Lygodactylus winki Vences & Multzsch & Gippner & Miralles & Crottini & Gehring & Rakotoarison & Ratsoavina & Glaw & Scherz 2022, sp. nov

Lygodactylus winki sp. nov. Lygodactylus sp. 18— Gippner et al. (2021). Holotype. ZSM 47 /2016 (MSZC 0075), adult male, collected by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D.H. Nomenjanahary, and J. Rabearivony, at Ampotsidy, 15.7 km NNW of Bealanana (8.7 km NNW of Beandrarezona), northern Madagascar, at geographical coordinates S14.41900, E48.71883, 1364 m a.s.l., on 22 December 2015 (Fig. 15). Paratypes. ZSM 48 /2016 (MSZC 0110), adult male, collected by same collectors and at same locality as holotype, at geographical coordinates S14.42843, E48.72285, 1315 m a.s.l., on 29 December 2015; UADBA-R 70856– 70859 (MSZC 0011, 0019, 0023, 0077), two males and two females, respectively, collected by same collectors and at same locality as holotype, between coordinates S14.41455 –14.42317, E48.71149 –48.71916, 1320–1404 m a.s.l., on 18–22 December 2015; ZSM 1763 /2010 (ZCMV 12502), by M. Vences, D. R. Vieites, R. D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison, at Bemanevika, Antsirakala campsite, geographical coordinates S14.43061, E48.60179, 1466 m a.s.l., on 27 June 2010; ZSM 555 /2014 (DRV 6288), collected by F.M. Ratsoavina, D. R. Vieites, M. Vences, R. D. Randrianiaina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison at Andrevorevo, geographical coordinates S14.3464, E49.1028, 1717 m a.s.l., on 21 June 2010. Diagnosis. Lygodactylus winki sp. nov. corresponds to a lineage forming part of the subclade A5 of Domerguella, and is one of only two representatives of this subclade known to reach northern Madagascar. The lowest genetic divergences of the lineages are 8.7% uncorrected 16S distance to L. guibei and 10.3% to L. miops. It is characterized by the presence of very distinct lateral spine-like scales at the base of the tail in males, as are found in several representatives of subclade A5 but not in other Domerguella. It can also be assigned to the subgenus Domerguella by an undivided mental scale with two postmentals, absence of a claw on the first finger, and 7 preanal pores in males. Within Domerguella, the new species is one of only two species of subclade A5 known from northern Madagascar. It differs from the other nominal species of Domerguella occurring in the same general area as follows: from L. expectatus by non-enlarged dorsolateral scales (longitudinal count of dorsal scales>185 vs. L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape with less elongated limbs (relative hindlimb length 0.49–0.54 vs.>0.55); from L. madagascariensis, L. petteri, and L. salvi by a lower longitudinal count of ventral scales (83–98 vs.>100); from L. roellae and L. hapei, by a higher longitudinal count of dorsal scales (187–222 vs. 159–179); and from L. tantsaha by a lower longitudinal count of dorsal scales (187–222 vs. 239–240). Furthermore, the new species differs from all of these species of the subclades A1–A4 by the presence of a distinct lateral spine at the base of the tail, especially large in males but also clearly recognizable in females (vs. more weakly expressed or absent in the other species). From the other two nominal species in subclade A5 (L. guibei and L. miops, according to current taxonomy; see above) the new species differs as follows: from L. miops by a lower longitudinal count of ventral scales despite minimal overlap (83–98 vs. 98–113); and from L. guibei by apparently relatively longer hindlimbs (HIL/SVL 0.49–0.54 vs. 0.42–0.49). Further comparative examination of specimens also revealed a different head shape in L. winki compared to L. guibei, with apparently more expressed supraocular bulges (also visible in specimens in life; Fig. 14 vs. Fig. 20). Additional measurements taken on selected specimens in good, fully comparable state of preservation (Table 3) revealed that L. winki individuals have proportionally longer and higher heads than L. guibei, with non-overlapping values (relative snout tip to tympanum distance in percent, 24.7–28.8 vs. 23.0–24.6; relative head height in percent, 12.5–14.2 vs. 11.1–12.4; see Table 3). L. winki sp. nov. does not share haplotypes in CMOS or RAG1 with L. miops, and only one instance of haplotype sharing in RAG1 is detected with L. guibei. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. This species is dedicated to Michael Wink, pharmacologist, herpetologist, ornithologist and professor emeritus of the University of Heidelberg, in recognition for his support of research in squamate systematics. The name is a patronym (i.e., a noun in the genitive case). Description of the holotype. Adult male, hemipenes everted, in good state of preservation, tail is broken and missing, second toe on the left forelimb is removed as source of tissue for molecular analysis (Fig. 15). SVL 29.5 mm, TAL 15.6 mm; for other measurements see Table 1. Head and neck short, head broader than body. The distance from the tip of the snout to the anterior border of the eye (3.9 mm) is less than the interorbital distance anteriorly (4.0 mm), and greater than the distance between the eye and ear opening. Snout covered with granular scales larger than those on the rest of the dorsum. Nostril surrounded by five scales: rostral, first supralabial, and three supranasals. Mental scale undivided; no contact between posterior projection of mental scale and first infralabial; two symmetrical postmental scales with four postpostmental scales; six infralabial scales; seven supralabial scales; two internasal scales; granular dorsal scales; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, smaller than on head and tail, the scales on limbs can be slightly larger; 222 dorsal scales longitudinally along the body; 93 ventral scales between mental and cloaca; venter with large homogeneous smooth scales; first finger present but very small, not bearing a claw; three pairs of subdigital lamellae on the fourth toe; eight not very distinct dorsolateral tubercles, consisting of one scale; seven preanal pores; tail without whorls; large lateral spines at the base of the tail. Based on available photographs (Fig. 20), the holotype displayed a light marbled yellow grayish dorsal coloration with more brownish flanks before preservation. Distinct yellow spots are present on the head, the flank, the limbs, and the tail. Two black markings are present on the shoulder on either side of the body, reminiscent of double scapular semi-ocelli. Along the spine a narrow brown line reaches from the neck to the base of the tail. Adjacent to this line two pairs of symmetrical dark spots are present (forming two disrupted chevrons), one pair on forelimb level and one pair 10 mm posterior (Fig. 20D). After six years of preservation in ethanol, the preserved specimen is more uniformly brownish and most of the marbled pattern is faded. The ventral side is fawn with few small brown spots, most of them in the gular region. Variation. Males display a light marbled yellow grayish dorsal coloration with more brownish flanks and yellow spots. The dorsal and ventral coloration on females is darker without a pattern except for a few dark dorsal spots (Fig. 20E and 20I in comparison to Fig. 20A–G). Three additional specimens (two males [ZSM 555 /2014, ZSM 48 /2016], one female [ZSM 1763 /2010]) were examined. The SVL ranges between 29.8 and 33.4 mm with the female being the largest. The two males have a TAL of 37.8 and 45.0 mm. The relative hindlimb length is 0.49 to 0.53 with the female having the smallest. The female also has the smallest eyes relative to the size of the body and with a medium size smaller tubercles at the tail base than the males, which have large tubercles. The number of dorsal scales ranges between 187 and 208. The ventral scales range between 83 and 98. Natural history. The holotype was collected in primary rainforest, on the trunk of a big tree, 0.2 m above the ground. The paratypes from Ampotsidy were mostly collected at night sleeping on leaves, twigs, or vines. UADBA-R 70856 was collected in the afternoon, on the ground during heavy rain. Distribution. L. winki is known from three localities in the North and Sambirano regions in northern Madagascar: (1) the type locality, Ampotsidy, (2) Andrevorevo, and (3) Bemanevika.