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

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
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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
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
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