Paroedura manongavato, Piccoli & Belluardo & Lobón-Rovira & Oliveira Alves & Rasoazanany & Andreone & Rosa & Crottini, 2023

Piccoli, Costanza, Belluardo, Francesco, Lobon-Rovira, Javier, Oliveira Alves, Ivo, Rasoazanany, Malalatiana, Andreone, Franco, Rosa, Goncalo M. & Crottini, Angelica, 2023, Another step through the crux: a new microendemic rock-dwelling Paroedura (Squamata, Gekkonidae) from south-central Madagascar, ZooKeys 1181, pp. 125-154 : 125

Paroedura manongavato sp. nov.

Figs 2 , 4 , 6 , 7A, C , 8

Remarks.

This species was previously referred to as Paroedura bastardi D by Miralles et al. (2021) and Paroedura sp. aff. bastardi Lineage D UCS by Belluardo et al. (2021). A single individual (ZCMV 12790) collected at Anja Reserve was molecularly examined in Miralles et al. (2021). Further samples have been analysed in Belluardo et al. (2021), which contributed to extend the species distribution to Tsaranoro Valley (ca. 25 km south of Anja Reserve).

Type locality.

Anja Reserve, 21.85098°S, 46.84270°E, elevation ca. 950 m a.s.l., Ambalavao, Fianarantsoa District, Haute Matsiatra Region, Madagascar.

Type material.

Holotype. ZSM 9/2023 (ACZCV 0300; ACZC6935; ACP2761), adult female from Anja Reserve, 21.85098°S, 46.84270°E, elevation ca. 950 m a.s.l., Ambalavao, Fianarantsoa District, Haute Matsiatra Region, Madagascar, collected on the 27 November 2014 by Franco Andreone, Angelica Crottini and Gonçalo M. Rosa.

Paratypes. UADBA R-uncatalogued (ACZCV 0528; ACZC10442; ACP4725), undetermined individual collected at Anja Reserve, 21.85223°S, 46.84404°E, elevation ca. 970 m a.s.l., on the 15 November 2018, by Francesco Belluardo, Javier Lobón-Rovira and Gonçalo M. Rosa; UADBA R-uncatalogued (ACZCV 0777; ACZC10941; ACP4991), juvenile individual collected at Tsaranoro Valley, Forêt Sacrée, 22.08491°S, 46.77545°E, elevation ca. 945 m a.s.l., on the 16 December 2018, by Francesco Belluardo, Javier Lobón-Rovira and Malalatiana Rasoazanany; and UADBA R-uncatalogued (ACZCV 0782; ACZC10953; ACP4998), female individual collected in Tsaranoro Valley, Forêt Sacrée, 22.08530°S, 46.77589°E, elevation ca. 955 m a.s.l., on the 16 December 2018, by Francesco Belluardo, Javier Lobón-Rovira and Malalatiana Rasoazanany.

Diagnosis.

Paroedura manongavato sp. nov. can be distinguished from the other species in the Paroedura genus by the presence of three broad light crossbands on the dorsum in juveniles (the first one between forelimbs, the second one at mid-body and the third one between hind limbs) versus four light crossbands in all other species, with exception of the members of the P. bastardi clade and P. oviceps . Juvenile colouration in P. vahiny is not known. It can be distinguished from P. gracilis by absence (versus presence) of a white tip on the original tail, absence (versus presence) of a raised vertebral ridge on the dorsum and shorter forelimbs, which do not extend forward beyond tip of snout (versus exceeding the snout); from P. masobe by smaller (versus distinctively large) eyes and absence (versus presence) of a dorsal row of paired spines on the original tail; from P. fasciata , P. homalorhina , P. hordiesi , P. vahiny and P. spelaea by the presence of spines on the original tail (versus absence); from P. gracilis , P. homalorhina , P. kloki , P. maingoka , P. masobe , P. oviceps (from its type locality Nosy Be), P. picta , P. spelaea , most P. tanjaka individuals and P. vahiny by the presence of prominent dorsal tubercles arranged in regular longitudinal rows (versus rather irregular rows of dorsal tubercles).

Within the Paroedura bastardi clade, P. manongavato sp. nov. is characterised by the unique combination of the following characters: (1) presence of prominent dorsal-enlarged keeled scales arranged in regular longitudinal rows, (2) presence of three broad light crossbands on the dorsum in juveniles (unknown in subadults), (3) presence of spines on the original tail, (4) nostrils separated from rostral scale by prenasals, (5) presence of a curly-bracket-shaped marking in the occipital region, (6) mediodorsal scale rows of the snout tip forming two rows of enlarged scales separated by a third median row (SnoutS, state “s”, sensu Miralles et al. (2021)), (7) prenasal scales separated by two scales in contact with rostral, (8) minimum of six interorbital scales separating the eyes, (9) 30-34 dorsal enlarged keeled scales from the middle tip of the nuchal curly-bracket to the base of the tail in adults, (10) minimum of 19 transversal dorsal-enlarged keeled scales at mid-body in adults, (11) body dorsally brown with ochre patches organised into crossbands bordered by a thin dark brown line in adults, (12) uniform colouration of toes, (13) absence of concave anterior edge in the “butterfly” pattern (sensu Miralles et al. (2021)) on the head in both juveniles and adults, (14) presence of central vacuity in the light patch on the head in both juveniles and adults and (15) body size (SVL = 68.3-73 mm).

Paroedura manongavato sp. nov. differs from P. neglecta and P. tanjaka by having the nostrils separated from the rostral scale by prenasals (versus nostrils in contact with the rostral); from P. guibeae by the presence of a curly-bracket-shaped marking in the occipital region (versus absence), by uniform colouration of toes (versus striped/bicolour toes), by mediodorsal scale rows of the snout tip forming two rows of enlarged scales separated by a third median row (versus mostly forming two rows of scales in contact; SnoutS, state “c”, sensu Miralles et al. (2021)) and by a larger adult body size (SVL> 68.3 mm versus SVL <60 mm); from P. ibityensis by a larger adult body size (SVL> 68.3 mm versus SVL <61 mm); from P. bastardi by the presence of a curly-bracket-shaped marking in the occipital region (versus absence) and by the absence of concave anterior edge in the “butterfly” pattern on the head in both juveniles and adults (versus presence only in juveniles).

Paroedura manongavato sp. nov. differs from P. rennerae , with which it is found in sympatry in Anja and Tsaranoro, by having the mediodorsal scale rows of the snout tip forming two rows of enlarged granules separated by a third median row (versus mostly forming two rows of scales in contact), by having prenasal scales separated by two scales (versus one scale), by a minimum of six interorbital scales separating the eyes (versus a maximum of five interorbital scales separating the eyes), by 30-34 dorsal-enlarged keeled scales from the middle tip of the nuchal curly-bracket to the base of the tail in adults (versus 29-31), by a minimum of 19 transversal dorsal-enlarged keeled scales at mid-body in adults (versus 15-17), by a body dorsally brown with ochre patches organised into crossbands bordered by a thin dark brown line in adults (versus a thick blackish line), by presence of a central vacuity in the “butterfly” pattern on the head in juveniles and adults (versus absence) and by having a smaller body size (SVL 68.3-73 mm versus SVL 73.6-80.9 mm). Despite an overall less spiky appearance of the dorsal body, Paroedura manongavato sp. nov. has a distinctly spikier regenerated tail than P. rennerae (Fig. 7 ). This trait, together with the dorsal colouration, seems to be the easiest way to differentiate these two species, when found in sympatry.

Description of the holotype.

Adult female (SVL = 68.3 mm) in good condition (Fig. 2B ), with two ventral incisions.

Head triangular (HW = 15.3 mm; HL = 19.9 mm; HH = 8.7 mm) and neck distinct. Rostral scale rectangular, much wider than tall and as wide as mental. Nostrils separated from rostral by prenasals. Two enlarged prenasals in contact with rostral and first supralabials, both separated by a single scale. 9/9 smooth and distinctly enlarged supralabials, followed by two pairs of smooth smaller scales, followed by two keeled scales above the mouth commissure. Canthal ridges well developed with a distinct median depression. Scales covering canthal ridges, loreal, temporal and periphery of the parietal region distinctly enlarged and slightly keeled. Eyes desiccated (ED = 5.6 mm). Dorsal head pholidosis posterior to the eyes juxtaposed, similar in size to interorbital scales (distE = 3.1 mm). Ear opening is a vertical slit (EO = 2.8 mm). 9/9 smooth and distinctly enlarged infralabials. First three infralabials slightly larger than others on both sides. Mental bell-shaped, bordered posteriorly by a pair of elongated, large and hexagonal postmentals. Each postmental in contact with six scales: mental, one postmental, first infralabial, one enlarged lateral gular, one smaller posterolateral gular and one larger central gular. Other gulars small and juxtaposed.

Scales covering the dorsal side of neck heterogeneous, with enlarged, markedly-keeled scales regularly separated from each other transversally and longitudinally by a row of 2-3 small, smooth and juxtaposed scales. Scales covering the lateral side of neck small, smooth and juxtaposed. Dorsally, 19 rows of enlarged markedly-keeled scales counted transversally at mid-body, regularly separated from each other transversally and longitudinally by a row of 1-2 smaller, keeled and juxtaposed scales. Vertebral line with a single distinct row of smaller keeled scales. Thirty-three dorsal-enlarged keeled scales from nape to tail. Ventral scales of neck, chest and abdomen flat, roundish and slightly imbricated.

Original tail (TL = 49 mm), slightly flattened dorsoventrally. Tail scales mostly keeled and very spiny. Dorsal pygal scales similar to dorsal body scales, only slightly more prominent. Regular transverse rows (whorls) with eight very spiny pygal scales per whorl. The first three whorls, with lateral pygal scales smaller and keeled. After the third whorl, lateral scales smooth. Ventral pygal section of tail with a pair of postcloacal sacs. Ventral scales of the tail between spiny pygal scales small and flat.

Dorsal scales of forelimbs and hind limbs mostly keeled. Ventral scales of forelimbs slightly smaller than surrounding ventral scales of the body. Ventral scales of hind limbs similar in size to surrounding ventral scales of the body. Hands (HAL = 7.1 mm) and feet (FoL = 9.3 mm) with proximal subdigital scales in rows of mostly two. Digits distinctly expanded at tips. One pair of squarish terminal adhesive pads. Claws curving downwards between terminal pads of digits.

After nine years in ethanol (Fig. 2 ), the overall colouration of the preserved holotype is similar to the colouration in life (Fig. 8 ). Background colours of the holotype are ochre and light brown. Melanophores are present in both the ochre and light brown body areas and their different concentrations across the body define contrasting darker brown markings. Head dorsally overall ochre with irregular light brown markings and with densely packed melanophores that define the curly-bracket-shaped transverse stripe in the nuchal region. This continues to reach the eyes on both sides. Other densely-packed areas of melanophores define two light brown blotches behind the eyes and a central vacuity in the parietal region. Altogether, this pattern of densely packed melanophores define the “butterfly” pattern on the head. Pair of light brown lateral bands running from second supralabial to the anterior corner of the eyes and from the posterior corner of the eyes to the nuchal region, where they merge with the background light brown body colouration. Body dorsally light brown with six irregular ochre patches. These are delimited by a line of densely-packed melanophores, that borders each patch externally and sometimes internally, creating irregular darker brown markings. These patches are the following (from head to cloaca): one narrow on the neck and discontinuous at mid-body (3.3 mm width along vertebral axis at mid-body), one at the forelimb insertion (4.8 mm), one posterior to the forelimb insertion along the vertebral axis (3.1 mm), one distinctly broader bow-tie-shaped at mid-body (9.1 mm), one discontinuous anterior to the hind limbs (with the aspect of three longitudinal short bands not in contact, 4.3 mm) and a very distinct patch between the hind limbs (6.3 mm). Dorsal colouration fading to ventral colouration along body flanks (this includes the irregular ochre patches). Tail colour similar to dorsum, with twelve regularly-distanced ochre whorls. These are separated by light brown whorls defined by scales bordered by one or two stripes of densely-packed melanophores. Dorsal surfaces of forelimbs and hind limbs light brown with irregular dark markings determined by densely-packed melanophores that confer a marbled appearance. Ventral colouration (throat, chest, abdomen, ventral parts of forelimbs and hind limbs) ochre. Ventral scales generally with less than ten melanophores each, number decreasing towards mid-body. Infralabials (except the first on left side and except the first three on right side) and peripheral gulars on both sides of throat pigmented for at least one third of their surface, giving an overall brownish colour. Peripheral gular scales on both sides of throat light brown and with melanophores over at least one third of their surface. Ventral surface of the tail ochre with irregular dark markings determined by densely-packed melanophores that confer a marbled appearance. Scales of the ventral surfaces of hands and feet with densely-packed melanophores, giving a dark brown colouration.

Etymology.

The specific epithet is a noun in apposition to the genus name, derived from the Malagasy words “manonga” (ma-noon-ga) meaning "to climb", and “vato” (va-too) meaning “rock”, because the species dwells on large granitic boulders. Additionally, the name evokes rock climbing, as the area, especially around Tsaranoro, has many well-known sites for this sport.

Natural history, habitat and distribution.

Based on genetically verified records, Paroedura manongavato sp. nov. is known from two localities, Anja Reserve and Tsaranoro Valley Forest ( Forêt Sacrée), which are separated by ca. 25 km and are located on the south-central plateau of Madagascar, south of the city of Ambalavao. The holotype was found at dusk active on a large boulder at the entrance of Anja Reserve. In Anja, other individuals of this species were found active both during day and night on granitic boulders within patches of semi-arid deciduous forest and in large cavities below these boulders, in a quite humid environment. The individuals ACZC10441 (ACP5940) and UADBA R-uncatalogued (ACZCV 0528; ACZC10442; ACP4725) were found at night moving on the walls of a cave-like recess created by large granitic boulders at ca. 1.5 m above the ground. ACZC10441 jumped on the leaf litter once spotted. The individuals UADBA R-uncatalogued (ACZCV 0782) and UADBA R-uncatalogued (ACZCV 0777) were found at night on boulders in the most internal part of Tsaranoro Forest. Paroedura manongavato sp. nov. seems associated with granitic boulders within semi-arid deciduous forest and it occurs in close syntopy with the morphologically similar P. rennerae .

Conservation and proposed IUCN Red List status.

The Extent of Occurrence (EOO) and Area of Occupancy (AOO) computed for this species are 3.032 km2 and 12 km2, respectively. We propose a classification of Paroedura manongavato sp. nov. as Critically Endangered (CR), under the criteria B1ab(iii) of the IUCN Red List guidelines ( IUCN Standards and Petitions Committee 2022). This proposed evaluation is based on the narrow EOO, the severely fragmented habitat where the species is encountered and the observed decline in the extent and quality of the habitat.

Figures

Figure 2. Morphological diagnostic characters and colourations of Paroedura rennerae and P. manongavato sp. nov. A morphological diagnostic characters defined and examined in this study (DorL, DorT and SeP, for definitions, see Methods section) and head pattern details (sensu Miralles et al. (2021)) of P. manongavato sp. nov. holotype B comparison of dorsal and ventral colouration of a preserved specimen of Paroedura rennerae (ZSM 8 / 2023, ACZCV 0804), after four years in 70 % ethanol and preserved holotype of P. manongavato sp. nov. (ZSM 9 / 2023, ACZCV 0300), after nine years in 70 % ethanol. Colours calibrated with ColorChecker C comparison of character SeP in P. rennerae (right) and P. manongavato sp. nov. (left). Photographs by CP and JLR. Scale bar: 10 mm (B).

Figure 4. Maximum Likelihood phylogenetic tree of the Paroedura bastardi clade inferred with the COI alignment. Bootstrap values above 80 in bold, below 70 removed. Lineages colours follow Miralles et al. (2021) and match the map, scatterplot of the PCA, haplotype networks and multilocus Bayesian phylogeny. Photograph of P. manongavato sp. nov. by JLR.

Figure 6. Multilocus Bayesian phylogeny of the Paroedura bastardi clade including the new species P. manongavato sp. nov. Colours match the lineages of map, scatterplot of the PCA, ML COI tree and haplotype networks. Photograph of P. manongavato sp. nov. by JLR.

Figure 7. Dorsal or dorsolateral views, and colourations in life of juveniles and adults of Paroedura rennerae and P. manongavato sp. nov. A dorsal or dorsolateral view of P. rennerae from Ambatomainty (male, ACP 5020; female, ACP 5021; juvenile, ACP 5022), from Anja (female, ACP 4723), from Sakaviro (juvenile, ACP 4959; female, ACP 4960) and from Tsaranoro (ACP 4975, ACP 4978, both undetermined); dorsolateral view of P. manongavato sp. nov. from Anja (undetermined, ACP 4725; juvenile, ACP 4991) and from Tsaranoro (female, ACP 4998) B dorsal colouration of P. rennerae (male, ACP 4722) from Anja Reserve C dorsal colouration of P. manongavato sp. nov. (male, ACP 5940) from Anja Reserve. Note the overall less spiky aspect of P. manongavato sp. nov. and the less contrasted ochre markings on dorsum, representing the two diagnostic characters that more likely help to discriminate between these two species when occurring in sympatry. Photographs by JLR.

Figure 8. Living holotype of Paroedura manongavato sp. nov. (ZSM 9 / 2023, ACZCV 0300), from Anja Reserve, photographed at night on the reserve boulders A dorsolateral view of the individual B dorsal view of the individual with detail on the original tail C detail of the head. Photographs by AC.