Lygodactylus morii, Vences & Multzsch & Gippner & Crottini & Glaw & Köhler & Rakotomanga & Rasamison & Raselimanana, 2024

Vences, Miguel, Multzsch, Malte, Gippner, Sven, Crottini, Angelica, Glaw, Frank, Köhler, Jörn, Rakotomanga, Sandratra, Rasamison, Solohery & Raselimanana, Achille P., 2024, Taxonomizing a truly morphologically cryptic complex of dwarf geckos from Madagascar: molecular evidence for new species-level lineages within the Lygodactylus tolampyae complex, Zootaxa 5468 (3), pp. 416-448 : 432-435

Lygodactylus morii sp. nov.

( Fig. 5–7 , 10–12 )

Remark. This species corresponds to populations previously considered as L. tolampyae sensu stricto by Gippner et al. (2021).

Holotype. ZSM 501/2001 (field number FGMV = MV 2001.300), adult male ( Fig. 5 , 10 ), collected by M. Vences, D.R. Vieites, G. Garcia, V. Raherisoa, and A. Rasoamamonjinirina, near Ampijoroa, Ankarafantsika National Park , Madagascar (approximate geographical coordinates 16.3°S, 46.82°E), on 22 February 2001.

Paratypes. A total of nine specimens (only genotyped individuals included in paratype series). ZSM 151/2013 (ZCMV 14173), one female, collected by F. Ratsoavina, J. Erens, and E. Rajeriarison, between Soalala and Mahajanga (at geographical coordinates 16.05298°S, 45.81034°E, 8 m a.s.l.), on 3 January 2013; ZSM 152/2013 (ZCMV 14172), one male, collected by F. Ratsoavina, J. Erens, and E. Rajeriarison at Katsepy (Ankiririka; 15.71289°S, 46.21650°E, 18 m a.s.l.), on 4 January 2013; ZSM 224/2018 (FGZC 5701), ZSM 225/2018 (FGZC 5705) and ZSM 226/2018 (FGZC 5706), three females, collected by F. Glaw, D. Prötzel, N.A. Raharinoro, R.N. Ravelojaona, A. Razafimanantsoa, J. Forster, K. Glaw, T. Glaw, and C. Zanotelli at Antsanitia (15.567710°S, 46.423104°E, ca. 10 m a.s.l.), on 28 March 2018; ZSM 227/2018 (FGZC 5727), female, collected by F. Glaw, D. Prötzel, N.A. Raharinoro, R.N. Ravelojaona, A. Razafimanantsoa, J. Forster, K. Glaw, T. Glaw, and C. Zanotelli at Antrema (15.716604°S, 46.216802°E, 90 m a.s.l.), on 31 March 2018; ZSM 159/2022 (ZCMV 15703), UADBA-ZCMV 15704, UADBA-ZCMV 15705, one male and two females, collected by M. Vences, S. Rakotomanga, S. Rasamison, and P. Galán near Ampijoroa, Ankarafantsika National Park (approximate geographical coordinates 16.3°S, 46.82°E), on 12 November 2022.

Referred additional material. ZSM 150/2013 (ZCMV 14165), male, collected by F. Ratsoavina, J. Erens, and E. Rajeriarison between Soalala and Mahajanga (at geographical coordinates 16.05298°S, 45.81034°E, 8 m a.s.l.), on 3 January 2013 (not included in paratype series as it belongs to genetic lineage Ic). ZSM 491/2001, ZSM 502/2001, ZSM 503/2001, one male and two females, collected in February 2001 by M. Vences, D.R. Vieites, G. Garcia, V. Raherisoa, and A. Rasoamamonjinirina, near Ampijoroa, Ankarafantsika National Park (not included in paratype series as species assignment not confirmed by molecular data) .

Diagnosis. L. morii sp. nov. is characterized as member of the L. tolampyae complex (and thereby distinguishable from all other Malagasy Lygodactylus not belonging to the complex) by combination of a mental scale semi-divided by a suture, broad contact of the posterior projection of the mental scale with the first infralabial scale, and three postmental scales; furthermore, it is characterized by the absence of whorls on the tail, and a typical look of the head with relatively large eyes. It is distinguished from L. tolampyae sensu stricto (as defined herein; mitochondrial lineages A+B and possibly G) by significantly higher mean counts of dorsal scales despite widely overlapping ranges (mean 246 vs. 229; ranges 225–269 vs. 204–241; Mann-Whitney U-test P<0.001), and possibly by a tendency towards broader posterior contact between the mental scale and the first infralabial scale, and by fewer (1 vs. 2–3) internasal scales in the majority of specimens. From a molecular perspective, the new species is characterized by numerous diagnostic nucleotide positions in the mitochondrial 16S rRNA gene: MolD identified a robust diagnostic nucleotide combination of an ‘A’ in the site 1072, ‘C’ in the site 1091, and ‘G’ in the site 1108 (positions relative to the full 16S rRNA gene of Phelsuma guimbeaui ).

Description of the holotype. Adult male, hemipenes everted, in moderate state of preservation, left hind limb removed as tissue source for molecular analysis, tail missing. SVL 29.4 mm, TAL 4.3 mm (as the tail is mutilated); for other measurements, see Table 2 . Body broader than head, possibly due to a flattened preservation state. The distance from the tip of the snout to the anterior border of the eye (3.7 mm) is less than the anterior interorbital distance (4.2 mm), and greater than the distance between the eye and ear opening (2.5 mm). Snout covered with granular scales larger than those on the dorsum. Nostril surrounded by five scales: rostral, first supralabial, one postnasal and two supranasals. The mental scale is semi-divided; contact between posterior projection of mental scale and first infralabial scale is roughly 40% of the infralabial scale length; three symmetrical postmental scales, followed by seven post-postmentals; six infralabial scales; eight supralabial scales; two internasal scales; dorsal scales granular; dorsum with small, homogeneous, granular, and unkeeled scales of similar size to those on trunk, no distinct size difference of scales on limbs; 245 dorsal scales longitudinally along the body; 102 ventral scales between mental and cloaca; venter with larger homogeneous smooth scales; first finger present, small, but bearing a claw; three pairs of subdigital lamellae on the fourth toe; no dorsolateral tubercles; seven preanal pores; no observable lateral spines at the base of the tail.

Based on the available photograph ( Fig. 5 ), in life, the holotype displayed a brownish colouration with darker regions on the dorsum and limbs. Above the spinal cord was a repeated darker pattern that continued on the tail. After each instance of the pattern, there was a spot of lighter colouration next to the middle stripe. The flanks were also darker in colour. The legs appeared to be lightly striped with a dark stripe followed by a lighter brownish region. The darker stripe at the flank continued on the head and transforms into a thin stripe, crossing the eye and ending shortly before the nostrils. The greyish colour visible in the picture was probably the result of a skin shedding, which just happened or was about to happen soon. The ventral side appeared to be whitish. In the throat region there were three visible tubercles of one scale each. After 22 years in ethanol ( Fig. 10 ), the specimen is overall darkened in colouration with the darker region still discernible from the lighter regions. The particular dorsal pattern, however, is not visible, except for the base of the tail and the limbs. Between the eyes is a whitish stripe, bordered by thin, darker stripes. The ventral side is whitish in colouration, with dark brown and light brown spots on the underside throat. The bluish spot between the limbs is probably coming from the organs, visible through the skin.

Variation. Similar to other Malagasy Lygodactylus , L. morii sp. nov. also has a striped colour morph that, according to our observations in the field, is rare; among at least 20 individuals seen in the wild in the Ankarafantsika/ Ampijoroa area, only one striped individual was found (male paratype ZSM 159/2022; see Fig. 6B ); it is characterized by a light brown to beige dorsolateral stripe that starts behind the eye and becomes gradually broader towards midbody, then narrows again; the stripes on either side of the body fuse on the tail and make up an overall light brown colour of the tail. The ventral side is usually uniformly white, especially on the throat, which can have a few scattered black spots in some individuals; this sparse spotting may extend onto the chest. The underside of the tail and especially the area of the precloacal pores can have a yellowish colour. The tail base is swollen in males due to the presence of hemipenes, but no lateral tail base tubercles are visible (as they would be characteristic for some species of the subgenus Domerguella ). Males appear to be smaller than females (SVL 28.2–29.4 vs. 28.7–32.8; see Table 2 ). For variations of other morphometric and scalation characters, see Table 2 .

Etymology. We dedicate this species to Akira Mori, Kyoto University, in recognition of his contributions to reptile biology and especially his leading role in the study of reptiles in Ankarafantsika National Park. The species epithet is a noun in the genitive case.

Natural history. This is a very common species in Ankarafantsika National Park ( Fig. 13 ) and apparently also occurs in other dry deciduous forests in the region, close to the North West coast. At Ankarafantsika, individuals were observed active during the day on relatively thin tree trunks at low perch heights of 1‒2 m, but were mostly observed at night, sleeping on thin terminal twigs ( Fig. 7 ), in agreement with observations in Mori et al. (2006). These authors recorded activity in the dry season but stated that individuals were more easily found during the rainy season.

Distribution. According to current data, reliably genotyped specimens of mitochondrial lineage Ia are known from (1) the type locality, Ankarafantsika National Park (including Ampijoroa, Ampondrabe forest, and unspecified sites outside of the protected area), (2) Antsakabe forest (Mariarano), (3) Antsanitia, (4) probably Betsako, (5) Katsepy (Ankiririka), (6) an unnamed place between Soalala and Mahajanga, and (7) Antrema. Furthermore, the only sample of lineage Ib was collected at (8) Bongolava Plateau, and a sample of Ic was collected from Besalampy (Andranomanintsy Forest).

Figures

FIGURE 5. Male holotype of Lygodactylus morii sp. nov., ZSM 501/2001 (FGMV 2001.300) from Ampijoroa,Ankarafantsika National Park, in life, photographed in February 2001.

FIGURE 6. Individualsof Lygodactylus morii sp. nov. from nearAmpijoroa (AnkarafantsikaNational Park)in life, photographed in November 2022.A. specimen with sample number MVTIS 33402/33488 (not collected). B. paratype ZSM 159/2022 (ZCMV 15703).

FIGURE 7. Four individuals of Lygodactylus morii sp. nov. photographed in November 2022 at different sites near Ampijoroa (Ankarafantsika National Park) in life, resting at night on terminal twigs of trees as is typical for the L. tolampyae complex. Individuals not individually assignable to sample or voucher specimen numbers, but corresponding to sequences included in Fig. 1.

FIGURE 10. Preserved holotypes of the three new species of the Lygodactylus tolampyae complex described herein, photographed in May 2023.

FIGURE 11. Ventral views of the heads of the holotypes of the three new species of Lygodactylus described herein, and one representative specimen of L. tolampyae. The images on the right show the mental region magnified and strongly sharpened to improve visibility of sutures between scales. For the holotype of L. morii sp. nov., the typical arrangement of scales in the mental region in this species complex is labelled: M, mental scales (semi-divided by incomplete sutures), PM, post-mentals (typically three), IL1, first infralabial (which maintains a broad contact with the mental). Scale bars (referring to the images on the left) represent 1 mm.

FIGURE 12. Lateral views of the heads of the holotypes of the three new species of Lygodactylus described herein, and one representative specimen of L. tolampyae. The scale bar represents 1 mm.

FIGURE 13. Habitat of species of the Lygodactylus tolampyae complex. A. Dry deciduous forest at Ampijoroa (Ankarafantsika) where L. morii sp. nov. occurs in high densities. B. View of remains of dry deciduous Ampondrabe forest (habitat of L. morii sp. nov. and L. herilalai sp. nov.) with large areas of forest destroyed by fire in the foreground. C, D. Transitional forest on Sahamalaza Peninsula, habitat of L. schwitzeri sp. nov. E. Forest in the Tsingy de Bemaraha, habitat of L. tolampyae. All photos by the authors except A by P. Galán.