely clarified (Crottini et al. 2009). All phylogenetic studies pub-
lished to date agree (1) on the monophyly of Malagasy scincines, (2) on the existence of two main clades (Whiting
et al. 2004; Schmitz et al. 2005; Crottini et al. 2009) and (3) on the paraphyly of the genus Amphiglossus even after
the exclusion of those species now included in Madascincus. The genus Amphiglossus currently includes 21 Mala-
gasy species, with one species (A. johannae) endemic to the Comoro Islands and another one (A. valhallae) for the
Glorieuses Islands (Brygoo 1983; Glaw & Vences 2007). Three main clades within Amphiglossus have been identi-
fied, with three other genera (Androngo, Pygomeles and Voeltzkowia) nested within them (Schmitz et al. 2005;
Crottini et al. 2009), which suggests that further work is needed to elucidate their taxonomy.
The largest species (A. astrolabi and A. reticulatus) belong to a strongly supported monophyletic group within
the Amphiglossus clade (Schmitz et al. 2005; Crottini et al. 2009). Their snout-vent length (SVL) in adults can
exceed 200 mm, being also larger than any other known Malagasy skink (Fig. 1). Both species are mainly aquatic,
being found near streams and active during day and night (pers. obs.). Amphiglossus astrolabi shows a continuous
distribution in rainforests from north of the Masoala Peninsula towards the southeastern tip of Madagascar, while
A. reticulatus is known to occur in transitionary and dry forest environments, mostly in the north and central west-
ern portions of Madagascar (Brygoo 1980, Glaw & Vences 2007). A certain degree of variation has been observed
in A. reticulatus, which led to the description of A. waterloti (Angel 1930). However, recent genetic analyses sug-
gest that A. waterloti is genetically similar to A. reticulatus (Schmitz et al. 2005), therefore, until further taxonomic
evidence is provided A. waterloti should be considered as a junior synonym of A. reticulatus (Glaw & Vences
Malagasy scincine lizards show a wide diversity of morphologies, with different degrees of limb or digits
reduction, body elongation or body size (Andreone & Greer 2002; Raselimanana & Rakotomalala 2003; Crottini et
al. 2009). Although most species show a relatively dull overall coloration, with different degree of patterning on
the flanks, some species have a characteristic bright coloration including orange to reddish patterns on (1) the tail
(e.g. frequent in Madascincus igneocaudatus), (2) the ventral side of the posterior part of the body (e.g. in A. ano-
syensis), (3) the flanks (e.g. A. crenni) or (4) the whole body (e.g. in Pseudoacontias menamainty). Only A. crenni
shows a very contrasting pattern between the bright pink/reddish lateral and ventral sides of the body and tail, with
a wide dark dorsal stripe that extends from the snout to the tip of the tail.
During several zoological surveys conducted within the Réserve Spéciale (RS) of Ambohijanahary (December
1999), the RS of Marotandrano (November 2004) and the recently delimited new Protected Area of Makira (June
2009), we discovered an unknown large and colorful skink. This new taxon is distinguished from the other species
by several morphological characters, and it is further characterized by a remarkable contrasting coloration pattern,
in particular by a large dark brown dorsal band, in contrast with an orange coloration on the flanks, and a pinkish
ventral face (coloration faded after several months in preserving solution). Here, we describe this new species of
the genus Amphiglossus, compare its morphology to the most similar species in this genus, and provide new molec-
ular data to support its distinctiveness.
Material and methods
Molecular sampling. Fifty-two new mitochondrial and nuclear DNA sequences were determined from nine sam-
ples belonging either to the new species or to the Amphiglossus astrolabi / reticulatus group, and were deposited in
GenBank (Table 1). These sequences were incorporated to the dataset published by Crottini et al. (2009) (with the
exclusion of Amphiglossus crenni and Pseudoacontias menamainty for which only mitochondrial sequences were
available), both to ensure the distinctiveness of the new species, and to infer its phylogenetic affinities within the
Malagasy genera of Scincine. One lygosomine (Tiliqua) and one scincine (‘Eumeces” sensu lato) were used as out-
Zootaxa 2918 © 2011 Magnolia Press · 49
groups. Among the non-Malagasy skinks, previous more inclusive studies (Whiting et al. 2004; Schmitz et al.
2005) suggested that species of the genus Eumeces sensu lato are relatively close to the Malagasy radiation. For
these two outgroup taxa, concatenated "chimera" sequences of different species were compiled from GenBank (see
Crottini et al. 2009).
Laboratory techniques. 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
three fragments of the 12S rRNA, 16S rRNA and ND1 genes. Additionally, fragments of five nuclear DNA genes
(nuDNA) were amplified: brain-derived neurotrophic factor (BDNF); recombination activating gene 2 (Rag2); -
enolase (enol); oocyte maturation factor (C-mos) and phosducin (PDC). Standard polymerase 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). See Crottini et al. (2009) for
primers and PCR conditions used. The successfully amplified products were purified using ExoSAP-IT purifica-
tion kit according to the manufacturer’s instruction. Purified PCR templates were sequenced using dye-labeled
dideoxy terminator cycle sequencing on an ABI 3130 automated DNA sequencer.
Analysis of molecular data. All obtained DNA sequences were edited and checked for errors using Codon-
Code Aligner (v. 2.0.6, Codon Code Corporation). No stop codons were found in protein coding genes. The data
matrix included 34 samples representing 32 taxa with an aligned sequence length of 3936 base pairs (Table 1). Four
additional specimens, for which not all the genes could be successfully sequenced, were included in a separate
analysis based on a reduced number of markers. Maximum parsimony (MP) and partitioned Bayesian inference
searches based on the full concatenated dataset, were performed to infer trees. We used PAUP* 4.0b10 (Swofford
2002) to perform MP analyses with 100 random addition sequence replicates, equal character weighting, tree bisec-
tion and reconnection (TBR) branch swapping, and gaps coded as missing data. Nodal support was obtained using
bootstrap analyses, with 10000 replicates, 10 random addition sequences replicates and TBR branch swapping.
Partitioned Bayesian analyses were performed using the 21 partitions and the same parameters as previously used
by Crottini et al. (2009) with MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003). We performed one run of 20 million
generations (started on random trees) and four incrementally heated Markov chains (using default heating values)
each, sampling the Markov chains at intervals of 1000 generations. The first 10 million generations were conserva-
tively discarded and 10000 trees were retained post burn-in and summed to generate a majority rule consensus tree.
Genetic divergences were estimated with MEGA 4.1 (Tamura et al. 2007) by calculating uncorrected p-distances
from the 16S and ND1 genes.
Morphological characters and coloration. Specimens were collected using opportunistic searches and pitfall
traps (see Raxworthy & Nussbaum, 1994). The specimens captured were euthanized with a 4% chloro-butanol
solution, tissue samples were collected and conserved in pure ethanol for molecular studies, and specimens were
subsequently preserved in 70% ethanol (with exception of the UADBA specimens that have been fixed in a 12%
formalin solution before the final conservation in alcohol). Specimens examined for the present study are deposited
in the Göteborg Natural History Museum, Göteborg, Sweden (GNM); Museo Nacional de Ciencias Naturales,
Madrid, Spain (MNCN); Muséum National d’Histoire Naturelle, Paris, France (MNHN); Département de Biologie
Animale, Université d'Antananarivo, Madagascar (UADBA); Zoologisches Forschungsmuseum Alexander
Koenig, Bonn, Germany (ZFMK) and Zoologische Staatssammlung München, Germany (ZSM). All additional
specimens used for comparisons with the new species are listed in the Appendix 1.
Measurements of specimens were recorded to the nearest 0.1 mm using a dial caliper (except the tail length
which was measured with a string). Meristic, mensural and qualitative characters examined here are routinely used
in the taxonomy of Scincidae, such as scale counts, presence or absence of homologous scale fusions or the vari-
ability in color patterns. Scale nomenclature, scale counts, and measurements used in the morphological analyses
essentially follow Andreone & Greer (2002). Nuchal scales are defined as enlarged scales of the nape, occupying
transversally the place of two or more rows of dorsal cycloid scale (see Miralles 2006). The ventral scales are
counted in a single row from the postmentals to the preanal scales (both included in the count), with mental scale
excluded. The frontal scale is considered hourglass-shaped when constricted by first supraocular, bell-shaped
when this is not the case (Greer & Shea 2000).
For several voucher specimens, color pictures were taken to record alive natural coloration. Drawings were
made using Adobe Illustrator CS2 and a WACOM graphic tablet CTE-640, after photographs were taken through a
ZEISS stereomicroscope SteREO Discovery V12.
50 · Zootaxa 2918 © 2011 Magnolia Press
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52 · Zootaxa 2918 © 2011 Magnolia Press
Diagnosis of the new species is focused on distinguishing it from all other large-size species (SVL more than
130 mm, see fig. 1) of Amphiglossus that may be morphologically similar, and/or that share the same coloration
pattern, namely Amphiglossus astrolabi (Duméril & Bibron), A. ardouini (Mocquard), A. crenni (Mocquard), A.
reticulatus (Kaudern) and A. mandokava Raxworthy & Nussbaum.
Additionally, we took the opportunity of the present contribution to publicize photographs of the holotype of
Amphiglossus crenni (MNHN 1906.60) that were taken by E. R. Brygoo in 1976, and that we have recently redis-
covered in the archives of the Laboratoire des Reptiles et Amphibiens at the MNHN. These unpublished documents
constitute a precious testimony given that they represent the only available pictures for this type specimen that has
been lost between 1977 and 1982 (see Brygoo 1987). A set of six photographs of A. crenni have been deposited in
Morphobank (available at under the accession numbers M53279 to M53284, two of them
being published in the present paper.
FIGURE 1. Maximum snout vent-length (mm) recorded in Malagasy species of Amphiglossus. Data essentially from Glaw &
Vences (2007), excepted for A. crenni (based on Andreone & Greer 2002) and for A. meva and A. mandokava (present study).
Taxonomy and description of new species
The new species described herein has been recognized using an integrative taxonomic approach. The concept of
integrative taxonomy (sensu Dayrat 2005) is based on the General lineage species concept (de Queiroz 1998,
2007), rejects the superiority of any particular set of characters (morphological, behavioural, molecular) over oth-
Zootaxa 2918 © 2011 Magnolia Press · 53
ers, and advocates the combined and integrated use of various such methods (Padial et al. 2010). In the present
case, both morphological data (qualitative and quantitative scalation characteristics, coloration pattern) and molec-
ular data (phylogenetic position, genetic distances) congruently support the distinctiveness of this new species:
Amphiglossus meva sp. nov.
(figs. 2, 3)
Holotype. MNCN 44648 (field no. ZCMV 11324), collected in the western portion of the Makira plateau, close to
a campsite locally named Angozongahy, at 15°26'13.3''S 49°07'07.0''E, 1009 m above sea level, district of Mandrit-
sara, region of Sofia, province of Mahajanga, northeastern Madagascar, by D.R. Vieites, M. Vences, F. Ratsoavina
and R.-D. Randrianiaina on 28 June 2009. The holotype is in a good state of preservation; it was fixed and pre-
served in alcohol. At the time of preservation the specimen was shedding, which explains the somewhat faded col-
Paratypes (n=10). One adult specimen, MNCN 44650 (field no. DRV 5947), a subadult, MNCN 44649 (field
no. DRV 5885), and a juvenile, ZSM 0487/2009 (field no. ZCMV 11323), collected by the same collectors and at
the same locality as the holotype; a juvenile, UADBA 29402 (field no. APR 05957), collected on 24 November
2004 in the transitional rainforest at Riamalandy, 16°17.1’S 48°48.9’E, 850 m elevation within the RS of
Marotandrano, Region of Sofia, Madagascar, by A. P. Raselimanana; a juvenile, UADBA 29403 (field no. APR
05958) and an adult male, UADBA 29404 (field no. APR 05959), collected at the same date and in the same area
as above, but both were found together in a different rotten log, by A. P. Raselimanana; an adult male, UADBA
29405 (field no. APR 06021) captured on 26th November 2004 in the transitional rainforest at Riamalandy, same
conditions as above, by A. P. Raselimanana; an adult female, UADBA 29406 (field no. APR 06039) and two juve-
niles, UADBA 29407 and UADBA 29408 (field no. APR 06040 and APR 06041), collected in the same rotten logs
on 27th November 2004 in transitional forest at Riamalandy, 16°16.9’S 48°49.1’E, 800 m elevation within the RS
of Marotandrano, Region of Sofia, Madagascar, by A. P. Raselimanana.
Additional specimens (n=2). Two additional specimens were collected in the RS of Ambohijanahary. We
refrain to include these specimens in the type series given that (1) this locality is far away from both the Makira
reserve and the RS of Marotandrano, (2) these specimens have a narrower brown dorsal stripe than those from
Makira and from the RS of Marotandrano (6 scales rows vs. 10) and (3) no tissue sample from this locality was
available for molecular analysis. These specimens are: an adult male, UADBA 12209 (field no. RD 1225) and an
adult female, UADBA 12210 (field no. RD 1269) in excellent condition of preservation, collected on 18 and 19
December 1999, in the “forêt d’ Ankazotsihitafototra”, 18°15.7’S 45°25.2’E, 1150 m elevation within the RS of
Ambohijanahary, Region of Bongolava, Madagascar, by D. Rakotomalala and S. M. Goodman.
Diagnosis. A member of the phenetic Amphiglossus/Madascincus group which differs (1) from the Malagasy
genera in the subfamily Lygosominae (Cryptoblepharus and Trachylepis) by the presence of entirely movable and
scaly eyelids (versus fused immovable eyelids forming spectacles over the eyes in Cryptoblepharus; or movable
eyelids with a translucent disk or window in the lower eyelid in Trachylepis), absence of prefrontals (present in
both Cryptoblepharus and Trachylepis), and lack of frontoparietal scales (present in Trachylepis); (2) from all the
other Malagasy scincine genera by the presence of four legs.
Within the Amphiglossus/Madascincus group, it is placed in the lineage called Amphiglossus (sensu Crottini et
al. 2009) by molecular data. Within Amphiglossus, it is distinguished from all the other species by a combination of
(1) a relatively large size (SVL of adults from 126 to 150 mm); (2) a characteristic pattern of coloration with dark/
grey dorsum contrasting with bright orange to yellowish flanks and ventrum, including the ventral side of the tail;
(3) absence of a postnasal scale; (4) presubocular frequently absent, (5) presence of a single elongated tertiary tem-
poral bordering lower secondary temporal.
Among large-sized Amphiglossus (A. astrolabi, A. reticulatus, A. ardouini, A. mandokava, A. crenni), the new
species can be distinguished from the superficially similar Amphiglossus astrolabi (see fig. 4A, 5A–D) by showing
significantly shorter fingers and toes with lower numbers of lamellae under fourth finger (6–8 versus 10–13) and
fourth toe (11–13 versus 15–21); a smaller size (SVL max = 150 mm versus 226 mm); more compact head; a lower
number of ventrals (91–96 versus 99–113); absence of postnasal; presubocular frequently absent (versus always
present, most often two on each sides). From A. reticulatus (see fig. 5E–H) it can be distinguished by showing sig-
54 · Zootaxa 2918 © 2011 Magnolia Press
nificantly shorter limbs in proportion to body size; smaller size (SVL max = 150 mm versus 212 mm); a less prom-
inent parietal area and more compact head; a lower number of ventral scales (91–96 versus 95–108) and of scale
rows around midbody (32–36 versus 39–41); absence of postnasal; by the uniform dark dorsal and light ventral col-
oration (versus complex patterns). From A. ardouini it differs by the absence of postnasals (versus presence), a
higher number of scale rows around midbody (32–36 versus 31–33), shorter fingers with a lower number of lamel-
lae under fourth finger (6–8 versus 7–10) and toe (9–13 versus 17–21), a uniform dark dorsal and light ventral col-
oration (versus complex patterns, including dark transversal dark stripes in the anterior part of body). From A.
mandokava it differs by the absence of postnasals (versus present), a lower number of ventrals (91–96 versus 103–
120) and paravertebrals (95–101 versus 129–141), by the uniform dark dorsal and light ventral coloration (versus
complex patterns, including dark transversal dark stripes in the anterior part of body). From A. crenni (see fig. 4B,
6), it differs by the absence of postnasals (versus presence); a more compact body with 32–36 scales around mid-
body (versus a slender elongated body with 26–28 scales around mid-body), and pentadactyl limbs (versus
extremely reduced limbs, usually with two toes and two fingers, but sometimes with up to four). See also table 2
for a summary of morphological characteristics of the new species. Furthermore, the new species differs from all
Amphiglossus and Madascincus species for which DNA sequences were available, by high sequence divergences
in mitochondrial and nuclear genes (see below).
TABLE 2. Comparison of some characteristics distinguishing the new species from other “large-sized” (SVL 130 mm) and/or
superficially similar species of Amphiglossus. 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
1Partly based on Angel (1942) and Brygoo (1983). 2 Based on Andreone & Greer (2002) and on the photographs of the lost
holotype taken by Brygoo in 1976 (see fig. 6). 3Partly based on Raxworthy & Nussbaum (1993). 4Two types of color patterns
are presently distinguished: (1) the “bicolor pattern” with a dorsal side uniformly dark contrasting with a ventral side uniformly
light, and (2) the “variegated patterns” that may be composed by dark transversal or longitudinal stripes, dash lines, or reticula-
tions on a lighter background.
A. ardouini1A. astrolabi A. crenni2A. mandokava3A. reticulatus A. meva
SVL max (mm) 137 226 164 171 212 150
Color pattern4variegated bicolor bicolor variegated bicolor
or variegated bicolor
Postnasals Present
N sides
Presubocular N=0
n sides:
100 %
100 %
100 %
N lamellae
4th finger
n sides:
11.59 ± 0.81
(31) 5–6
5.38 ± 0.92
8.36 ± 0.84
6.85 ± 0.67
N lamellae
4th toe
n sides:
18.16 ± 1.27
10.20 ± 0.92
14.19 ± 1.22
11.23 ± 1.11
N ventral
scale rows min–max:
105.41 ± 4.40
(17) 103–120
116.20 ± 7.40
102.13 ± 4.7
93.23 ± 1.48
N paravertebral
scale rows min–max:
103.12 ± 2.82
136.75 ± 5.44
103.13 ± 4.45
98.30 ± 2.13
N longitudinal
rows at mid–body
35.53 ± 1.06
36.80 ± 1.10
40.00 ± 0.53
34.38 ± 1.12
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FIGURE 2. Drawings of the head of the holotype of Amphiglossus meva sp. n. (MNCN 44648): (A) dorsal view, (B) ventral
view, (C) lateral view, (D) close up of the ocular region. Scale bar = 2 mm.
56 · Zootaxa 2918 © 2011 Magnolia Press
FIGURE 3. Photographs of Amphiglossus meva sp. n.: (A) picture of the holotype specimen in life (MNCN 44648) and (B)
four freshly euthanasied specimens including juveniles and an adult, all from the Makira reserve ; (C) living picture of an adult
specimen from Ambohijanahary (UADBA 12209). Photographs by M. Vences and D.R. Vieites (A-B) and Harald Schütz (C),
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FIGURE 4. Photographs of two species of Amphiglossus sharing with A. meva a bicolor pattern and a relatively large size: (A)
A. astrolabi, from Ranomafana and (B) A. crenni, from Analabe. Photographs taken by F. Glaw and Franco Andreone, respec-
58 · Zootaxa 2918 © 2011 Magnolia Press
FIGURE 5. Drawings of the head of (A-D) the lectotype of Amphiglossus atrolabi (MNHN 5256) and of (E-H) a specimen of
A. reticulatus (MNHN 1931.77, holotype of Scelotes waterloti Angel, 1930). Scale bar = 4 mm.
Description of the holotype. MNCN 44648 (field number ZCMV 11324) (Fig 2, 3A). In general appearance,
a medium to large-sized Amphiglossus skink, assigned to this (paraphyletic) genus on the basis of molecular phylo-
genetic relationships and presence of pentadactyl limbs; snout–vent length (143.0 mm), 6.5 times head length (21.9
mm), almost as long as tail length (136 mm, tip regenerated). Both pairs of limbs short, pentadactyl, with very short
fingers; as a proportion of SVL, front limb 14% (20–19.5 mm) and rear limb 17% (24.5–26.2 mm).
Snout bluntly rounded in both lateral and medial aspect; rostral wider than high contacting first supralabials,
nasals and supranasals. Paired supranasals in median contact, contacting loreal. Frontonasal triangular, wider than
long, posterior side concave, contacting loreals and first suproculars. Prefrontals absent. Frontal quadrilateral, as
long as wide becoming wider at the posterior part. Supraoculars four, the second one is significantly reduced, the
first and third are barely in contact medially, the fourth supraocular is also reduced; first supraocular constricting
frontal (frontal hourglass-shaped sensu Andreone & Greer 2002), the first and the third supraoculars contacting
frontal. Frontoparietals absent. Interparietal present, well separated from supraoculars and of triangular shape, lon-
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ger than wide; parietal eye evident. Parietals contact posterior to interparietal. Two pairs of enlarged nuchals. Nasal
an anteriorly open ellipsis, just slightly larger than nostril, in contact with rostral, first supralabials and supranasals.
Postnasal absent, probably fused with first supralabials. Loreal single, longer than higher. Preocular single; presub-
ocular absent. Supraciliaries five, in continuous row, first and last pairs significantly larger and longer than the
intermediate ones; last pair projecting medially into supraocular series (thereby greatly reducing fourth supraocular
in size); upper palpebrals small except for last which projects dorsomedially slightly. Pretemporals two, both con-
tacted by parietal; postsuboculars two, the first reduced, upper contacting lower pretemporal, both contacting pen-
ultimate supralabial. Lower eyelid moveable, scaly; lower palpebrals small, longer than high, interdigitating with
large columnar scales of central eyelid; contact between upper palpebrals and supraciliaries direct but flexible, i.e.
palpebral cleft narrow. Primary temporal single. Secondary temporals two, upper long, contacting lower pretempo-
ral anteriorly and the first pair of nuchal posteriorly and overlapping lower secondary temporal ventrally; tertiary
temporal single, bordering lower secondary temporal, dorsoventrally elongated, and posteriorly followed by a scale
slightly smaller and similar in shape. Supralabials six, the fourth being the subocular which contacts scales of lower
eyelid. Postsupralabial single, external ear opening approximately half size of eye opening, circular to horizontally
suboval, with short, narrow, blunt lobules anteriorly (at least three evident, the first one being the biggest). Mental
twice wider than long; postmental diamond shaped, wider than long, contacting two infralabials. Infralabials five.
Three pairs of large chin scales, members of first pair nearly in contact medially, members of second pair separated
by one scale row, and members of third pair separated by five scale rows. Two asymmetrical postgenials posterolat-
erally 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 35; paravertebrals 98–99, including nuchals, similar in size to adjacent scales; ventrals 92, including the pre-
anals and postmentals; larger inner preanals overlap outer smaller; scales of midventral caudal series similar in size
to more adjacent scales. Both pairs of limbs pentadactyl; fingers and toes very short, clawed. Subdigital lamellae
smooth, single, with 7/8 subdigital lamellae beneath fourth digit of hands, 11/11 subdigital lamellae beneath fourth
digit of feet.
Color in life. The color in life is similar to the color in preservative as described below, except the scales of the
flanks that showed a faded orange coloration which was lost when preserved in ethanol. The orange was present in
the central portion of the scales, with the posterior border creamy-whitish.
Color in preservative. Background color of the upper side of the head, neck, back, limbs, and tail light grey/
brownish. Venter, lower side of head, throat, lower side of limbs, tail and flanks are creamish, with the flanks
slightly darker than venter. The limit between the dorsal coloration and the flanks shows a little contrast, which was
more evident in life. Dorsal scales show lighter posterior edges, and the grey/brownish coloration comprises ten
dorsal scales in wide. On the head, the area between the ear opening and the eye, including the two posteriormost
supralabial, post-supralabial and the lower temporals, are whitish (the same coloration as the throat), contrasting
with the rest of the dorsal side of the head. On all limbs, the dark coloration of the upper part does not connect with
the dorsum, having an area in the proximal part of the limb with light-cream coloration. The coloration of the palms
and feet is slightly darker than the ventral coloration. The rostral, first supralabials and the supranasals scales show
a contrasting milky or semi-translucid coloration, with a clearer whitish dot on the central part of the rostral.
Intraspecific variation. The following summary of the variation in meristic and mensural characters gives the
range for each of them, followed by the mean, ± the standard deviation, and sample size in parentheses. For some
bilateral characters, the sample size has been noted as the number of sides rather than specimens, and this is then
indicated after the sample size. Ventrals scales rows: 91–96 (93.23 ± 1.48, n=13); paravertebral scales rows: 95–
101 (98.30 ± 2.13, n=13); longitudinal scale rows at mid-body: 32–36 (34.38 ± 1.12, n=13); lamellae under 4th fin-
ger: 6–8 (6.85 ± 0.67, n sides=26); lamellae under 4th toe: 9–13 (11.23 ± 1.11, n sides=26); SVL adults: 126–
150mm (140 ± 8.0, n=7), with a minimal SVL of 68 mm recorded on a juvenile; supralabials (n sides=26): most
often six supralabial (88.46%), sometimes five (11.53%); postsupralabials (n sides=23): always single (100%);
infralabials (n sides=18): most often five (72.2%), sometimes four (16.7%) or six (11.1%); supraoculars (n
sides=26): most often four (84.6%), sometimes three (7.7%) or five (7.7%), number of supraoculars in contact with
the frontal (n sides=26): most often three (73.1%), sometimes two (19.2%) or four (7.7%); supraciliaries (n
sides=24): most often five (50%), sometimes six (29.2%) or seven (20.8%); loreals (n sides=26): always single
(100%); postnasals (n sides=26): always absent (100%).
Most of the specimens show the contrasting milky coloration on rostral, first supralabials and the supranasals
scales as in the holotype. Juveniles frequently show a whitish patch on the anterior supraciliary area. Specimens
60 · Zootaxa 2918 © 2011 Magnolia Press
from the western population (Ambohijanahary) have a narrower dark dorsal stripe (always six scale rows on the
neck, n=2) than those from Makira and Marotandrano (always ten scale rows on the neck, n=11). The life color-
ation of juveniles is similar to the adults, with a bright contrasting orange / pink on the flanks, venter, lower side of
head, throat, lower side of limbs, and tail. Nevertheless, in Makira, the pattern shown by the available series of
specimens suggests that the orange coloration fades in parallel with the age of the specimen, what does not seem to
be the case in the populations of Marotandrano and Ambohijanahary. In all specimens, the orange coloration disap-
peared after fixation, becoming cream or peach colored. See also table 2.
FIGURE 6. Photographs (A, B) of the lost holotype of Amphiglossus crenni (MNHN 1906.60; unpublished pictures taken by
E. R. Brygoo in 1976, archives of the Laboratoire des Reptiles et Amphibiens at the MNHN), and (C) a drawing made after the
photograph A. Scale borders hardly distinguishable and doubtful have been represented by dashlines. Scale bar = 2 mm.
Etymology. Meva, pronounced “mæva or mœva”, is a Malagasy word used to express beauty and refers to the
splendid bicoloration of this skink. It is used as a noun in apposition.
Phylogenetic position and genetic differentiation. The results of the phylogenetic analyses are summarized
in Figure 7. Unsurprisingly, the phylogenetic tree obtained is highly congruent with the one published by Crottini et
Zootaxa 2918 © 2011 Magnolia Press · 61
al. (2009). Although many of the most basal nodes of the genus Amphiglossus are insufficiently supported to reli-
ably infer the exact position of the new species within this taxon, some significant results may be formulated : (1)
both MP and Bayesian analyses gave results congruently supporting the placement of A. meva within the genus
Amphiglossus” sensu Crottini et al. (2009), with a bootstrap support value of 79% and a posterior probability of
1.00; and (2) the results show that Amphiglossus astrolabi is more closely related to Amphiglossus reticulatus
(88%; 1.00) than to Amphiglossus meva.
The uncorrected p-distances (Table 3) estimated between Amphiglossus meva and Amphiglossus astrolabi (p-
distances ranging between 7.4% and 8.3% for the 16S gene, and between 14.4% and 15.3% for the ND1 gene) or
between Amphiglossus meva and Amphiglossus reticulatus (5.5% to 5.8%, and 11.3% to 12.0%, respectively) are
of the same order of magnitude as those observed between Amphiglossus astrolabi and Amphiglossus reticulatus
(6.00% to 7.10%, and 12.90% to 14.20%, respectively).
TABLE 3. Summary of genetic divergence (uncorrected p-distances) within the “Amphiglossus astrolabi / meva / reticulatus
clade, estimated from 16S and ND1 sequences. Range, followed by mean ± standard deviation and sample size (inside paren-
theses) are given for both intra- and interspecific comparisons.
Habitat and distribution. In the Makira reserve, despite the use of pitfall lines, no specimen of A. meva was
collected in pitfall traps. Instead, all individuals were caught in a flat valley area not too far from a stream, at sites
that during the rainy season are probably flooded, but that during our visit (in the dry period) did not have any
water or wet soil. All specimens were found within or under large logs, which typically were largely rotten and
which maintained a certain degree of humidity without being soaked with water. The collecting site was on the
main plateau of Makira, which is made up by a vast rainforest area at an altitude between 900-1200 m above sea
level. The collecting site was characterized by a herpetofaunal composition typical for the mid-altitude eastern
rainforests of Madagascar, whereas within a few kilometers, on the western slopes of the plateau, a drastic ecotone
towards the drier areas of western Madagascar occurs, characterized by numerous herpetofaunal elements typical
for western and northern Madagascar, such as the frogs Mantidactylus ambreensis and M. ulcerosus and the gecko
Paroedura oviceps (a comprehensive account of the results of our survey at Makira will be published elsewhere).
In the RS of Ambohijanahary, the specimens UADBA 12209 and 12210 (not included in the type series) were cap-
tured in the same pitfall bucket, which was part of a line within the bottom of a forested valley, 5 m away from a
small stream, and 50 m away from the forest edge. In this area, the vegetation is the typical mid-elevation (1150 m)
primary rainforest, within the category western humid forest (Moat & Smith 2007). The RS of Ambohijanahary is
located along the extreme western portion of the Central Highlands, within the Bongolava chain, approximately 80
km NW of the town Tsiroanomandidy. This forest shows some transitional vegetation elements between two phyto-
geographic subdivisions of the Western and Central Domains (Nicoll & Langrand 1989), and can be attributed to
the humid vegetation (Moat & Smith 2007). The specimens were collected from the RS of Marotandrano at Riama-
landy, located in the northern region of Madagascar, about 12.5 km SSW of the Commune rurale de Marotandrano.
The habitat is a closed canopy midelevation (800–850m) transitional rainforest with humid vegetation (Moat &
Smith 2007) associated with taller trees and forest floor rich in organic material with thick leaf litters and diverse
detritus. All specimens were found in large and humid rotten logs, in valley closed canopy rainforest during refuge
examination. Three pitfall lines with drift fence were used during the survey, but no individuals of this species were
16S ND1
Intraspecific distances within :
Amphiglossus astrolabi
Amphiglossus meva
Amphiglossus reticulatus
0.2 – 2.1 (1.37 ± 1.02 ; 3)
0.0 (0.0 ; 3)
0.0 – 0.7 (0.47 ± 0.40 ; 3)
2.0 – 2.1 (2.00 ± 2.04 ; 6)
0.0 – 0.2 (0.13 ± 0.11 ; 3)
0.0 – 2.7 (1.80 ± 1.55 ; 3)
Interspecific distances between:
A. astrolabi / A. meva
A. astrolabi / A. reticulatus
A. meva / A. reticulatus
7.4 – 8.3 (7.70 ± 0.45 ; 9)
6.0 – 7.1 (6.52 ± 0.41 ; 9)
5.5 – 5.8 (5.60 ± 0.15 ; 9)
14.4 –15.3 (14.94 ± 0.31 ; 12)
12.9 –14.2 (13.62 ± 0.65 ; 12)
11.3 –12.0 (11.60 ± 0.27 ; 9)
62 · Zootaxa 2918 © 2011 Magnolia Press
FIGURE 7. Phylogenetic position of Amphiglossus meva sp. nov. within the Malagasy scincine. The present tree has been
inferred from a Bayesian analysis combining five nuclear (BDNF, Rag2, enol, C-mos, PDC) and three mitochondrial (12S, 16S
rRNA, ND1) genes, with posterior probabilities followed by the bootstrap support values >50% from Maximum Parsimony
analysis. Red lines indicate the positions of four specimens as resulted from a separate analysis including a reduced number of
markers (see table 1).
Zootaxa 2918 © 2011 Magnolia Press · 63
FIGURE 8. Distribution map of Amphiglossus meva. White circles represent localities sampled for the phylogenetic analyses.
64 · Zootaxa 2918 © 2011 Magnolia Press
Previous herpetological surveys using pitfall traps in the same reserve (Raxworthy, unpublished; Biodev,
unpublished), the Central Highland sites of the RS d'Ambohitantely (C.J. Raxworthy and collaborators and S.M.
Goodman and collaborators, unpublished), Ankazomivady forest (Goodman et al., 1998), Parc National d’Andrin-
gitra (Raxworthy et al., 1996), and Andranomay forest (Raselimanana 1998), the west part of Parc National du
Tsingy de Bemaraha (Bora et al. 2010), Kirindy forest (Raselimanana 2008, Raxworthy et al, unpublished), the
southwestern PK 32 forest (Raxworthy et al., umpublished,), Parc National de Zombitse (Raxworthy et al. 1994),
and Vohibasia (Goodman et al. 1997), and the northwestern RS d'Ankarafantsika (Ramanamanjato & Rabibisoa,
2002; Raselimanana, 2008) did not provide any evidence for the occurrence of A. meva n. sp. although numerous
other burrowing species (skinks and frogs) were collected in these surveys. The species appears to show a prefer-
ence for large rotten logs retaining a certain degree of humidity but in general in parts of the rainforest with rela-
tively dry soils. Based on our finding in Marotandrano and Makira, this new species is not rare but probably has a
quite strict ecological specificity with respect to its microhabitat (Fig. 8).
Ecological notes. In Ambohijanahary, two specimens were captured on two consecutive days, both in the same
pitfall trap, 5 m away from a 2 m wide stream. They may have been a breeding pair. No other individuals were
found in the RS d’Ambohijanahary despite an additional 165 trap days with pitfall devices. The specimens from
Marotandrano were all captured during refuge examination including rotten logs excavation and removal of barks
and leaf litter accumulated under taller and big dead trees. In Makira, a group of three individuals was found
together under a big log. Several larvae of coleopterans, other insects and termites were found in the same micro-
habitat, suggesting that this new skink may feed on these preys. In contrast to the other large species within the
same genus, A. meva was never found in water.
Threats and Conservation status. Habitat loss due to slash and burn agriculture, bush fires, and wood extrac-
tion are the main pressures on the Ambohijanahary and Marotandrano reserve. Forests at these reserves, especially
Ambohijanahary, are extensively fragmented. Although this forest is classified as a Réserve Spéciale, no manage-
ment plan has been proposed. The reserve and surrounding areas are known to be the domain of zebu cattle thieves
(dahalo). The forested areas within the reserve are often used by local people as a site to shelter stolen zebu. The
importance of this form of refuge has provided some protection to the remaining forest. Moreover, MNP has an
office and agents in Marotandrano Village and a coordination office is operational in Mandritsara. On the contrary,
Makira reserve currently appears to be relatively well preserved at its western edge, where the type locality of the
new species is located. Despite the common use of the forest for cattle grazing, in 2009 we could not detect major
forest destruction nor human settlements directly in the forested area. Efforts need to be undertaken to maintain this
apparently stable situation and to reduce forest destruction at the eastern lowland borders of Makira, which appears
to be relatively intense in some areas.
Madagascar is one of the richest and more biologically diverse places on Earth, and its diversity is still largely
unknown with many new species being described every year and many more yet to be discovered (Köhler et al.
2005; Vieites et al. 2009). There are several reptile species in Madagascar with bright coloration; some of them
have been considered aposematic. For example, several snakes such as Stenophis citrinus Domergue and Liophid-
ium pattoni Vieites, Fanomezana, Ratsoavina, Randrianiaina, Nagy & Vences present unique color patterns consist-
ing of alternation of yellow and black cross bands or red, yellow and blue in contrast with black respectively. None
of these species are poisonous, and the factors triggering the evolution of their aposematic coloration is unclear. In
skinks there are several examples of bright coloration that may be related to escaping or warning predators. Skinks
such as Madascincus igneocaudatus have a contrasting bright red tail, which can be autotomized if a predator bites
it, while other species show an overall bright coloration like A. crenni or Pseudoacontias menamainty that could be
considered as aposematic. However, these species are highly fossorial and as far as it is known they are not poison-
ous. It is unclear if the bright coloration evolved as a warning signal to predators, but also other burrowing taxa are
known to have aposematic coloration (Wollenberg & Measey 2009).
Both morphological and coloration characters distinguish A. meva sp. nov. from any other skink species from
Madagascar. It is remarkable that such a large and conspicuous species of vertebrate has not been detected until
now, despite this new species being relatively widespread on the island. This suggests that many more species of
Zootaxa 2918 © 2011 Magnolia Press · 65
vertebrates can still be expected from Madagascar and encourages more field and taxonomic work on its fauna.
Despite the species is only known from three sites, we have found many specimens in a short period. Until more
data is gathered, we suggest to consider this species as Data Deficient for conservation purposes following IUCN
We are grateful to Theo Rajoafiarison, Jim and Carol Patton, Emile Rajeriarison, Fanomezana Mihaja. Ratsoavina
and Roger-Daniel Randrianiaina who helped collecting specimens at Makira, to Florent and François Randriana-
solo from the WCS for their help and companionship in the field at Makira, and to our drivers Claude and Samy for
safely carrying us to the basis of the Makira slopes. Steven M. Goodman helped in the capture of several specimens
and made valuable comments on earlier versions of this manuscript. Harald Schütz took the photograph of the liv-
ing specimen from Ambohijanahary. Franco Andreone, Edouard R. Brygoo and Frank Glaw contributed photo-
graphs of Amphiglossus crenni and A. astrolabi. We also thank the Département de Biologie Animale, Université
d’Antananarivo, Madagascar National Park team in Mandritsara, the brigade of Gendarmerie in Marotandrano that
ensuring our security during the field survey, the Ministères des Eaux et Forêts, Antananarivo, and the Technical
University of Braunschweig. Many thanks to Annemarie Ohler and Ivan Ineich (MNHN), Frank Glaw (ZSM) and
Göran Nilson (GNM) for providing us access to collection specimens. We are grateful to the Malagasy authorities
for research and export permits, and to the Wildlife Conservation Society for supporting our research activities at
Makira. The field inventory in Ambohijanahary was funded by a grant from the National Geographic Society to
S.M. Goodman and D. Rakotondravony, and the mission to Marotandrano was generously supported by the Mac-
Arthur Foundation in the context of the RAP–Gasy project. We are indebted to the Ecology Training Program and
the World Wide Fund for Nature, Antananarivo, for logistical support. Funding was provided by a postdoctoral
research fellowship of the Alexander von Humboldt Foundation and by a SYNTHESYS grant (FR–TAF–842) to
AM, by the Volkswagen Foundation to MV, and by a Spanish Ministry of Science and Innovation grant
(CGL2009–10198) to DRV.
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APPENDIX 1. List of additional specimens examined.
Amphiglossus ardouini (4)
Antsiranana Province : MNHN 1896.417, MNHN 1897.32 (respectively paralectotype and lectotype of Sepsina
ardouini Mocquard, 1897), “Diego Suarez” (=Antsiranana) ; ZSM 1554-2008, Montagne des Francais, ca. 2.3 km
WNW Andavakoera ("trapsite 3"), 250m (12°19,575'S 49°20,495'E), coll. by E. Randriamalala on 02.04.2006 ;
ZSM 1555-5008, Ambombofofo-Region (Frontier base camp), 28m (12°05,528'S 49°19,485'E) coll. by S. Megson
on 11.11.2006.
Amphiglossus astrolabi (16).
Fianarantsoa province: MNHN 1930.341, Ikongo massif, coll. by R. Decary; MNHN 1965.284, .285, .286, .287,
probably all collected at Pic Ivohibé near Andringitra Massif, according to the catalogue of the Paris museum with
the notion "Vivarium at the IRSM (?)" thus probably kept in captivity for some time before preservation. ZSM 201/
2003, Ranomafana, near Hotel Manja, 19.1.2003, coll. by F. Glaw, M. Puente, L. Raharivololoniaina, M. Thomas
and D.R. Vieites. Toamasina province: MNHN 1906.59, Fanovana. MNHN 1983.518, region of the Alaotra lake,
coll. by Therefieu, 24 mai 1961. ZSM 1557/2008, near Andasibe, 02/03. 2008, coll. by M. Vences. Unknown
exact localities: MNHN 5256, coll. by Quoy & Gimard; MNHN 1937.84, coll. by Lavanden; MNHN1991.4274.
MNHN 8880, 8880A, 8880B, Mandraka? Institut de Recherche Scientifique de Madagascar (IRSM) (?) – this
specimen might originate in fact from Mandraka (located between Antananarivo and Moramanga) but could also
originate from other parts of Madagascar and been collected by A, Peyrieras and subsequently maintained in cap-
tivity at his farm in Mandraka. MNHN 8389, with the untraceable locality information "Mosbaie", coll. by L. Rou-
Amphiglossus crenni (2).
Toamasina province: MNHN 1906.60, Fanovana (holotype specimen of Scelotes crenni Mocquard, 1906); this
specimen is considered to be lost since 1982, according to E.R. Brygoo, in the collection catalogue at the MNHN;
MNHN 1980.1190, region of the Alaotra lake.
Amphiglossus mandokava (4).
Antsiranana province: ZSM 208-2003, Montagne d’Ambre, 800-900m, coll. By F. Glaw, R.D. Randrianiaina &
A. Razafimanantsoa on 19.02.2003 ; ZSM 312-2004, Montagne d’Ambre, 900m (12°30'S 49°10'E), coll by F.
Glaw, M. Puente, R. Randrianiaina & A. Razafimanantsoa on 27.02.2004 ; ZSM 2167-2007, Montagne d'Ambre,
Cascade Antakarana, coll. P. Bora on 13.03.2007 ; ZSM 2232-2007, Montagne d’Ambre, coll. by P. Bora & I.
Knoll on 03./04.2007.
Amphiglossus reticulatus (8).
Antsiranana province: MNHN 1956.57, Ambilobé, coll. by J. Guibé; MNHN 1931.77 Ambilobé, coll. by Water-
lot (holotype of Scelotes waterloti Angel, 1930). Mahajanga Province: MNHN 1978.2703, 1978.2704, Forêt de
Boza, Antsohihy, coll. by Anthony Randriamihanta, in march 1977 ; ZSM 527-2001, 528-2001 529-2001, Ankara-
fantsika (Ampijoroa) coll. by M. Vences, D.R. Vieites, G. Garcia, Raherisoa, Rasoamamonjinirina from 25 to
28.2.2001. GNM 1520, St Marie de Marovoay (holotype of Sepsina reticulata Kaudern, 1922, only examined from
high resolution photographs).
... 71. (Miralles et al. 2011) 72. (Mori 2002) 73. ...
... can get up to 70 mm snout to vent (Bauer 2004), therefore capable of taking small toads. The scincid was Amphiglossus astrolabi, one of the largest scincids on Madagascar (Miralles et al. 2011). ...
... Amphiglossus astrolabi is likely to encounter D. melanostictus if D. melanostictus ventures out from disturbed areas. Amphiglossus astrolabi avoid disturbed areas, are found near water bodies and will actively forage during night and day (Miralles et al. 2011;Raxworthy and Nussbaum 1993). Unlike the other squamates the only aspect of their Na + ,K + -ATPase successfully sequenced was the alpha 1 isoform. ...
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Invasive and introduced species can pose major ecological challenges to vulnerable native wildlife. Biodiversity hotspots, in particular, require protection from this significant cause of species loss. One hotspot, Madagascar, is experiencing the accidental introduction of a potentially ecologically damaging species – the toxin carrying bufonid toad, Duttaphyrnus melanostictus. The presence of these toxic invaders drives fears that if such a species gains a foothold widespread poisoning of Malagasy predators could occur, mirroring the invasion of Australia by Rhinella marina. This includes numerous endemic and endangered species. The mechanism by which the toxin acts upon organisms has been previously identified via the study of toxin resistant versus toxin non-resistant taxa. Specific amino acid substitutions are required on the organism’s Na+/K+–ATPase for them to be resistant to bufonid toxin. This solution to combat the toxin is widely consistent across taxa providing a method to discover and predict toxin resistance or vulnerability. Here I investigate the Na+/K+–ATPase gene to detect vulnerability of a selection of Malagasy fauna to the toxics of Duttaphrynus melanostictus. It is discovered that no tested species on Madagascar have the capacity to survive ingestion of the novel toxin. The vulnerability is found in all examined species, including snakes, frogs, lizards, lemurs and tenrecs. The results suggest that the invasive Duttaphrynus melanostictus is liable to have significant impact on Malagasy fauna.
... 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. ...
... The BI tree based on the complete concatenated dataset is highly congruent with previously published phylogenies [1], [7], [26]. It retrieves the two main clades A and B (cf. [36]) within Malagasy scincines (Fig 2). ...
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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.
... 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.
... We focused on some of the most diagnostic and commonly used characteristics in skink taxonomy, allowing us to obtain a more inclusive overview of generic diversity and evolutionary trends: (1) number of presacral vertebrae (indirect proxy of body elongation [length and shape]); (2) number of scale rows around midbody (related to body width); (3) number of ventral scales between the mental and the cloaca (indirect proxy of body elongation [length and shape]) and (4) number of lamellae under the fourth toe (indirect proxy of toe length, and therefore limb reduction). Data were collected from the available literature (Angel, 1942;Andreone and Greer, 2002;Brygoo, 1980Brygoo, , 1981aBrygoo, ,b, 1983Brygoo, , 1984aBrygoo, -d, 1985Brygoo, , 1987Glaw and Vences, 2007;Miralles et al., 2011c;Miralles and Vences, 2013;Raxworthy and Nussbaum, 1993) and complemented by examination of museum specimens (Table SM5). To ascertain the number of presacral vertebrae, 36 specimens of 16 species of Amphiglossus were radiographed using a Faxitron PathVision (Faxitron Bioptics LLC). ...
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.
... Madascincus igneocaudatus - Glaw and Vences (2007, partim), Crottini et al. (2009;partim), Miralles et al. (2011a, c, partim;2011b); Madascincus igneocaudatus "igneocaudatus-C clade" - Miralles and Vences (2013); Madascincus sp. "igneocaudatus" central clade - Miralles et al. (2015). ...
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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.
... The definition of the genus Madascincus herein follows previous molecular work [49][50][51][52], encompassing all species of an exclusively four-legged lineage that is sister to the legless genus Paracontias. Throughout the manuscript we use species names (scientific binomina) largely following current taxonomy (see File S1). ...
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Delimiting and describing species is fundamental to numerous biological disciplines such as evolution, macroecology, and conservation. Delimiting species as independent evolutionary lineages may and often does yield different outcomes depending on the species criteria applied, but methods should be chosen that minimize the inference of objectively erroneous species limits. Several protocols exploit single-gene or multi-gene coalescence statistics, assignment tests or other rationales related to nuclear DNA (nDNA) allele sharing to automatically delimit species. We apply seven different species delimitation protocols to a taxonomically confusing group of Malagasy lizards (Madascincus), and compare the resulting taxonomies with two newly developed metrics: the Taxonomic index of congruence C tax which quantifies the congruence between two taxonomies, and the Relative taxonomic resolving power index R tax which quantifies the potential of an approach to capture a high number of species boundaries. The protocols differed in the total number of species proposed, between 9 and 34, and were also highly incongruent in placing species boundaries. The Generalized Mixed Yule-Coalescent approach captured the highest number of potential species boundaries but many of these were clearly contradicted by extensive nDNA admixture between sympatric mitochondrial DNA (mtDNA) haplotype lineages. Delimiting species as phenotypically diagnosable mtDNA clades failed to detect two cryptic species that are unambiguous due to a lack of nDNA gene flow despite sympatry. We also consider the high number of species boundaries and their placement by multi-gene Bayesian species delimitation as poorly reliable whereas the Bayesian assignment test approach provided a species delimitation highly congruent with integrative taxonomic practice. The present study illustrates the trade-off in taxonomy between reliability (favored by conservative approaches) and resolving power (favored by inflationist approaches). Quantifying excessive splitting is more difficult than quantifying excessive lumping, suggesting a priority for conservative taxonomies in which errors are more liable to be detected and corrected by subsequent studies.
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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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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.
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Madagascar is home to more than 10 000 plant species, 80% of which occur nowhere else in the world. With natural vegetation ranging from rainforest to unique spiny forest, Madagascar’s range of plant diversity makes it one of the world's most important biodiversity hotspots. In common with many other tropical countries, the flora of Madagascar is extremely threatened not only by habitat destruction for agriculture, fuelwood, building materials and so on, but also, in the case of certain species, by over-collection for the horticultural trade. The CEPF Madagascar Vegetation Mapping Project is a three-year project (2003–2006), funded by the Critical Ecosystem Partnership Fund (CEPF) and managed jointly by The Royal Botanic Gardens, Kew, Missouri Botanical Garden, and Conservation International’s Center for Applied Biodiversity Science. The project is innovative in a number of ways. It employs state-of-the art remote sensing technology and methodologies to delimit Madagascar’s vegetation. It represents an all-inclusive collaboration between specialists from a wide range of botanical and conservation institutions, which has ensured the most thoroughly ground-truthed vegetation map ever compiled for Madagascar. Finally, through a series of workshops, it incorporates detailed consultations with the conservation community to ensure that the final products are of maximum relevance and utility to conservation planners and managers. An accurate and updated vegetation map is imperative for conservation planning and natural resource management in Madagascar. It is also essential that the data on which such a map is based be made freely available, so that conservation organisations, government departments, academic institutions and other stakeholders can use them as an up-to-date standard dataset on which to base their activities. The electronic version of this atlas is available on Kew’s website (, and local experts were invited to continually improve and update the map. In order for a vegetation map to fulfil its intended role it must accurately delimit areas with various vegetation types as they currently exist, and assign those areas to objective categories that can be easily recognised in the field and that reliably reflect fundamental biological differences (primarily structural features, for example, physiognomy). Madagascar is becoming increasingly aware of the need to protect its biodiversity. The most immediate use of this vegetation map in conservation is likely to be by protected area managers who wish to understand the flora of their designated areas. It will also provide a valuable baseline for monitoring longer-term changes in vegetation inside and outside protected areas. However, Madagascar also provides an exceptionally high rate of species discovery and description, and this atlas will be used by field biologists attempting to identify potential sampling sites for biodiversity surveys, which will in turn yield data that is critical for biogeographic research and conservation planning. At the 2003 World Parks Congress, Madagascar’s President Marc Ravalomanana emphasised his country’s commitment to conservation by announcing its intent to triple the size of its existing protected area network. This admirable effort to prevent the extinction of many of Madagascar’s endemic species has become known as the ‘Durban Vision’. In order to ensure effective preservation of Madagascar’s biodiversity, the identification of sites for these new protected areas should follow a systematic process. A recent workshop on systematic conservation planning (November 2005, Antananarivo) highlighted the importance of using habitat types and indicators of habitat quality in addition to species distribution data when conducting conservation prioritisation analyses, concluding that this is the best way to produce robust conservation solutions. Because only a small proportion of Madagascar’s species have had their distributions documented, the vegetation types identified by this mapping project are good surrogates for habitat diversity and for the majority of the biota, which is so little known. In addition, conservation practitioners, including NGOs and donors, need information on trends in natural vegetation cover and quality in order to assess the outcomes of their conservation work. The Convention on Biological Diversity includes trends in the extent of habitats among its headline indicators for tracking progress towards the 2010 target (SBSTTA, 2004). The immediate focus of the Durban Vision group will be on establishing new protected areas (map 1) in remaining native vegetation, although subsequent attention could productively turn to managing that vegetation, and the habitat quality categories in the atlas provide valuable information. The atlas also provides important up-to-date information on native vegetation cover and quality, which maximises its potential to aid planning for future habitat restoration activities.
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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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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.
— 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.