Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar Author Ratsoavina, Fanomezana M. Author Gehring, Philip-Sebastian Author Scherz, Mark D. Author Vieites, David R. Author Glaw, Frank Author Vences, Miguel text Zootaxa 2017 2017-11-14 4347 3 journal volume 31537 10.11646/zootaxa.4347.3.2 ccdea397-919f-4cfc-b0ea-e417a7ad6701 1175-5326 1048750 4D06B7E6-6FEE-42E5-9C8C-0ABAC8B0A6BE Uroplatus fotsivava sp. nov. ( Figures 4–5 ) Remark. This new species was previously referred as Uroplatus sp. F (1) by Raxworthy et al. (2008b) , as U. sp. 1 by Ratsoavina et al. (2011) , and as U. ebenaui [Ca1] by Ratsoavina et al. (2012 , 2013 , 2015 ). Holotype . ZSM 1830/2010 (field number ZCMV 12279 ), adult male , collected in the Analabe forest on the Ambodimanga mountain near Antambato village , Antsiranana Province , northern Madagascar , 14.5048°S , 48.8760°E , 1361 m above sea level, on the night of 6 June 2010 , by M. Vences , D. R.Vieites , R. D. Randrianiaina , F. M. Ratsoavina , F. Randrianasolo , S. Rasamison , A. Rakotoarison , E. Rajeriarison and T. Rajoafiarison . Paratypes. ZSM 1831/2010 (ZCMV 12280) and ZSM 639/2014 (DRV 6067), two adult males, and ZSM 1829/2010 (ZCMV 1227), an adult female, all with the same collecting data as the holotype; ZSM 646/2014 (DRV 6326), adult male, collected at Ambodikakazo, 14.2098°S, 48.8981°E, 1411 m above sea level, on the night of 16 June 2010 by F. M. Ratsoavina and F. Randrianasolo; ZSM 648/2014 (DRV 6263), adult female, collected at Vinanitelo forest, Ambinanitelo, 14.2098°S, 48.9702°E, 1280 m above sea level, on the night of 22 June 2010 by D.R. Vieites, M. Vences, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison; ZSM 647/2014 (DRV 6409), adult female, collected close to the village of Bemanevika, 14.3600°S, 48.5903°E, 1538 m above sea level, on the night of 28 June 2010 by M. Vences, D.R. Vieites, R. D. Randrianiaina, F. M. Ratsoavina, F. Randrianasolo, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison; ZSM 645/2014 (DRV 6324), adult female, collected at the Ambodikakazo campsite, 14.2098°S, 48.8981°E, 1473 m above sea level, on the night of 17 June 2010 by F. M. Ratsoavina and F. Randrianasolo; ZSM 51/2016 – 55/2016 (MSZC 57, 81, 104, 113, 141) and UADBA uncatalogued (MSZC 67 and 118), three males and four females from Ampotsidy, 15 km north of Bealanana, (14.412–14.422°S, 48.712–48.722°E), 1293–1489 m above sea level, collected at night between the 18th December 2015 and 4th January 2016 by M. D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D. H. Nomenjanahary and J. Rabearivony. Etymology. The specific epithet fotsivava is a composite of the Malagasy words for white (fotsy) and mouth (vava). The name refers to the unpigmented oral mucosa of the species, as it was the first character we used to distinguish this new species to the other forms occurring in the Tsaratanana massif. The name is an invariable noun in apposition. Diagnosis. Uroplatus fotsivava sp. nov. is included in the Uroplatus ebenaui group of small-sized leaf-tailed geckos due to its triangular head with supraocular spines, laterally compressed body, and short 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 much smaller size (adult SVL 49.5–70.7 mm versus at least 85 mm), lack of lateral dermal fringing on any part of the body, and lateral compression of the body (versus depressed body shape with lateral dermal fringes), and a relatively much shorter tail. The evident triangular head, rather smooth skin and short tail distinguish U. fotsivava from U. alluaudi , U. guentheri , U. malahelo and U. pietschmanni . Within the U. ebenaui group, the new species differs from U. finiavana , U. malama and U. phantasticus by a shorter tail (adult TAL/SVL 0.15–0.32 vs. 0.42 in U. finiavana , 0.72 in U. malama , 0.62–0.76 in U. phantasticus ) and by a narrower tail (TAW/SVL 0.07–0.10 vs. 0.14–0.16 in U. finiavana , 0.24 in U. malama , 0.16–0.20 in U. phantasticus ). It is further distinguished from U. ebenaui , U. malama and U. phantasticus by its unpigmented oral mucosa (versus at least partly blackish pigmented oral mucosa). Uroplatus fotsivava shares an unpigmented oral mucosa with its sister species U. fiera , and has overlaps in values of relative tail length and tail width with this species. However, the ratio TAW/TAL distinguishes almost all specimens of the two lineages (0.17–0.31 in U. fotsivava vs. 0.19–0.21 in U. fiera ; except for MSZC 0 118, all specimens of U. fotsivava have values of 0.23 or larger). Furthermore, U. fotsivava in comparison to U. fiera has a higher number of supralabials and associated scales (28–31 versus 27–28; U-test, P = 0.005) and a tendency of fewer lamellae under the third toe (5–8 versus 7–8; U-test, P = 0.065). Description of the holotype. Adult male in good condition with an intact tail and everted hemipenes. SVL 56.0 mm, tail length 16.2 mm, maximum tail width 4.2 mm, for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis recognizable and concave; snout sloping strongly and continuously downward anteriorly; snout weakly depressed, short (1.2 times longer than eye diameter); eyes large (eye diameter 4.6 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.0 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 but with miniscule spines on the hindlimbs, and practically no spines on the forelimbs; forelimb reaches beyond tip of snout when adpressed forward and approaches the groin when adpressed backwards along body (forelimb length/axilla–groin distance 25.2/26.6 mm = 95%), hindlimb reaches beyond axilla when adpressed forward along body (hindlimb length/axilla–groin distance 30.7/26.6 mm = 115%); original tail length 29% of snout–vent length, membranous borders of the denticulated tail more or less symmetrical, somewhat trapezoidal (i.e. broadening on either side and then narrowing again posteriorly), completely absent from the slightly spatulate tail-tip. Nares separated from each other by at least nine small granular scales, from the first supralabial by one scale, and from the rostral scale by one scale; first supralabial taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 22/20 (right/left) true supralabials and 21/20 infralabials (as defined in methods above); 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 interspersed with a few larger and partly raised tubercles, except for the irregular lines on the head and body which consist of series of slightly enlarged scales. 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, angular and medially straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Two additional transverse lines, consisting of slightly enlarged scales, are recognizable on the frontal region of the head. Numerous spines on the posterior and postorbital part of the head, on hindlimbs (ca. 30 per limb), a dermal flap on each knee, bearing a spine; no flaps or spines on elbow and forelimb; a prominent pointed flap on the posterior portion of each upper eyelid (supraocular spine); posterior border of eye fringed. Weakly hollowed axillary pits present. Scales of the posterior ventral abdomen enlarged and arranged more uniformly than other scales, almost forming proper rows. Hemipenes everted, bearing two lobes. Calyx with protuberance bearing honeycomb-like structures, especially on the asulcal side. Area of sulcus spermaticus is smooth. Each lobe with a dense field of pointed papillae at its apex. Coloration. After six and a half years of preservation in 70% ethanol the color pattern remains the same as in the living animal ( Fig. 5 ) but its vivacity and intensity have faded. All dorsal surfaces are mottled beige in base color, with individual flecks of burnt umber. The tail is as the dorsum at its base, but beyond the hemipenes is dorsally lighter beige, with few small brownish spots. The dorsum has weakly defined posteroventrally oriented lighter and darker beige chevrons, running parallel to the ribs beneath them. Several burnt umber flecks are noteworthy, especially an hourglass-shaped marking on the snout, and a strong dark fleck on the mid-dorsum, as well as several asymmetrical flecks on other parts of the body, including the arms, legs, and nape. Several whitish flecks that were visible on the animal in life ( Fig. 4 ) are no longer visible in preservative, particularly those on the flank. The V-shaped area on the nape is almost indistinguishable. Two light cream markings defined by brown borders are present below each eye, one at the posterior corner and one at the mid-eye, both descending to the supralabials. The supralabials are generally beige in color, but a few are burnt umber. The infralabials weakly alternate between cream and burnt umber. The venter, including the ventral limbs except the lower hindlimbs, is a lighter beige than the dorsum, and much less mottled. The venter bears several round cream spots defined by brown borders, with three pairs of such markings in the pectoral region, and other such spots on the posterior abdomen. The tail is slightly darker below than above, and has a few small cream flecks bordered with brown, and a few burnt umber spots. The oral mucosa is unpigmented and whitish in preservative. In life the eye was golden brown with dark striations. Variation. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2–26, and tends to be associated with the overall spiny-ness of the specimens; those with more head spines tend to have more arm and leg spines as well, and generally have more heterogeneous scalation. Forelimbs can bear no spines, or as many as six, with at most a slightly raised scale on the elbow but never a clear spine. Hindlimbs can bear 2–30 spines, and the dermal knee flap almost always bears a spine. In males the tail is highly denticulated and tends to be asymmetrical, but it can be symmetrical as well; in females it is without denticulations and trapezoidal; this might reflect a general pattern of sexual dimorphism in tail shape in the U. ebenaui group that warrants further investigation (see also Ratsoavina et al. 2015 ). Internarial scales range from eight to nine, supralabials from 18–22, and infralabials from 17–21. The line on the head connecting the supraocular spines can be curved to a point, or almost straight. The coloration of the type series is immensely variable, and includes browns to reds, with some specimens being strongly contrastingly colored, while others have almost homogeneous brown coloration, sometimes with dark and light markings. Ventral coloration is equally variable, with the venter sometimes being cream, sometimes being brown, and the chin sometimes bearing weak posteriorly-oriented chevrons. The V-shape on the nape can be extremely dark and thus highly contrasting, or hardly noticeable. Cream markings below the eye can be present or absent. The identity of all paratypes from Ampotsidy (MSZC series) was ascertained by sequencing a fragment of the mitochondrial 12S rRNA gene (see Results); they showed no or negligible differentiation compared to the samples included in our main molecular phylogenetic analysis. ZSM 52/2016 (MSZC 0141) clustered in a separate haplotype lineage with ZSM 648/2014 from Ambinanitelo, whereas all other Ampotsidy samples clustered in the Analabe/Bemanevika haplotype lineage. TABLE Ɩ. Measurements in mm, anđ two scale counts (SUPL anđ LAM3T) taken ƭrom all Uroplatus specimens incluđeđ in this stuđy. HT, holotype; PT, paratype; NA, not available, NM, not measuređ. See Materials anđ Methođs ƭor other abbreviations. Specimens without museum numbers will be catalogueđ in UAĐBA. All measurements taken by E.M. Ratsoavina except those markeđ with * (measurements anđ counts taken by M. D. Scherz), ** (measurements taken by A. Miralles), # counts taken by stuđent assistants.

Locality	Sex	Museum number	Field number	SVL	TAL	TAW	HLƖ	HL2	HW	FORL	HIL	NTL	IRPL	SUPL #	LAM³T #
U. fotsivava sp. nov.
Tsaratanana (Analabe)	male	ZSM 639/2014 (PT)	ĐRV 6067	ƽ9.2	16.9	4.4	17.8	16.1	13.4	23.6	23.ƽ	9.3	2.7	29	7
Tsaratanana (Ambođikakazo)	male	ZSM 646/2014 (PT)	ĐRV 6326	68.0	21.6	6.7	20.7	18.9	14.9	27.2	33.6	7.8	NM	31	8
Tsaratanana (Ambinanitelo)	ƭemale	ZSM 648/2014 (PT)	ĐRV 6263	70.7	10.7	NA	19.7	17.8	14.6	26.1	3ƽ.8	11.ƽ	2.9	30	7
Bemanevika	ƭemale	ZSM 647/2014 (PT)	ĐRV 6409	ƽ7.2	14.ƽ	4.ƽ	17.4	1ƽ.8	13.4	23.0	30.6	11.2	2.3	28	7
Tsaratanana (Ambođikakazo)	ƭemale	ZSM 64ƽ/2014 (PT)	ĐRV 6324	70.2	18.0	ƽ.7	19.ƽ	18.2	1ƽ.2	27.7	36.4	12.2	2.6	29	7
Tsaratanana (Analabe)	ƭemale	ZSM 1829/2010 (PT)	ZCMV 12277	68.8	19.2	ƽ.0	19.ƽ	18.9	14.6	26.0	33.3	12.8	1.8	28	ƽ
Tsaratanana (Analabe)	male	ZSM 1830/2010 (HT)	ZCMV 12279	ƽ6.0	16.2	4.2	18.1	16.3	13.6	2ƽ.2	30.7	8.3	1.9	29	7
Tsaratanana (Analabe)	male	ZSM 1831/2010 (PT)	ZCMV 12280	67.3	18.3	4.9	19.1	18.1	1ƽ.ƽ	2ƽ.8	38.2	10.3	3.1	30	6
Ampotsiđy	ƭemale	* UAĐBA uncatalogueđ (PT)	MSZC 0 118	63.ƽ	17.8	3.1	18.0	16.ƽ	13.1	2ƽ.2	29.0	13.7	ƽ.0	NM	NM
Ampotsiđy	ƭemale	* UAĐBA uncatalogueđ (PT)	MSZC 0 0 67	60.3	NA	NA	18.1	12.0	14.6	19.0	26.1	8.1	1.6	NM	NM
Ampotsiđy	male	* ZSM ƽƽ/2016 (PT)	MSZC 0 113	ƽ2.3	16.7	3.9	1ƽ.4	14.6	11.8	19.4	24.8	14.ƽ	1.0	NM	7
Ampotsiđy	male	* ZSM ƽ2/2016 (PT)	MSZC 0 141	ƽ9.9	17.6	4.ƽ	18.0	16.9	13.8	23.6	28.4	8.2	2.1	NM	7
Ampotsiđy	ƭemale	* ZSM ƽ4/2016 (PT)	MSZC 0 0 81	66.2	16.8	4.ƽ	20.1	18.2	1ƽ.8	23.7	28.6	11.ƽ	2.2	NM	7
Ampotsiđy	ƭemale	* ZSM ƽ1/2016 (PT)	MSZC 00ƽ7	69.1	16.7	ƽ.2	18.ƽ	17.4	14.1	24.ƽ	29.6	11.3	2.ƽ	NM	6
Ampotsiđy	male	* ZSM ƽ3/2016 (PT)	MSZC 0 104	49.ƽ	NA	NA	1ƽ.9	14.9	11.7	19.ƽ	24.3	7.6	2.3	NM	7
U. kelirambo sp. nov.
Tsaratanana Camp 2	male	ZSM 641/2014 (HT)	ĐRV 6192	ƽ7.3	2ƽ.4	2.ƽ	18.3	16.6	13.4	24.6	30.8	1ƽ.1	4.3	30	7
Tsaratanana Camp 2	male	ZSM 642/2014 (PT)	ĐRV 62ƽ0	ƽ3.1	20.7	2.3	16.8	1ƽ.9	12.8	23.8	29.7	9.9	3.7	28	7
Tsaratanana Camp 2	male	ZSM 643/2014 (PT)	ĐRV 62ƽ1	62.8	23.4	3.1	17.9	16.8	13.7	27.1	34.9	9.4	2.7	27	7
Tsaratanana Camp 2	ƭemale	ZSM 1832/2010 (PT)	ZCMV 12388	63.4	22.3	2.6	17.2	16.8	14.4	2ƽ.2	31.4	10.7	2.4	30	6
U. fiera
Eierenana region	male	ZSM 1126/2003 (PT)	EGMV 2002.3097	63.6	18.1	3.6	20.1	17.7	14.2	20.6	2ƽ.9	8.1	2.8	27	8
Eierenana region	male	ZSM 211/2002 (PT)	MV 2001.1403	ƽƽ.9	19.1	3.7	17.7	17.0	13.9	23.9	33.8	10.1	2.0	27	7
Ambatovy region	ƭemale	UAĐBA-R 708ƽ0 (PT)	ERC 6ƽ1	67.2	18.ƽ	3.9	20.3	18.ƽ	1ƽ.3	28.1	3ƽ.ƽ	11.3	3.2	27	7
Eierenana	male	ZSM 212/2014 (PT)	ERC 618	60.4	17.ƽ	3.4	17.6	16.4	13.8	28.6	34.2	6.6	2.2	28	8
Eierenana	male	ZSM 213/2014 (HT)	ERC 619	66.2	19.9	3.8	19.6	19.2	1ƽ.3	29.1	36.3	9.7	3.1	27	7
Ambatovy region	male	UAĐBA-R 70849 (PT)	ERC 6ƽ0	78.0	24.4	4.7	21.8	20.3	16.6	30.1	41.1	8.3	3.2	28	8

……continued on the next page TABLE Ɩ. (Continueđ)

Locality	Sex	Museum number	Field number	SVL	TAL	TAW	HLƖ	HL2	HW	FORL	HIL	NTL	IRPL	SUPL #	LAM³T #
U. finiavana Montagne đAAmbre	male	ZSM 113ƽ/2003 (PT)	EGMV 2002.3083	ƽ8.0	NA	NA	18.8	17.2	13.1	26.1	34.6	10.7	2.7	30	6
Montagne đAAmbre	ƭemale	ZSM 1134/2003 (PT)	EGMV 2002.2390	ƽ2.0	24.8	ƽ.6	1ƽ.2	1ƽ.1	11.4	22.4	30.1	9.8	2.8	29	6
Montagne đAAmbre	male	ZSM 1133/2003 (PT)	EGMV 2002.2388	63.ƽ	32.8	6.8	18.0	17.ƽ	13.3	2ƽ.0	34.ƽ	NA	NA	29	6
Montagne đAAmbre	male	ZSM 1132/2003 (PT)	EGMV 2002.2387	61.0	NA	NA	18.8	17.8	13.2	27.0	3ƽ.8	10.0	1.8	30	7
U. phantasticus Eierenana region	male	ZSM 1127/2003	EGMV 2002.3100	ƽ1.7	39.1	9.6	1ƽ.7	14.8	11.7	21.3	29.9	7.1	1.2	30	ƽ
Eierenana region	male	ZSM 1128/2003	EGMV 2002.3103	ƽ8.8	44.4	11.4	17.6	17.1	12.6	24.ƽ	34.9	10.1	2.3	30	6
Eierenana region	male	ZSM 212/2002	MV 2001.1402	ƽƽ.3	40.0	10.8	16.6	16.1	13.1	24.9	34.7	14.1	2.1	31	6
AnBozorobe	ƭemale	ZSM 471/2010	EGZC 4341	ƽƽ.1	34.5	10.1	1ƽ.7	14.7	12.2	21.8	26.1	11.4	1.7	28	6
AnBozorobe	male	ZSM 472/2010	EGZC 438ƽ	ƽ8.3	NA	NA	16.7	1ƽ.7	11.6	24.6	33.2	4.1	NM	27	ƽ
Tarzanville	male	ZSM 473/2010	EGZC 4ƽ0ƽ	66.3	46.0	12.4	18.9	18.2	13.8	26.8	36.6	12.1	2.9	31	7
Ranomaƭana	male	ZSM 934/2003	EGMV 2002.14ƽ2	60.4	37.6	9.8	17.8	17.2	13.0	2ƽ.ƽ	30.1	8.4	2.7	29	6
Ranomaƭana	ƭemale	ZSM 93ƽ/2003	EGMV 2002.14ƽ3	76.1	NA	NA	20.8	19.8	1ƽ.7	29.9	39.9	16.ƽ	2.9	29	6
Emerina	ƭemale	**BMNH 1946.8.26.64 (HT)	ƽ6.8	NA	NA	16.8	NM	11.6	21.4	31.1	NM	NM	NM	NM
U. ebenaui Eorêt đAAmbre	male	ZSM 2028/2008	FGZC 1876	49.6	18.6	4.2	1ƽ.4	14.7	11.8	19.2	27.2	3.4	0.9	32	6
Eorêt đAAmbre	female	ZSM 2029/2008	FGZC 3116	59.8	12.9	3.ƽ	17.7	16.7	13.3	22.ƽ	30	2.2	0	32	5
Eorêt đAAmbre	male	ZSM 2030/2008	FGZC 31ƽ3	4ƽ.6	1ƽ.4	4	1ƽ.1	13.ƽ	11.6	18.3	27.4	1.7	1.1	29	6
Nosy Be	ƭemale	ZSM 2498/2009	ZCMV 13013	63.1	16.1	4.1	19.1	17.2	14.4	21	32.1	2.ƽ	0.2	28	6
Manongarivo	male	ZSM 8ƽ6/2003	EGMV 2002.826	ƽ6.ƽ	22.ƽ	3	16.9	16.6	13.4	23.8	34.2	1.8	0.6	29	6
Manongarivo	male	ZSM 997/2003	EGMV 2002.2403	ƽ1.4	16.4	2.ƽ	1ƽ.8	14.ƽ	11.6	19.2	30	3	0	29	6
Nosy Be	male	NA	EGZC 3747	ƽƽ.ƽ	12.1	3.3	17.3	11.1	13.1	23.6	31.1	4.3	0	29	6
Eanambana	male	UAĐBA-R 708ƽ1	ERC 611	ƽ4.1	19.7	3.6	20.6	1ƽ.3	12.3	20.1	30.7	1.1	0.0	28	6
U. malama Beampingaratra /Anđreoky	male	ZSM 2860/2010	ZCMV 12128	77.ƽ	ƽ6.1	18.4	21.3	20.6	1ƽ.4	31.2	41.9	4.2	1.ƽ	28	7

FIGURE 4 . Photographs of specimens of Uroplatus fotsivava sp. nov. in life. (a) Male holotype ZSM 1830/2010 (ZCMV 12279) from Analabe; (b) male paratype, ZSM 639/2014 (DRV 6067) from Analabe; (c-d) female paratype, ZSM 647/2014 from Bemanevika; (e) showing unpigmented oral mucosa as typical for the species; (f) female paratype ZSM 54/2016 (MSZC 81) from Ampotsidy; (g) female paratype ZSM 51/2016 (MSZC 57) from Ampotsidy; (h) female paratype MSZC 118 (to be catalogued in UADBA) from Ampotsidy. Distribution. Summarizing all sites vouchered by molecular data, the species is known from six sites, all in mid- to high-elevations of the Tsaratanana area: (1) the type locality, Analabe forest; (2) Ambodikakazo; (3) Ambinanitelo; (4) Bemanevika; (5) Ampotsidy; and (6) Manarikoba forest (Antsahamanara campsite; located at 1100 m above sea level; Andreone et al. 2009 ). Taken together, these localities span an elevational range of 1100– 1538 m a.s.l. Natural history. The holotype was found active and perching at night on a tree branch of the Analabe forest, as were the remaining specimens from this locality. The forest is isolated at the top of the Ambodimanga mountain, with some degradation especially towards the edge. It is a humid evergreen forest characterized by a few big trees and dense understory, and a series of small streams running down the mountain. In Ampotsidy individuals of the species were typically found in bushes and small trees, up to a maximum of four meters above the ground. Several females encountered in December 2015 were gravid, suggesting a mating period coinciding with December to January, or the start of the local rainy season.