Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar’s rainforest Author Vences, Miguel m.vences@tu-braunschweig.de Author Multzsch, Malte m.multzsch@tu-braunschweig.de Author Gippner, Sven Author Miralles, Aurélien Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire Naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, Paris, France Author Crottini, Angelica 0000-0002-8505-3050 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485 - 661 Vairão, Portugal & Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, 4099 - 002 Porto, Portugal & BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485 - 661 Vairão, Portugal acrottini @ cibio. up. pt; https: // orcid. org / 0000 - 0002 - 8505 - 3050 acrottini@cibio.up.pt Author Gehring, Philip-Sebastian 0000-0002-8266-2184 Faculty of Biology / Biologiedidaktik, University Bielefeld, Universitätsstrasse 25, 33615 Bielefeld, Germany sebastiangehring @ web. de; https: // orcid. org / 0000 - 0002 - 8266 - 2184 sebastiangehring@web.de Author Rakotoarison, Andolalao 0000-0003-2620-440X Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar andomailaka@gmail.com Author Ratsoavina, Fanomezana M. Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar Author Glaw, Frank glaw@snsb.de Author Scherz, Mark D. mark.scherz@gmail.com text Zootaxa 2022 2022-09-01 5179 1 1 61 http://dx.doi.org/10.11646/zootaxa.5179.1.1 journal article 135489 10.11646/zootaxa.5179.1.1 fab2423d-f82d-4543-8720-86cd568c18b6 1175-5326 7040745 70366A84-EBDE-427D-B525-09E5A2D81EB5 Lygodactylus roellae sp. nov. Lygodactylus sp. 17: Gippner et al. (2021) . Holotype . ZSM 49 /2016 ( MSZC 0072 ), adult female, collected by M.D. Scherz , J. Borrell, L . Ball, T . Starnes, E . Razafimandimby, D.H . Nomenjanahary, and J. Rabearivony at Ampotsidy mountains , 15.7 km NNW of Bealanana ( 8.7 km NNW of Beandrarezona ), northern Madagascar , at geographical coordinates S14.41974 , E48.71935 , 1344 m a.s.l. , on 22 December 2015 ( Fig. 15 ). Paratypes . ZSM 556 /2014 ( DRV 6289 ), adult male, collected by F.M. Ratsoavina , D. Vieites , M. Vences , R . D. Randrianiaina , S. Rasamison , A. Rakotoarison , E. Rajeriarison, and T . Rajoafiarison at Andrevorevo , a site south of the Tsaratanana Massif , northern Madagascar , at geographical coordinates S14.3464 , E49.1028 , 1717 m a.s.l. , on 21 June 2010 ; UADBA-R 70855 ( MSZC 0010 ), adult female, with the same collection data as the holotype but collected at S14.41878 E48.71896 , 1354 m a.s.l. on 18 December 2015 at 20h20 . Diagnosis. Lygodactylus roellae sp. nov. corresponds to a genetically highly distinct lineage from northern Madagascar that is the sister species of L. salvi described above, but differs by high genetic divergence and several scale counts. It belongs to subclade A3 within Domerguella as defined herein. 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 5 preanal pores in males. Within Domerguella , the new species is only known from two localities in northern Madagascar and differs from the other nominal species of Domerguella occurring in this part of the island as follows: from L. expectatus by a different color pattern, without scapular semi-ocellus and with a striped pattern apparently in most individuals (vs. scapular semi-ocellus usually present, and striped pattern unknown); from L. rarus by lack of regular crossbands on tail (vs. presence) and different body shape without elongated limbs (relative hindlimb length 0.41–0.45 vs.>0.55); from L. madagascariensis , L. petteri , L. salvi , and L. tantsaha by smaller longitudinal dorsal scale count (159–169 vs.>188) and smaller longitudinal ventral scale count (83–92 vs.>96). Genetically, the new species is highly distinct from all species in subclade A5, and differs at least from L. guibei by a less distinctly expressed lateral spine at the tail base of males (vs. presence of a distinct, large spine). Furthermore, the longitudinal dorsal scale count is smaller than in all known individuals of subclade A5. The new species differs from its sister lineage, L. salvi (described above), by a lower longitudinal dorsal scale count (159–169 vs. 211–217) and a lower ventral scale count (83–92 vs. 107–112). The two sister species also differ by a high genetic divergence of 11.2–12.6% in the 16S gene, and do not share haplotypes in RAG1 despite occurring in geographical proximity. For a distinction from additional species newly named and described herein, see the respective diagnoses below. Etymology. We are pleased to dedicate this beautiful gecko species to Beate Röll, in recognition for her substantial contributions to Lygodactylus biology and phylogeny. The name is a matronym (i.e., a noun in the genitive case). Description of the holotype . Adult female, in a good state of preservation, the right hind limb is partly removed as a source of tissue for molecular analysis ( Fig. 15 ). SVL 35.9 mm , TAL 39.6 mm ; for other measurements see Table 1 . Head and neck thick, body broader than head. The distance from the tip of the snout to the anterior border of the eye ( 3.8 mm ) is less than the interorbital distance anteriorly ( 4.2 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 asymmetrical postmental scale with five postpostmental scales; seven 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; 169 dorsal scales longitudinal along the body; 92 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; no dorsolateral tubercles; tail without whorls; small lateral spines at the base of the tail. Based on available photographs ( Fig. 18A–B ), the life coloration of the holotype exhibited a distinct pattern of dark brownish to yellowish stripes on the dorsum reaching from the eye to base of the tail. The stripes continue in an irregular pattern of more elongated dark and brighter spots on the caudal spine. The dark wide area on the back is slightly emarginated at the level of the hindlimbs before ending at the base of the tail. The flanks and limbs are gray to brownish ( Fig. 18A ). The ventral side is whitish with few small brown spots predominately on the throat and the tail ( Fig. 18B ). The specimen has darkened after 6 years of preservation in ethanol; however, the striped pattern is still distinctly visible. Variation. The coloration of the dorsum is characteristic with a distinct pattern of dark brownish to yellowish stripes on the dorsum with a variable strength of expression on the tail ( Fig. 18A and 18E ), which appears to weaken greatly or be lost upon regeneration ( Fig. 18C ). FIGURE 18. Photos of Lygodactylus roellae sp. nov. in life. A and B, adult female holotype ZSM 49/2016 (MSZC 0072) from Ampotsidy. C,D, adult female paratype UADBA-R 70855 from Ampotsidy. E, male paratype ZSM 556/2014 (DRV 6289) from Andrevorevo. TABLE 1. Selected morphometric and meristic data for voucher specimens of the Lygodactylus madagascariensis group examined for this study. All morphometric measurements (SVL, TAL, HIL, HW, ED, SED) in mm. All specimens in the table were genotyped by 16S sequences, except for historical types from BMNH, MNHN, and SMF, and two further individuals of L. expectatus . Abbreviations are those described in Materials and Methods section; additional abbreviations and symbols as follows: NA, not assessed or not applicable; * counts and measurements partly or fully taken by M. Puente ( Puente et al. 2009 ); § specimen not genotyped; # tail broken or incomplete; ## tail regenerated. HT (sep) and HT (spi) mark the holotypes of Lygodactylus septemtuberculatus Angel, 1942 and Microscalabotes spinulifer Boettger, 1913 , here considered to be junior synonyms of L. miops . First LCDS value for L. expectatus refers to the enlarged dorsolateral scales, the second value to the non-enlarged vertebral scales. See species account of L. fritzi sp. nov for the identity of the historical locality “Senbendrana”.

Specimen voucher	Field number	Locality	Type	Sex	SVL	TAL	HIL	HW	ED	SED	size STT
L. expectatus —subclade A1
MNHN 1990.1 *§	NA	Ankarana	HT	M	29.7	32.6	13.1	NA	NA	NA	0
MNHN 1990.2 * §	NA	Ankarana	PT	F	27.8	25.3	NA	NA	NA	NA	0
MNHN 1990.3 * §	NA	Ankarana	PT	NA	19.7	20.8	NA	NA	NA	NA	NA
MNHN 1933.168 * §	NA	Ankarana	F	27.5	0.0	NA	NA	NA	NA	NA
ZSM 282/2004	FGZC 541	Ankarana	M	27.7	16.8#	12.8	5.3	1.7	6.7	0
ZSM 492/2001 §	MV 2001.256	Ankarana	M	26.3	30.7	11.8	5.2	1.6	7.2	0
ZSM 284/2004	FGZC 543	Ankarana	M	29.4	38.1#	12.8	5.5	1.6	7.3	0
ZSM 1540/2008	FGZC 1649	Ankarana	M	29.2	31.5	13.9	5.8	1.5	8.0	0
ZSM 2143/2007	FGZC 1183	Ankarana	F	24.7	28.6	11.6	4.6	1.4	6.6	0
ZSM 283/2004	FGZC 542	Ankarana	F	24.3	31.8	11.0	5.1	1.4	6.7	0
UADBA 06059 * §	NA	Ankarana	NA	21.9	0.0	NA	NA	NA	NA	NA
L. rarus —subclade A1
MNHN 1990.6 * §	NA	Ankarana	HT	F	22.6	NA	12.9	NA	NA	NA	0
MNHN 1990.1887 * §	NA	Ankarana	NA	13.2	13.0	NA	NA	NA	NA	NA
MNHN 1990.1888 * §	NA	Ankarana	NA	22.5	13.4	NA	NA	NA	NA	NA
ZSM 860/2003	FGMV 2002.833	Ankarana	M	33.5	37.0	20.0	5.9	1.3	8.3	0
ZSM 913/2003	FGMV 2002.941	Ankarana	M	36.5	44.0	20.6	6.3	1.6	8.6	0
ZSM 1085/2003	FGMV 2002.837	Ankarana	F	35.5	23.0#	20.7	6.1	1.4	9.5	0
L. tantsaha sp. nov. (sp. 10)—subclade A2
ZSM 196/2018	MSZC 772	Montagne d’Ambre	HT	M	31.9	36.9#	15.8	6.3	1.6	8.5	0
ZSM 197/2018	MSZC 771	Montagne d’Ambre	PT	M	36.9	12.2#	16.8	6.4	1.8	9.1	1
L. hapei sp. nov. (sp. 26)—subclade A2
ZSM 298/2018	NA	Djohahely (Djangoa)	HT	F	26.3	27.4§	11.4	4.3	1.6	6.3	0

......continued on the next page TABLE 1. (Continued)

Specimen voucher	Field number	Locality	Type	Sex	SVL	TAL	HIL	HW	ED	SED	size STT
L. salvi sp. nov. (sp. 8)—subclade A3
ZSM 557/2014	DRV 6327	Ambodikakazo	PT	F	36.2	29.0#	17.2	6.8	1.8	8.7	1
ZSM 783/2001	FGMV 2001.74	Tsaratanana (Antsahamanara)	HT	M	29.9	24.9##	15.0	5.2	1.2	8.0	1–2
L. roellae sp. nov. (sp. 17)—subclade A3
ZSM 49/2016	MSZC 72	Ampotsidy	HT	F	35.9	39.6	14.7	6.4	1.8	8.7	1
ZSM 556/2014	DRV 6289	Andrevorevo	PT	M	36.0	45.3	16.1	7.0	1.7	9.0	1
L. madagascariensis —subclade A4
SMF 8937 §	NA	Nosy Be	HT	M	31.6	37.1	14.4	5.4	2.0	7.7	1
ZSM 782/2001	MV 2001.55	Tsaratanana (Andampy)	F	31.5	10.2#	13.1	5.6	1.5	8.1	1
ZSM 118/2019	FGZC 5511	Maromandria	M	34.0	38.7	14.3	5.9	1.9	8.3	1
ZSM 119/2019	FGZC 5515	Maromiandra	M	29.5	35.0	13.3	5.7	1.7	8.0	1
ZSM 120/2019	FGZC 5596	Nosy Be (Lokobe)	F	28.5	35.0	13.6	4.9	1.7	7.1	1
ZSM 813/2003	FGMV 2002.721	Manongarivo	M	31.7	36.2	12.6	5.2	1.7	7.6	1
ZSM 832/2003	FGMV 2002.778	Manongarivo	M	30.4	40.9#	14.5	5.8	1.7	7.4	1
ZSM 270/2004	FGZC 518	Montagne d’Ambre	F	30.2	29.0	14.1	5.3	1.7	7.4	0–1
L. hodikazo sp. nov. (sp. 23)—subclade A4
ZSM 77/2006	FGZC 828	Tsingy de Bemaraha	HT	F	29.4	37.2	12.9	5.6	1.6	7.2	1–2
L. petteri —subclade A4
MNHN 1990.4 * §	1346-L	Montagne d’Ambre	HT	F	35.0	12.0#	16.9	NA	NA	NA	1
MNHN 1990.5 §	1347-L	Montagne d’Ambre	PT	M	33.2	0.0	NA	NA	NA	NA	1
ZSM 914/2003	FGMV 2002.942	Montagne d’Ambre	F	36.7	42.1	16.0	6.2	1.9	8.8	1
ZSM 2087/2007	FGZC 1065	Montagne d’Ambre	M	37.8	48.0	19.0	7.1	2.0	9.4	1
ZSM 194/2018	MSZC 454	Montagne d’Ambre	F	30.3	15.5#	15.3	6.0	1.5	8.0	0
ZSM 195/2018	MSZC 485	Montagne d’Ambre	M	38.5	34.6##	18.4	6.7	2.0	9.4	0
L. guibei —subclade A5
MNHN 1933.156 * §	NA	Moramanga	PT	M	30.2	36.3	NA	NA	NA	7.7	2–3
MNHN 1993.60 * §	NA	Andasibe	HT	M	30.8	37.0	13.8	NA	NA	7.8	3
ZSM 5117/2005	FGZC 2689	Andasibe	F	34.0	41.7	15.3	6.5	1.9	8.0	2
ZSM 525/2009	ZCMV 11210	Makira (Angozongahy)	F	31.3	17.9#	15.3	6.0	1.7	7.7	1–2
ZSM 319/2004	FGZC 613	Moramanga region	F	35.8	31.3	15.6	6.7	2.3	10.0	1–2

......continued on the next page TABLE 1. (Continued)

Specimen voucher	Field number	Locality	Type	Sex	SVL	TAL	HIL	HW	ED	SED	size STT
ZSM 320/2004	FGZC 616	Moramanga region	M	34.9	36.8	15.4	6.7	2.2	8.7	3
ZSM 321/2004	FGZC 617	Moramanga region	M	33.2	39.0	14.5	6.0	2.0	8.0	2
ZSM 330/2004	FGZC 627	Moramanga region	F	39.5	29§	16.5	6.7	2.2	9.0	2–3
ZSM 349/2006	ZCMV 2484	-	F	30.1	10.2#	13.4	5.6	2.0	7.7	2–3
ZSM 349/2010	FGZC 4362	Anjozorobe	M	37.6	46.5#	15.3	7.0	2.0	9.1	2
L. miops —subclade A5
BMNH 1946.8.22.55 §	NA	“Senbendrana”	HT	F	29.9	21.4#	12.8	5.3	2.1	7.4	2
MNHN 1893.63 * §	NA	Moramanga	HT (sep)	F	29.0	19.0##	13.1	NA	NA	NA	2
SMF 8931 * §	NA	Moramanga	HT (spi)	F	28.5	27.6#	14.2	5.1	2.2	7.0	1–2
ZSM 351/2010	FGZC 4295	Ambodivoahangy	F	29.2	28.8	13.7	5.6	1.8	7.4	1–2
ZSM 731/2003	FGMV 2002.456	Ranomafana	M	27.2	3.6#	12.8	5.0	1.6	7.0	1–2
ZSM 732/2003	FGMV 2002.458	Ranomafana	F	30.4	30.8	13.2	5.2	1.7	6.7	1–2
ZSM 733/2003	FGMV 2002.459	Ranomafana	F	29.8	31.7	13.8	5.2	1.7	7.5	2
ZSM 352/2006	ZCMV 3350	Mahakajy Reserve	F	30.8	10.4#	15.1	6.0	1.9	7.6	2
ZSM 116/2004	FGZC 205	Andohahela	M	26.2	5.5#	11.7	5.7	1.5	7.0	2
ZSM 117/2004	FGZC 206	Andohahela	F	31.2	16.5#	15.1	6.0	1.8	7.6	2
ZSM 165/2004	FGZC 307	Manantantely	M	30.0	29.8	13.0	5.6	2.3	7.6	2
ZSM 352/2010	FGZC 4503	Anosibe An’Ala	F	29.4	30.5	13.4	5.6	1.6	7.0	2
ZSM 175/2016	FGZC 5039	Vohimana	F	30.1	33.2#	13.1	5.4	1.8	7.4	1–2
ZSM 376/2016	ZCMV 14832	Sampanandrano	F	28.9	16.3#	12.1	5.3	1.6	7.0	1
L. fritzi sp. nov. (sp.. 11)—subclade A5
ZSM 651/2009	ZCMV 8902	Ankanin’ny Nofy	F	26.4	26.5	10.7	4.9	1.8	6.8	0–1
L. winki sp. nov. (sp. 18)—subclade A5
ZSM 555/2014	DRV 6288	Andrevorevo	PT	M	33.0	45.0	17.4	7.1	2.0	8.6	3
ZSM 48/2016	MSZC 110	Ampotsidy	PT	M	29.8	37.8	14.9	6.0	1.9	7.7	3
ZSM 1763/2010	ZCMV 12502	Bemanevika	PT	F	33.4	18.7#	16.2	6.5	1.8	8.2	2
ZSM 47/2016	MSZC 0075	Ampotsidy	HT	M	29.5	15.6#	15.8	5.5	1.8	7.3	3
L. ulli sp. nov. (sp. 21)—subclade A5
ZSM 154/2005	FGZC 2811	Marojejy	HT	M	28.8	2.6#	13.6	5.3	1.7	6.7	2

......continued on the next page TABLE 1. (Continued)

Specimen voucher	Field number	Locality	Type	Sex	PPM	INFL	SUPL	IN	PCL	NDT	LCDS	LCVS
L. expectatus —subclade A1
MNHN 1990.1 *§	NA	Ankarana	HT	M	4	5	5	1	7	4	122/162	95
MNHN 1990.2 * §	NA	Ankarana	PT	F	4	6	6	1	0	0	NA	96
MNHN 1990.3 * §	NA	Ankarana	PT	NA	4	5	6	1	0	0	NA	94
MNHN 1933.168 * §	NA	Ankarana	F	4	5	6	2	0	0	NA	NA
ZSM 282/2004	FGZC 541	Ankarana	M	4	7	8	1	5	4	127/164	91
ZSM 492/2001 §	MV 2001.256	Ankarana	M	4	6	7	1	7	5	124/139	98
ZSM 284/2004	FGZC 543	Ankarana	M	4	6	7	1	5	3	128/157	92
ZSM 1540/2008	FGZC 1649	Ankarana	M	4	6	7	1	7	4	127/155	96
ZSM 2143/2007	FGZC 1183	Ankarana	F	4	5	6	1	0	4	130/154	86
ZSM 283/2004	FGZC 542	Ankarana	F	4	6	7	2	0	3	125/162	87
UADBA 06059 * §	NA	Ankarana	NA	4	6	6	1	0	0	NA	NA
L. rarus —subclade A1
MNHN 1990.6 * §	NA	Ankarana	HT	F	4	NA	NA	3	0	215	125
MNHN 1990.1887 * §	NA	Ankarana	NA	4	6	7	3	0	0	NA	NA
MNHN 1990.1888 * §	NA	Ankarana	NA	4	5	6	1	0	0	NA	NA
ZSM 860/2003	FGMV 2002.833	Ankarana	M	5	5	6	1	7	0	260	120
ZSM 913/2003	FGMV 2002.941	Ankarana	M	4	5	7	2	7	0	224	119
ZSM 1085/2003	FGMV 2002.837	Ankarana	F	5	7	8	2	0	0	223	139
L. tantsaha sp. nov. (sp. 10)—subclade A2
ZSM 196/2018	MSZC 772	Montagne d’Ambre	HT	M	4	7	7	3	7	1	239	111
ZSM 197/2018	MSZC 771	Montagne d’Ambre	PT	M	5	7	7	1	7	6	240	97
L. hapei sp. nov. (sp. 26)—subclade A2
ZSM 298/2018	NA	Djohahely (Djangoa)	HT	F	5	7	8	3	0	0	179	87
L. salvi sp. nov. (sp. 8)—subclade A3
ZSM 557/2014	DRV 6327	Ambodikakazo	PT	F	4	6	7	2	0	5	211	112
ZSM 783/2001	FGMV 2001.74	Tsaratanana (Antsahamanara)	HT	M	4	7	7	3	6	6	217	107

......continued on the next page TABLE 1. (Continued)

Specimen voucher	Field number	Locality	Type	Sex	PPM	INFL	SUPL	IN	PCL	NDT	LCDS	LCVS
L. roellae sp. nov. (sp. 17)—subclade A3
ZSM 49/2016	MSZC 72	Ampotsidy	HT	F	5	7	7	2	0	0	169	92
ZSM 556/2014	DRV 6289	Andrevorevo	PT	M	6	6	8	1	5	6	159	83
L. madagascariensis —subclade A4
SMF 8937 §	NA	Nosy Be	HT	M	4	6	8	2	7	0	246	106
ZSM 782/2001	MV 2001.55	Tsaratanana (Andampy)	F	5	7	8	3	0	4	205	106
ZSM 118/2019	FGZC 5511	Maromandria	M	4	8	9	1	7	5	236	121
ZSM 119/2019	FGZC 5515	Maromiandra	M	4	8	9	2	7	5	251	138
ZSM 120/2019	FGZC 5596	Nosy Be (Lokobe)	F	4	7	8	2	0	3	219	128
ZSM 813/2003	FGMV 2002.721	Manongarivo	M	5	7	9	2	7	4	258	126
ZSM 832/2003	FGMV 2002.778	Manongarivo	M	4	6	7	1	7	0	244	124
ZSM 270/2004	FGZC 518	Montagne d’Ambre	F	4	7	8	2	0	4	258	109
L. hodikazo sp. nov. (sp. 23)—subclade A4
ZSM 77/2006	FGZC 828	Tsingy de Bemaraha	HT	F	4	8	10	1	0	3	231	109
L. petteri —subclade A4
MNHN 1990.4 * §	1346-L	Montagne d’Ambre	HT	F	4	6	6	1	0	0	189	102
MNHN 1990.5 §	1347-L	Montagne d’Ambre	PT	M	4	7	6	1	7	5	NA	105
ZSM 914/2003	FGMV 2002.942	Montagne d’Ambre	F	4	7	7	2	0	0	222	113
ZSM 2087/2007	FGZC 1065	Montagne d’Ambre	M	4	7	7	1	7	3	209	101
ZSM 194/2018	MSZC 454	Montagne d’Ambre	F	4	7	8	1	0	0	214	101
ZSM 195/2018	MSZC 485	Montagne d’Ambre	M	4	7	8	1	7	0	214	103
L. guibei —subclade A5
MNHN 1933.156 * §	NA	Moramanga	PT	M	4	6	6	1	7	1	198+	99+
MNHN 1993.60 * §	NA	Andasibe	HT	M	4	NA	NA	2	6	9+	170	88
ZSM 5117/2005	FGZC 2689	Andasibe	F	5	6	7	2	0	5	206	94
ZSM 525/2009	ZCMV 11210	Makira (Angozongahy)	F	4	7	7	1	0	3	220	95
ZSM 319/2004	FGZC 613	Moramanga region	F	4	7	8	1	0	3	195	87
ZSM 320/2004	FGZC 616	Moramanga region	M	6	7	9	1	6	4	180	91

......continued on the next page TABLE 1. (Continued)

Specimen voucher	Field number	Locality	Type	Sex	PPM	INFL	SUPL	IN	PCL	NDT	LCDS	LCVS
ZSM 321/2004	FGZC 617	Moramanga region	M	4	6	6	2	7	4	201	87
ZSM 330/2004	FGZC 627	Moramanga region	F	4	7	7	1	0	0	181	89
ZSM 349/2006	ZCMV 2484	-	F	4	6	7	1	0	0	191	109
ZSM 349/2010	FGZC 4362	Anjozorobe	M	5	6	6	2	5	6	191	89
L. miops —subclade A5
BMNH 1946.8.22.55 §	NA	“Senbendrana”	HT	F	6	7	7	3	0	3	233	113
MNHN 1893.63 * §	NA	Moramanga	HT (sep)	F	5	5	7	1	0	3	225	102
SMF 8931 * §	NA	Moramanga	HT (spi)	F	5	7	8	3	0	5	240	107
ZSM 351/2010	FGZC 4295	Ambodivoahangy	F	5	6	8	2	0	4	205	102
ZSM 731/2003	FGMV 2002.456	Ranomafana	M	4	7	7	3	7	4	236	106
ZSM 732/2003	FGMV 2002.458	Ranomafana	F	4	8	8	2	0	3	237	108
ZSM 733/2003	FGMV 2002.459	Ranomafana	F	6	7	8	2	0	3	237	113
ZSM 352/2006	ZCMV 3350	Mahakajy Reserve	F	3	6	8	2	0	4‒5	220	98
ZSM 116/2004	FGZC 205	Andohahela	M	4	7	8	2	7	4	207	109
ZSM 117/2004	FGZC 206	Andohahela	F	5	8	8	2	0	4	229	108
ZSM 165/2004	FGZC 307	Manantantely	M	5	6	7	2	8	6	233	106
ZSM 352/2010	FGZC 4503	Anosibe An’Ala	F	6	6	7	2	0	4	216	105
ZSM 175/2016	FGZC 5039	Vohimana	F	5	6	7	2	0	5	242	106
ZSM 376/2016	ZCMV 14832	Sampanandrano	F	5	6	6	2	0	4	210	102
L. fritzi sp. nov. (sp.. 11)—subclade A5
ZSM 651/2009	ZCMV 8902	Ankanin’ny Nofy	F	4	7	8	2	0	2	247	98
L. winki sp. nov. (sp. 18)—subclade A5
ZSM 555/2014	DRV 6288	Andrevorevo	PT	M	4	7	7	1	7	6‒8	187	93
ZSM 48/2016	MSZC 110	Ampotsidy	PT	M	4	7	8	2	7	5	208	83
ZSM 1763/2010	ZCMV 12502	Bemanevika	PT	F	4	7	8	1	0	6	204	98
ZSM 47/2016	MSZC 0075	Ampotsidy	HT	M	4	6	7	2	7	8	222	93
L. ulli sp. nov. (sp. 21)—subclade A5
ZSM 154/2005	FGZC 2811	Marojejy	HT	M	5	6	7	2	7	2	253	110

The examined male paratype specimen (ZSM 556/2014) is almost the same size as the holotype with a SVL of 36.0 mm and has a longer tail ( 45.3 mm ) and hindlimbs (HIL/SVL 0.45). On the fourth toe it has four instead of three subdigital lamellae, which differs from all other examined specimens. Unlike the holotype it has tubercles between the limbs (6) that consist of one scale each. It has fewer dorsal (159) and ventral scales (83). These differences could be due to different sex or just random variation. Natural history. Specimens of this species were collected sleeping at night on roosts up to 1 m above the ground in Ampotsidy. UADBA-R 70855 was found sleeping on the tip of a Pandanus frond. It occurs in close sympatry with Lygodactylus winki sp. nov. , described below. Distribution. L. roellae is known from (1) the type locality Ampotsidy and (2) Andrevorevo.