Corresponding author: Aurélien Miralles (
Academic editor: Johannes Penner
In a previous study,
The genus
Muséum National d’Histoire Naturelle, Paris National History Museum, London Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main Université d’Antananarivo, Département de Biologie Animale
nuclear DNA mitochondrial DNA
Comparison of phylogenetic trees of the genus
Haplotype network reconstructions for the four nuclear genes (BDNF, PDC, CMOS and RAG2). For each marker, circles represent haplotypes (size proportional to the number of individuals), black lines represent mutational steps and black dots missing haplotypes, white curves represent connections between haplotypes found co-occurring in heterozygous individuals, and white numbers represent the number of individuals in which the respective haplotypes were found co-occurring. Single locus fields of recombination (pools of co-occurring haplotypes) are represented by grey rectangles (redrawn from
Despite numerous conflicts among the seven methods of species delimitation (
Considering an integrative taxonomic approach, the distinctiveness of the two clades
Both clades represent monophyletic units fully supported by both the
The
Both clades are unambiguously morphologically diagnosable from their respective sister clade and from all the other species of
The genetic distance values between these two clades and their respective sister clades are relatively high, with p-distances ranging from 7.3 to 9.0% (16S) and 16.2 to 17.7% (ND1) between
Drawings of the lateral and dorsal views of the heads of most of the species of
Photographic plate showing most of the recognized species of
Comparison of the most relevant morphological characters, plus additional data on the altitudinal distribution and reproductive mode of
|
||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
|
|
|
||||||||||
|
8–11 | 8–11 | 5–7 | 5–8 | 5–8 | 8–11 | 3–5 | 6–7 | 7–8 | 6–9 | 6–9 | |
9.0 ± 0.9 | 9.1 ± 0.9 | 6.0 ± 0.8 | 7.0 ± 0.6 | 6.3 ± 1.0 | 9.8 ± 0.7 | 3.9 ± 0.6 | 6.4 ± 0.5 | 7.6 ± 0.5 | 7.5 ± 0.7 | 7.3 ± 0.8 | ||
(50) | (14) | (13) | (40) | (9) | (16) | (8) | (11) | (22) | (26) | (27) | ||
|
15–22 | 15–18 | 9–13 | 12–16 | 12–15 | 16–20 | 5–8 | 16–19 | 18–23 | 16–22 | 15–20 | |
18.1 ± 1.4 | 16.4 ± 1.3 | 11.3 ± 1.5 | 14.1 ± 1.2 | 13.8 ± 1.1 | 17.5 ± 1.2 | 6.8 ± 1.3 | 17.5 ± 0.8 | 20.6 ± 1.3 | 18.5 ± 1.5 | 17.9 ± 1.2 | ||
(54) | (13) | (10) | (52) | (12) | (15) | (9) | (13) | (20) | (22) | (28) | ||
|
68–83 | 73–78 | 55–63 | 56–61 | 59–63 | 63–66 | 52–60 | 75–80 | 65–73 | 74–78 | 70–88 | |
76.7 ± 4.4 | 75.7 ± 1.8 | 58.3 ± 3.0 | 58.8 ± 1.2 | 60.2 ± 1.5 | 64.3 ± 1.0 | 57.6 ± 3.3 | 77.9 ± 1.6 | 68.7 ± 2.1 | 75.8 ± 1.2 | 81. 3 ± 4.0 | ||
(21) | (7) | (7) | (27) | (6) | (8) | (5) | (7) | (14) | (12) | (16) | ||
|
69–80 | 71–79 | 57–65 | 51–62 | 52–62 | 60–65 | 50–57 | 74–81 | 65–79 | 71–81 | 76–88 | |
74.7 ± 3.0 | 74.6 ± 3.7 | 59.7 ± 3.4 | 55.9 ± 2.9 | 57.7 ± 3.1 | 62.6 ± 2.1 | 53.6 ± 2.5 | 79.0 ± 2.3 | 68.7 ± 3.3 | 77.9 ± 2.6 | 82.7 ± 3.2 | ||
(26) | (7) | (7) | (28) | (7) | (8) | (5) | (7) | (14) | (13) | (15) | ||
|
24–26 | 22–24 | 22–26 | 24–26 | 22–26 | 28–30 | 18–20 | 26 | 24–26 | 24–26 | 30–32 | |
24.2 ± 0.6 | 23.3 ± 1.0 | 24.0 ± 1.2 | 24.1 ± 0.4 | 23.7 ± 1.5 | 29.0 ±4 2.1 | 19.6 ± 0.9 | 26.0 ± 0 | 24.1 ± 0.5 | 25.4 ± 0.9 | 31.6 ± 0.8 | ||
(28) | (7) | (7) | (27) | (6) | (8) | (5) | (7) | (14) | (13) | (16) | ||
|
– | – | – | – | – | – | – | 42.9% | 92.3% | 56.3% | 81.3% | |
– | – | – | 2% | 7.1% | 100% | 40.0% | 57.1% | 7.7% | 37.5% | 18.7% | ||
23.2% | 21.4% | 28.6% | 50% | 35.8% | – | 20.0% | – | – | 6.2% | – | ||
71.4% | 78.6% | 57.1% | 48% | 57.1% | – | 40.0% | – | – | – | – | ||
5.4% | – | 14.3% | – | – | – | – | – | – | – | – | ||
(56) | (14) | (14) | (58) | (14) | (16) | (10) | (14) | (26) | (22) | (32) | ||
|
100% | 100% | 100% | 100% | 100% | 100% | 100% | – | 89.3% | 100% | 94.4% | |
– | – | – | – | – | – | – | 100% | 10.7% | – | 5.6% | ||
(56) | (14) | (14) | (58) | (14) | (16) | (10) | (14) | (28) | (26) | (36) | ||
|
– | – | – | – | – | 87.5% | – | – | – | – | – | |
100% | 100% | 100% | 100% | 100% | 12.5% | 100% | 100% | 100% | 100% | 100% | ||
(28) | (14) | (7) | (28) | (7) | (8) | (10) | (7) | (14) | (13) | (16) | ||
|
100% | 100% | – | – | – | – | – | 100% | 100% | 100% | 47.2% | |
– | – | 100% | 100% | 100% | 100% | 100% | – | – | – | 52.8% | ||
(23) | (12) | (14) | (29) | (14) | (8) | (10) | (7) | (14) | (13) | (18) | ||
|
73.0 | 54.2 | 47.4 | 53.5 | 50.5 | 68.5 | 33.6 | 81.7 | 61 | 75 | 114.0 | |
56.3 ± 11.6 | 52.3 ± 2.1 | 42.0 ± 5.1 | 49.5±2.5 | 48.0 ± 2.4 | 60.1 ± 9.6 | 27.8 ± 8.2 | 72.3 ± 6.1 | 54.9 ± 3.1 | 66.0 ± 7.1 | 89.6 ± 10.8 | ||
(9) | (4) | (7) | (21) | (5) | (7) | (6) | (7) | (14) | (13) | (14) | ||
|
– | 7.1% | 7.1% | – | – | – | – | – | – | – | n/a | |
98.2% | 92.9% | 57.1% | 8.6% | 35.7% | 93.8% | 66.6% | 92.9% | 100% | 87.5% | |||
1.8% | – | 35.8% | 79.3% | 50.0% | 6.2% | 33.3% | 7.1% | – | 12.5% | |||
– | – | – | 12.7% | 14.3% | – | – | – | – | – | |||
(56) | (14) | (14) | (58) | (14) | (16) | (6) | (14) | (2) | (8) | |||
|
1.8% | – | – | – | 7.1% | – | 100% | – | 3.6% | – | – | |
98.2% | 100% | 100% | 100% | 92.9% | 100% | – | 100% | 96.4% | 100% | 100% | ||
(56) | (14) | (14) | (58) | (14) | (24) | (10) | (14) | (28) | (24) | (32) | ||
|
spectacled | spectacled | spectacled | spectacled | spectacled | scaly | scaly | scaly | scaly | scaly | scaly | |
|
viviparous | oviparous | ? | oviparous? | ? | oviparous | ? | ? | ? | ? | ? | |
|
≤ 500 m | ≥ 1500 m | ≤ 1000 m | ≤ 1000 m | ≤ 1000 m | ≤ 1000 | 500–1500 m | ≤ 500 m | ≤ 500 m | ≤ 500 m | ≤ 500 m |
Summary of the most relevant morphological characters differentiating each pair of species of : number of lamellae under 4th finger : number of lamellae under 4th toe : number of ventral scale rows : number of paravertebral scale rows : number of longitudinal scale rows at midbody : number of enlarged nuchal scales : presence or absence of postnasal scales : shape of the frontal scale : position of the subocular scale : aspect of the lower eyelid window : reproduction mode
|
|
|
|
|
|
|
|
|
||
---|---|---|---|---|---|---|---|---|---|---|
|
F, T, |
F, N, |
F, |
F, |
T, |
T, |
|
|
|
|
|
– | F, T, |
F, T, |
F, T, |
T, |
T, |
T, |
F, T, |
F, T, |
F, T, |
|
– | – | R | F, T, |
N, |
|
||||
|
– | – | – | F, T, |
N, |
|
||||
|
– | – | – | – | F, T, |
T, |
||||
|
– | – | – | – | – | none | none | T, |
T, |
T, |
|
– | – | – | – | – | – | none | T, |
||
|
– | – | – | – | – | – | – | T, |
T, |
|
|
– | – | – | – | – | – | – | – |
|
|
|
– | – | – | – | – | – | – | – | – |
|
Genetic divergences among
Inter-specific distances (%) | Intra-specific distances (%) | ||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
|
|
|
|
|
|
|
|
|
|
16S | ND1 | |
|
- | 19.8 | 17.9 | 19.2 | 17.5 | 16.6 | 17.3 | 16.9 | 12.5 |
|
10.5 | 0.1 | 1.1 |
|
10.1 | - | 21.1 | 21.2 | 21.3 | 19.0 | 21.3 | 20.0 | 21.1 | 21.2 | 20.5 | 0.0 | 0.0 |
|
8.2 | 10.1 | - |
|
18.9 | 17.3 | 18.3 | 20.8 | 19.5 | 19.2 | 19.3 | 1.5 | 3.5 |
|
9.3 | 9.9 |
|
- | 20.5 | 19.3 | 20.1 | 18.8 | 19.3 | 20.0 | 19.3 | 2.5 | 5.9 |
|
8.3 | 10.9 | 9.8 | 11.2 | - | 8.2 | 14.7 | 17.5 | 16.5 | 17.9 | 16.2 | 0.5 | 1.0 |
|
8.5 | 10.6 | 9.7 | 11.6 | 3.4 | - | 12.3 | 17.1 | 15.9 | 16.1 | 15.2 | 0.5 | 0.7 |
|
8.7 | 11.3 | 10.9 | 12.5 | 8.5 | 8.0 | - | 17.0 | 17.7 | 18.1 | 16.6 | 3.1 | 5.7 |
|
7.0 | 8.6 | 9.2 | 8.7 | 9.9 | 9.9 | 10.6 | - | 15.8 | 17.6 | 15.6 | 0.7 | 1.3 |
|
3.4 | 9.8 | 8.7 | 10.0 | 8.4 | 9.2 | 9.2 | 6.8 | - | 10.5 | 9.5 | 1.2 | 3.1 |
|
|
10.1 | 9.2 | 10.0 | 9.2 | 9.3 | 10.3 | 7.1 | 3.4 | - | 8.9 | 0.5 | 1.5 |
|
3.4 | 8.7 | 9.1 | 9.4 | 7.6 | 8.6 | 9.3 | 6.7 | 3.0 | 3.8 | - | 1.1 | 2.7 |
(n=23, all from Antsiranana province, northern Madagascar).
(n=2, not sequenced).
A member of the genus
Coloration of the holotype in preservative with a pair of lateral dark brown stripes (about two scales wide on the neck) relatively large and well defined anteriorly (overlapping rostral, mental, first four supralabials, loreals, and presuboculars), then progressively breaking up into two parallel very thin dashed lines posteriorly to forelimbs, hardly distinguishable from the rest of the dots covering the body. Dorsum and dorsal sides of forelimbs, hindlimbs and tail light bronze. The bronze dorsal field and flanks are covered by numerous little dark dots, each of them in the middle of a dorsal scale, in contact with its posterior edge; resulting in many thin dash lines (14 to 16 at midbody, including the dark lateral stripes), darker and more contrasted in the posterior part of the dorsum, then posteriorly becoming progressively indistinguishable from the background coloration of tail, and laterally, becoming progressively indistinguishable from the light coloration of the ventral field. No distinct border between the background coloration of the dorsal and the ventral sides. Immaculate whitish ventral field extending from lower side of head (mental excluded), throat, lower side of limbs and venter, to the ventral side of tail. Palms and soles barely darker than venter. Coloration in life was almost identical to the coloration in preservative, with the only significant difference being the presence of iridescent glints of scales and a venter with some violet-pinkish tint (cf. Figs
For variation in measurements and scale characters see Table
The specific epithet
The species is known from northernmost Madagascar including at least four localities (see localities of type specimens above and Fig.
Distribution maps for
A member of the genus
Coloration in preservative with upper side of the head, neck, back, limbs, and tail dark bronze. Venter, lower side of head, throat, lower side of limbs, and tail withish/cream. Lateral borders on the ventral side maculated by very small dark dots. Six very well defined very dark stripes run along the body, continuing along the first third of the tail, then abruptly ending where the tail is regenerated. Two thin blackish dorsal stripes formed by succession of contiguous dots start on the supranasal; at midbody, each dorsolateral stipe is less than one scale wide and both are separated by two rows of dorsal scales. Two wide dark brown upper lateral stripes; margins slightly darker and very sharp; about two scales wide at midbody and overlying three rows of scales; starting from the rostral, where the stripes all meet, extending on the upper half of each supralabial, the loreals, around the eyes, above ear opening, and above forelimbs and hindlimbs. Two thin dark lower lateral stripes, starting on the last infralabials, extending through the forelimb and hindlimb insertion. At midbody, each lower lateral stripe is less than one scale wide, with irregular margin. Four very light stripes run along the body, continuing along the first third of tail, then abruptly ending where the tail is regenerated. Two whitish dorsolateral stripes separate dark dorsal stripes from the upper lateral dark stripes; about one scale wide at midbody and overlying two rows of scales. Two whitish lateral stripes separating the dark upper lateral stripes from the dark lower lateral stripes; about one scale wide at midbody and overlying two rows of scales. Regenerated part of the tail cream, maculate with many small dark dots on the dorsal side. Palms and soles darker than the ventral side.
Life coloration for the holotype has not been documented, but it is apparently very similar to the coloration in preservative, with exception of the tail which is usually bright red, or pinkish brown in some specimens (cf. Fig.
For variation in measurements and scale characters see Table
The specific epithet
The species is known from the dry environments on two massifs in the central highlands of Madagascar, in Mont Ibity and in Itremo (Fig.
A list of the species currently recognized, including information supporting their respective taxonomic validity is presented in Table
a | postnasal always absent |
|
a' | postnasal mostly present (rarely absent in |
b |
b | 28 or more scale rows at midbody | c |
b' | 26 or less scale rows at midbody | d |
c | 30–32 scale rows at midbody, 76–88 paravertebral scale rows, 70–88 ventral scale rows, frontal and interparietal separated together |
|
c' | 28–30 scale rows at midbody, 60–65 paravertebral scale rows, 63–66 ventral scale rows, frontal and interparietal mostly fused together |
|
d | 18–20 scale rows at midbody, 8 or less lamellae under 4th toe, frequently less than 5 fingers, adult SVL < 35 mm, an atypical head shape with a short and acuminate snout and relatively large eyes | |
d' | 22–26 scale rows at midbody, 9 or more lamellae under 4th toe, always 5 fingers, SVL in adult > 40 mm | e |
e | Frontal bell-shaped, 65 or more ventral scale rows, 65 or more paravertebral scale rows, tail frequently red colored | f |
e' | Frontal hour-glass shaped, 63 or less ventral scale rows, 65 or less paravertebral scale rows, tail always brown colored | |
f | Two lateral dark brown stripes relatively large and well defined anteriorly, then progressively breaking up into two parallel very thin dashed lines posteriorly to forelimbs, hardly distinguishable from the rest of the dots covering the body, lower eyelid scaly | g |
f' | Four or six well defined and brightly contrasted dark stripes running along the body, lower eyelid spectacled | h |
g | 65–73 ventral scale rows |
|
g' | 74–78 ventral scale rows |
|
h | A relatively short and rounded snout, six well defined and contrasted stripes running along the body, 22–24 scale rows around midbody |
|
h' | A relatively long and pointed snout, four well defined and contrasted stripes running along the body, 24–26 scale rows around midbody |
|
List of the species presently recognized in the genus = Methods of species delimitation
Taxa | Morphological diagnosability | Monophyly | Species delimitation methods |
---|---|---|---|
|
Unambiguously diagnosable | Monophyletic ( |
Supported by six of seven |
|
Unambiguously diagnosable | Monophyletic ( |
Supported by all seven |
|
Unambiguously diagnosable | Monophyletic ( |
Supported by six of the seven |
|
Unambiguously distinguishable from most of the species (see below |
Monophyletic ( |
Supported by all seven |
|
Unambiguously distinguishable from most of the species (see species delimitation) | Not tested | Rare species so far not studied using molecular methods and therefore not included in |
|
Unambiguously distinguishable from almost all other species of |
Monophyletic ( |
Supported by six of the seven |
type locality |
|||
|
Unambiguously distinguishable from almost all other species of |
Monophyletic ( |
Supported by all seven |
|
Unambiguously diagnosable | Monophyletic ( |
Supported by all seven |
|
Unambiguously diagnosable | Monophyletic ( |
Supported by all seven |
|
Species of the |
Monophyletic ( |
Supported by five of the seven |
|
Species of the |
Monophyletic ( |
Supported by five of the seven |
|
Species of the |
Monophyletic ( |
Supported by five of the seven |
The different phylogenetic inferences applied (separated phylogenetic Bayesian analysis based on the
Two of these clades reveal a relevant biogeographical component: (1) The
In contrast, the
Research in Madagascar was made possible in the framework of collaboration agreements among the Université d’Antananarivo (Faculté des Sciences – Mention Biologie Animale), the Direction des Eaux et Forêts, Madagascar, the
List of specimens examined morphologically
Adobe PDF file
Specimens examined are listed including collection number, locality and collector information.