Finaritra! A splendid new leaf-tailed gecko (Uroplatus) species from Marojejy National Park in north-eastern Madagascar Author Ratsoavina, Fanomezana Mihaja Author Raselimanana, Achille P. Author Scherz, Mark D. Author Rakotoarison, Andolalao Author Razafindraibe, Jary H. Author Glaw, Frank Author Vences, Miguel text Zootaxa 2019 2019-01-21 4545 4 563 577 journal article 27671 10.11646/zootaxa.4545.4.7 6e699de7-3402-497b-ac7d-cf031d53b703 1175-5326 2618341 BBD8A9FD-BC9B-42D2-AD7D-CEB71ADF17DF Uroplatus finaritra sp. nov. Figs. 1–5 , Table 1 Uroplatus sp. G—Raxworthy et al. (2008) Uroplatus sp. Ca9— Ratsoavina et al. (2013) LSID: urn:lsid:zoobank.org:act: E6A1407B-7C9E-46F6-8FEC-D120A111A344 Holotype . UADBA-R 70489 (field number APR 12522), adult female, collected on the eastern slope of Marojejy by a tributary of the Ampanasatongotra River , Marojejy National Park , Sava Region , northern Madagascar , at ca. 14.4559°S , 49.7749°E ( 780 m a.s.l. ) on the night of 15 May 2016 by A. P. Raselimanana. Paratypes . UADBA-R 70493 (field number APR 12591), adult male , UADBA-R 70490 ( APR 12590), adult male, and UADBA-R 70491 ( APR 12589), adult female, three specimens collected in the same locality as the holotype ( 750–810 m a.s.l. ) on 18 May 2016 by A. P. Raselimanana ; UADBA-R 70492 ( APR 12691), adult male, and UADBA-R 70494 ( APR 12692), juvenile female, two specimens collected on the western slope of Marojejy by Tsiasisa River , 2.2 km east of Antanimbaribe , Marojejy National Park , Sava Region , northern Madagascar , at ca. 14.5074°S , 49.6153°E ( 845 m a.s.l. ) on the night of 28 May 2016 by A. P. Raselimanana ; UADBA-R 70501 ( MSZC 0253 ), subadult female , ZSM 458 /2016 ( MSZC 0271 ), subadult female, two specimens collected near the path below Camp 1 ( Camp Mantella ) of Marojejy National Park , Sava Region , northern Madagascar , at ca. 14.438°S , 49.776°E ( 450 m a.s.l. ) on 21–23 November 2016 by M. D. Scherz , M. Bletz , A. Rakotoarison , J. Razafindraibe , A. Razafimanantsoa , and M. Vences . Etymology. Finaritra is a Malagasy word for special greetings but also means healthy and happy. We refer to our delight in describing this splendid and exceptionally large species from a clade of generally small-sized leaftailed geckos. The name is an invariable noun in apposition. Diagnosis. Uroplatus finaritra sp. nov. is assigned to the Uroplatus ebenaui group based on its relatively small size and its triangular head with supraocular spines, and overall leaf-mimicking aspect. It is characterised by (1) a dark red oral mucosa, (2) a comparatively large body size ( 82.5–95.3 mm SVL in adults), (3) 7–8 lamellae (exceptionally 6) under the third finger and toe, and (4) a long (length 53–65% of SVL), wide (width 16–18% of SVL), leaf-shaped tail. It differs from all members of the U. fimbriatus group ( U. fimbriatus , U. giganteus , U. henkeli , U. sikorae , and U. sameiti ) and U. lineatus by its smaller size (SVL 82.5–95.3 mm vs. 85–200 mm ) and the lack of lateral dermal fringes on any part of the body (vs. presence except in U. lineatus ). Also, Uroplatus finaritra sp. nov. diverges from the species of the U. fimbriatus group by their lack of lateral body compression. The prominent triangular head, supraocular spines, and rather smooth skin distinguish Uroplatus finaritra sp. nov. from U. alluaudi , U. guentheri , U. malahelo and U. pietschmanni . Amongst the U. ebenaui group, the rather long tail (TAL/SVL 0.53–0.68) differentiates Uroplatus finaritra sp. nov. from U. ebenaui , U. fiera , U. fotsivava , and U. kelirambo (TAL/SVL 0.22–0.44), and furthermore it differs from U. fiera and U. fotsivava by a pigmented oral mucosa (vs. unpigmented). The new species differs from U. finiavana by a wider tail (TAW/SVL 0.16–0.18 vs. 0.05–0.14), and by its pigmented oral mucosa (vs. unpigmented). By its large tail and pigmented oral mucosa, the new species is most similar to U. malama and U. phantasticus . It differs from U. phantasticus by larger size (SVL 83–95 mm vs. 52–76 mm ), a generally shorter tail (TAL/SVL 0.53–0.65 vs. 0.62–0.76), a typically higher number of lamellae under the third finger and toe (6–8 vs. 5–7), and the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black). It differs from U. malama by a spinier integument in some adult males, by the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black), and probably by a narrower tail (TAW/SVL 0.16–0.18 vs. 0.24). In the mitochondrial phylogeny, the new species occupies an isolated position within the U. ebenaui group, without affinities to the morphologically similar U. phantasticus and U. malama (which are not each other's closest relatives either). The nuclear gene c-mos reveals haplotype sharing of the new species with U. kelirambo and two undescribed candidate species ( U. sp. Ca3 and Ca4) occurring in the same region of Madagascar , but morphologically very different by their short tails. TABLE 1 . Morphological measurements and scale counts of Uroplatus finaritra sp. nov. For abbreviations used, refer to Methods; additional abbreviations: M, male; F, female; sa, subadult; j, juvenile; HT, holotype ; PT, paratype ; na, not applicable (due to missing original tail); nm, not measured. All measurements in mm.

Museum Number	Field number	Sex	SVL	TaL	TAW	HL2	AGD	ED	EARD	SL	HL1	HW	FORL	HIL	NTL	IRPL	INS	SUPL	INFL	LAM3F	LAM3T
ZSM 458/2016	MSZC 0 271	sa F (PT)	71.2	48.4	13.8	15.6	34.2	5.4	1.0	9.2	21.9	15.1	28.5	43.1	10.6	1.1	8	28/28	23/26	8	8
UADBA-R 70493	APR 12591	M (PT)	91.3	na	na	24.2	49.1	7.2	0.9	10.5	25.7	19.4	35.4	47.8	8.3	1.5	10	26/25	22/23	7	8
UADBA-R 70492	APR 12691	M (PT)	82.5	53.8	14.2	21.4	40.5	6.8	1.2	9.5	22.5	17.7	34.5	42.9	7.3	1.1	9	26/29	24/24	8	8
UADBA-R 70490	APR 12590	M (PT)	94.7	56.4	15.3	22.1	42	6.6	1.3	9.4	23.2	18.1	33.0	46.6	9.6	1.7	10	25/27	21/21	8	8
UADBA-R 70491	APR 12589	F (PT)	93.7	51.4	16.8	23.4	50.5	6.7	1.6	10.2	24.9	19.2	32.2	46.2	13.2	1.7	9	25/26	21/21	7	8
UADBA-R 70489	APR 12522	F (HT)	95.3	50.9	17.0	25.3	50.2	6.5	1.1	10.8	26.1	20.2	33.4	41.4	9.9	0.7	9	25/26	25/25	7	8
UADBA-R 70494	APR 12692	j F (PT)	47.1	25.7	7.6	12.1	26.6	4.4	0.6	5.5	13.6	10.3	17.7	26.8	nm	nm	10	26/27	23/21	6	7
UADBA-R 70501	MSZC 0 253	sa F (PT)	59.5	38.2	9.8	16.2	27.6	5.8	0.8	7.5	18.1	13.7	23.9	35.0	nm	nm	10	29/29	25/21	7	8

FIGURE 1. (a) Bayesian inference phylogenetic tree of the Uroplatus ebenaui group based on a 2326 bp alignment of four concatenated mitochondrial gene fragments (ND4, 12S rRNA, 16S rRNA, COX1). Only posterior probability values of 0.94 or greater are displayed at nodes. An outgroup ( U. alluaudi ) was used for rooting but omitted from the tree for graphical reasons. (b) Haplotype network independently calculated from sequences of the nuclear gene c-mos. FIGURE 2. Holotype of Uroplatus finaritra sp. nov. (UADBA-R 70489, APR 12522) in dorsal view (left) and a paratype (UADBA-R 70491, APR 12589) in ventral view (right) view. Scale bar indicates 10 mm. Description of the holotype . Adult female in fair condition, with a fully intact tail, both of its hind limbs fractured ( Fig. 2 ). SVL 95.3 mm , tail length 50.9 mm , maximum tail width 17.0 mm; for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis indistinct and concave; snout sloping weakly downwards anteriorly; snout weakly depressed, fairly long (1.7 times longer than eye diameter); eyes large (eye diameter 6.5 mm ), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.1 mm ), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in ventral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and very few, miniscule spines on the hind limbs (one on the knee, as well as a small flap of skin), and two miniscule spines on the forelimb (on the elbow and near the wrist); forelimb reaches the canthus rostralis when adpressed forward and midbody when adpressed backwards along body (forelimb length/axilla–groin distance 33.4/ 50.2 mm = 67%), hindlimb reaches ¾ up the body when adpressed forward along body (hindlimb length/axilla–groin distance 41.4/ 50.2 mm = 83%); original tail length 53% of snout–vent length, membranous borders of the tail symmetrical, without any emarginations, long and leafshaped, with a thin, borderless spatulate tip. Toes bear small claws, with the distal phalange not much wider than the rest of the digit; the third finger bears 7 lamellae, the third toe 8 lamellae. FIGURE 3. Variation among specimens of Uroplatus finaritra sp. nov. from Marojejy National Park. (a) photo probably showing paratype UADBA-R 70494 (APR 12692), (b) paratype UADBA-R 70491 (APR 12589), (c) paratype UADBA-R 70490 (APR 12590), (d) holotype UADBA-R 70489 (APR 12522), (e) paratype UADBA-R 70501 (MSZC 0253), (f) paratype ZSM 458/2016 (MSZC 0271). All specimens shown except (c) are females; note the differences in tail notching between this specimen and the others, and the presence of hemipenial bulges. Nares separated from each other by at least 10 small granular scales, from the first supralabial by 3 scales, and from the rostral scale by 5 scales; first supralabial not taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 25/26 (right/left) supralabials and 25/25 infralabials; no enlarged postmental scales or chin shields; dorsal and ventral scales of head, neck, body, limbs, and tail small, granular, juxtaposed and largely of uniform size, except on the posterior ventral abdomen, where they are slightly larger than the rest of the body, and arranged almost uniformly. Two curved lines (rows of slightly enlarged scales) extending from the posterolateral parts of the head (nuchal region) converge on the neck forming a V-shaped pattern (neck triangular line). A similar, moderately distinct, fairly straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Another, less distinct line is present connecting the anterior angle of the eyes over the snout in a bowed line. The body possesses very few dermal spines—a prominent pointed supraocular spine, a single spine at the posterior angle of the head, and a small number of spines on each limb (described above). No axillary pits present. Colouration . After two years of preservation in 70% ethanol the colour is slightly faded ( Fig. 2 ). All dorsal surfaces of the specimen are a ruddy brown, fading to a salmon brown in the neck region and at the insertion of the hind limbs. A fine network of blackish lines is present across the body, with the following notable features: a somewhat straight line connecting the suprocular spines (described above); a v-shaped marking on the neck, which extends into a vertebral stripe that reaches all the way to the tail tip; a number of posteriorly oriented chevrons from the mid-dorsum to the belly, becoming thinner posteroventrally, three of which are most distinct. The exterior of the shank and outermost two toes are grey-black, whereas the inner foot is brown. The dorsal snout is darker than the rest of the head. The tail is not different in colour or pattern from the dorsal body. A distinct lateral line runs along the ventrolateral edges of the abdomen from the insertion of the leg to the axilla. The ventral colouration is uniformly orange-brown, with a faint, symmetrical pattern of dark lines on the chin. The soles of the hands and feet are grey. The ventral tail is the colour of the trunk mottled with grey splotches. A faint cream tear-like marking is present at the posterior corner of the eye above the supralabial. The oral mucosa is dark red, with a pink tongue and cream under the eyes and along the lower jaw. In life, the iris colour was silver externally and rusty around the pupil ( Fig. 3d ). Variation . Molecular data are also available from a previous paper ( Raxworthy et al. 2008 ) for one specimen (RAN 42274) from Marojejy as Uroplatus sp. G, included for 12S and cytochrome b as U . sp. Ca 9 in Ratsoavina et al. (2013) . The available sequence for 12S agrees with those of our specimens. The paratypes were sexed as adult males by the presence of everted hemipenes (UADBA-R 70493) or presence of distinctly enlarged tail base (UADBA-R 70490 and 70492), and distinctly notched tail edges; or as females based on the absence of hemipenes or hemipenial bulges, presence of visible oviduct or ovary structures after dissection, and generally smooth tail edges. In the smallest specimen (UADBA-R 70494) female inner reproductive organs were observed but were only weakly developed, and we therefore consider this specimen as juvenile. The sexual maturity of two further specimens (UADBA-R 70501 and ZSM 458/2016) is unclear, as they are considerably smaller than the large adult females (UADBA-R 70491 and 70489) but have similar gonad development. We tentatively consider these to be subadults. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2–9, and is more in males than in females, as males are more spiny than females in general, and have more heterogeneous scalation. Forelimbs can bear a minimum of two spines, or as many as eight, with a small spine present on the elbow. Hind limbs can bear 2–15 spines, and the dermal knee flap almost always bears at least a small spine, which is smaller in females than males. In males the tail is moderately emarginated, with semicircular emarginations that tend to be asymmetrical. In females the tail is without emarginations and has smooth edges. Internarial scales range from nine to ten, supralabials from 25–29, and infralabials from 21–25. The third finger bears 7–8 lamellae ( 6 in one juvenile specimen, UADBA-R 70494), the third toe bears 7–8 lamellae. The colouration is varied, as is commonly the case in Uroplatus species ( Figs. 3–4 ). The line on the head connecting the supraocular spines is generally bowed, but can be slightly pointed posteriorly, and in UADBA-R 70492, it is irregular at its midpoint. The dark vertebral stripe is only present in the holotype , UADBA-R 70491 and 70494, and in other specimens is either absent, or is cream and not dark (in UADBA-R 70492 and 70490). Overall dorsal colouration differs dramatically, from almost completely reddish brown (UADBA-R 70501) to highly mottled greys and blacks (UADBA-R 70493). Irregular flecks can be present or absent on any part of the body. The different colouration of the outer shank and foot is always present, and a cream tear-like marking below the eye is almost always present (absent in UADBA-R 70494 and 70501). The ventrolateral stripe is always present but not always distinct (indistinct in UADBA-R 70493). Ventral colouration is as variable as dorsal colouration. UADBA-R 70491 is extremely similar to the holotype , and to a lesser degree 70492, 70494, and 70501 resemble the holotype ventrally. UADBA-R 70490 and 70493 differ dramatically in being darker in colour, having irregular flecks of cream and reddish brown, and lacking the symmetrical markings on the chin. In life, the colouration of the iris is typically silvery but can be red ( Figs. 3c, f ), and the oral mucosa was consistently dark red ( Fig. 5 ). Distribution . At present this species has only been collected on the Marojejy Massif, at altitudes from 450–845 m a.s.l. Natural History. The holotype was collected active at night on a stem 0.5 m above the forest floor in nearly closed canopy humid forest at the bottom of a slope, in the vicinity of a Uroplatus lineatus specimen. UADBA-R 70490, 70491, and 70493 were collected on small branches and leaves between 1.5 and 2.5 m above the ground in similar forest. UADBA-R 70492 and 70494 were also active on branches 3–4.5 m from the forest floor in nearly closed-canopy humid forest with a thick understory and ferns. UADBA-R 70501 and ZSM 458/2016 were collected active at night on thin branches (up to 4 m above the ground) in closed-canopy, low-altitude rainforest. We consider the various collection localities to represent a single Threat-Defined Location in the sense of the International Union for the Conservation of Nature ( IUCN 2012 ), as we have recently done for several frog species from low- to mid-elevation forests in Marojejy ( Scherz et al. 2016 , Rakotoarison et al. 2017 ). In line with these assessments, we recognise that there is on-going habitat degradation due to illegal logging activity, especially at low altitudes in and around Marojejy National Park. The Extent of Occurrence and Area of Occupancy of the new species is certainly greater than 10 km 2 , but is likely less than 500 km 2 , and as such we recommend a status of Endangered for it under criterion B1ab(iii).