An inordinate fondness for inconspicuous brown frogs: integration of phylogenomics, archival DNA analysis, morphology, and bioacoustics yields 24 new taxa in the subgenus Brygoomantis (genus Mantidactylus) from Madagascar Author Scherz, Mark D. Author Crottini, Angelica CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485 - 661 Vairão, Portugal Author Hutter, Carl R. 0000-0001-6381-6339 Museum of Natural Sciences and Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, USA Ễcarl. hutter @ gmail. com; https: // orcid. org / 0000 - 0001 - 6381 - 6339 USAỄcarl.hutter@gmail.com Author Hildenbrand, Andrea Hauptstr. 13, 82234 Wessling, Germany Ễandrea @ gutachten-hildenbrand. de GermanyỄandrea@gutachten-hildenbrand.de Author Andreone, Franco Museo Regionale di Scienze Naturali, Via G. Giolitti, 36, 10123 Torino, Italy Author Fulgence, Thio Rosin 0000-0001-7205-7282 Natural and Environmental Sciences, Regional University Centre of the SAVA Region (CURSA), Antalaha, Madagascar Ễthiorosinf @ yahoo. fr; https: // orcid. org / 0000 - 0001 - 7205 - 7282 MadagascarỄthiorosinf@yahoo.fr Author Köhler, Gunther Author Ndriantsoa, Serge Herilala Author Ohler, Annemarie Author Preick, Michaela Author Rakotoarison, Andolalao Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar Author Rancilhac, Loïs Author Raselimanana, Achille P. Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar Author Riemann, Jana C. Author Rödel, Mark-Oliver Author Rosa, Gonçalo M. Author Streicher, Jeffrey W. Author Vieites, David R. Author Köhler, Jörn Author Hofreiter, Michael Author Glaw, Frank Author Vences, Miguel text Megataxa 2022 2022-12-15 7 2 113 311 http://dx.doi.org/10.5281/zenodo.6687412 journal article 217510 10.11646/megataxa.7.2.1 8af27bde-abaa-4f0a-8f84-8b6cc8a97034 2703-3090 7441023 2FD8C310-6486-4592-92F6-5EB894EBD6AC Mantidactylus alutus ( Peracca, 1893 ) Type material .— Rana aluta Peracca, 1893 is based on 25 syntypes according to the original description, 14 of the colour morph ‘forma’A and 11 of the ‘forma’ B.According to Frost (2021) , Rana aluta Peracca, 1893 includes the following syntypes : MZUT An725 and An729 , MNHN 1894.1–2 , and specimens in BMNH , all from ‘dintorni di Andrangoloaka e dalla vicina valle dell’Umbi’ . However, Gavetti and Andreone (1993) designated MZUT An725.1 as lectotype and redescribed this specimen including morphological measurements (summarised in Table 4 ). They listed 11 paralectotypes of colour morph A ( MZUT An725.2–12 ) and 13 of colour morph B ( MZUT An729 ), resulting in a total of 25 type specimens, although their numbers of individuals attributed to both colour morphs differ from the original description ( Peracca 1893 ). However, since the total number of type specimens is in accordance with the number mentioned by Peracca (1893) the claim by Blommers-Schl̂sser and Blanc (1991) concerning the presence of two other syntypes at Paris ( MNHN 1894.1 and 1894.2 ) does not seem to be correct ( Gavetti & Andreone 1993: 106 ) and the same must be assumed for the specimens claimed by Boulenger (1895 ‘1894’) to have been received by the BMNH from Peracca. An additional non-type specimen is MZUT An917 from Andrangoloaka ( Gavetti & Andreone 1993 ), demonstrating that more than the 25 types were available in the MZUT collection. The MNHN and BMNH specimens are therefore not paralectotypes. Identity .—In this study we obtained genetic data via barcode fishing from specimen MNHN 1894.1 from Andrangoloaka, marked as ‘type’ of M. alutus in the MNHN catalogue and provided by M.G. Peracca according to the MNHN catalogue, but, as discussed above, probably not representing one of the paralectotypes . The 16S sequence of this specimen clustered among specimens from the central highlands of Madagascar that are typically considered as M. alutus (BlommersSchl̂sser 1979; Blommers-Schl̂sser & Blanc 1991; Glaw & Vences 1992a , 1994 , 2007). Although this information does not refer to the lectotype of the species, little doubts thus remain that the nomen M. alutus has been correctly applied to this small-sized lineage from the central highlands. FIGURE 10. Mantidactylus curtus in life, in dorsolateral and ventral view. (a,b) Female specimen (ZSM 768/2001 = FGMV 2001.489) from Itremo, photographed in 2001 (note the almost complete absence of femoral glands). (c,d) Probable female (note small femoral glands) from Antoetra, photographed in 2003. (e,f), Probable female (ZSM 758/2001 = FGMV 2001.423; note rather small femoral glands) from Mount Ibity/Col des Tapias, photographed in 2001. TABLE 5. Morphometric measurements (all in mm) of voucher specimens of the Mantidactylus curtus clade. Type status is given in square brackets after voucher number: HT, holotype; PT, paratype; LT, lectotype; PLT, paralectotype. An asterisk (*) marks lectotypes designated in the current paper. For abbreviations of measurements, see Materials and Methods. All specimens are probable adults, but sex is difficult to ascertain externally in many species of this clade due to weak expression of femoral glands in male specimens, and is therefore only given for some specimens where gonad examination was possible. NM, not measured. # Measurements taken by Gavetti and Andreone (1993)

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL
M. alutus
MZUT An725.1 # [LT]	NA	M	Andrangoloaka	23.4	8.9	8.7	3.6	3.2	2.4	2.0	NM	NM	5.8	NM	18.6	NM	NM
MNHN 1894.2	NA	M	Andrangoloaka	27.2	8.4	11.0	3.3	1.9	2.7	1.5	3.0	14.8	6.4	40.9	18.9	13.0	11.9
MNHN 1894.1	NA	M	Andrangoloaka	25.0	9.5	9.9	2.8	2.2	2.6	1.7	3.0	15.0	7.8	41.7	20.8	14.3	11.6
ZSM 194/2021	ACZCV 341	F	Ankaratra	31.0	10.4	11.6	4.0	2.9	2.2	1.9	3.2	18.1	9.0	NM	NM	NM	13.0
M. ambohimitombi ambohimitombi (Ca19)
BMNH 1947.2.26.25 [LT*]	NA	Ambohimito- mbo forest	65.0	25.1	24.0	6.6	4.1	4.5	NA	5.8	36.6	17.4	110.3	52.5	35.2	NM
BMNH 1947.2.26.26 [PLT]	NA	Ambohimito- mbo forest	63.9	23.1	23.0	6.6	4.8	4.7	NA	5.6	35.5	17.1	99.2	48.2	35.1	NM
ZSM 191/2021	ACZCV 335	F	Analafohy	50.6	16.8	17.5	5.9	4.0	3.9	4.0	4.8	27.3	13.0	NM	NM	NM	23.7
ZSM 192/2021	ACZCV 336	F	Analafohy	38.9	12.9	15.0	5.0	2.7	2.6	2.4	4.9	23.0	11.6	NM	NM	NM	20.0
ZSM 193/2021	ACZCV 340	F	Ankaratra	33.5	11.4	12.7	4.4	3.0	2.7	2.6	4.0	19.7	11.0	NM	NM	NM	17.3
ZMA 6865 (1062)	NA	F	Manjakotompo	43.1	14.0	15.8	3.7	2.5	3.6	2.9	3.8	23.1	10.4	67.6	34.2	22.3	22.9
ZSM 190/2021	ACZCV 334	M	Analafohy	37.7	14.0	15.2	5.4	4.1	3.0	2.4	3.6	22.3	11.4	NM	NM	NM	18.0
ZSM 195/2021	ACZCV 342	M	Ankaratra	35.3	12.5	14.1	4.4	2.7	2.6	2.2	3.6	21.3	11.0	NM	NM	NM	17.3
ZSM 368/2000	FG/MV 2000.87	M	Manjakatompo	32.6	12.0	13.7	3.7	2.5	2.6	1.8	2.7	22.8	9.9	60.4	29.1	21.0	17.7
ZSM 369/2000	FG/MV 2000.88	M	Ankaratra	37.2	13.8	14.9	3.8	3.1	2.8	2.3	3.3	23.8	10.5	63.6	30.7	21.0	18.7
ZMA 6863 (671)	NA	?	Manjakotompo	34.3	12.4	13.9	3.9	2.6	2.3	2.2	3.6	19.7	9.2	58.0	27.7	19.2	17.4
ZMA 6863 (672)	NA	?	Manjakotompo	36.7	13.4	15.3	3.7	2.7	2.8	2.7	3.7	25.3	10.9	70.1	33.4	22.8	20.9
ZMA 6866 (1192)	NA	?	Manjakotompo	41.0	15.1	17.0	4.7	3.6	2.9	2.7	4.6	21.2	11.2	59.7	29.6	21.0	18.9
ZMA 6866 (1193)	NA	?	Manjakotompo	30.7	11.1	12.7	2.9	2.1	2.5	2.4	3.4	17.3	8.1	52.1	25.3	17.5	16.2
M. ambohimitombi miloko ssp. nov.
ZSM 219/2005 [HT]	FGZC 2143	M	Ambohitantely	41.7	15.3	16.5	5.5	4.3	3.3	2.9	4.0	23.9	11.8	67.0	32.1	22.2	19.6
ZSM 237/2005 [PT]	FGZC 2172	F	Ambohitantely	49.4	16.5	17.8	5.8	4.1	4.0	3.1	4.4	25.5	12.5	75.2	36.4	25.5	22.9

...Continued on thte next page TABLE 5. (Continued)

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL
ZMA 6859 (675)	NA	?	Tampoketsa d’Ankazobe	32.9	10.9	12.6	3.1	2.2	2.2	2.1	3.3	20.8	9.1	52.8	25.9	27.4	16.5
ZMA 6859 (676)	NA	?	Tampoketsa d’Ankazobe	39.8	13.9	15.4	4.5	2.6	3.1	2.8	4.2	23.4	11.0	64.5	31.8	21.8	19.4
ZMA 6860 (910)	NA	?	Tampoketsa d’Ankazobe	39.0	13.7	14.7	4.1	2.1	3.0	3.1	4.0	23.8	11.1	69.4	33.1	23.4	21.2
ZMA 6860 (911)	NA	?	Tampoketsa d’Ankazobe	43.7	14.6	16.6	4.6	2.4	3.1	3.0	4.2	25.0	11.3	76.3	36.7	25.5	23.2
ZMA 6860 (912)	NA	?	Tampoketsa d’Ankazobe	43.8	15.4	16.6	4.9	2.5	3.0	2.7	4.3	23.9	11.6	70.9	34.2	23.7	21.1
ZMA 6860 (913)	NA	?	Tampoketsa d’Ankazobe	41.3	13.8	14.8	4.3	2.2	2.9	3.3	4.3	25.6	13.7	73.1	36.0	25.4	22.4
ZMA 6860 (914)	NA	?	Tampoketsa d’Ankazobe	35.9	14.5	15.0	3.9	2.6	3.1	2.3	3.4	24.2	11.2	65.9	31.5	22.9	19.7
ZMA 6860 (917)	NA	?	Tampoketsa d’Ankazobe	42.2	14.9	16.9	4.6	2.4	3.4	3.1	4.2	25.9	11.5	73.4	35.8	24.9	21.6
ZMA 6861 (732)	NA	?	Angavokely Carion	36.5	12.5	14.8	3.4	3.0	3.2	2.3	3.7	21.1	10.4	64.4	30.8	20.7	18.7
ZMA 6862 (1100)	NA	?	Angavokely Carion	41.5	13.9	15.8	4.3	2.7	2.9	2.6	4.2	22.9	10.3	64.9	30.0	20.8	19.6
ZMA 6862 (1101)	NA	?	Angavokely Carion	43.5	14.4	16.4	4.1	3.1	3.4	2.7	4.2	24.1	11.3	66.1	33.2	23.4	20.2
ZMA 6862 (1103)	NA	?	Angavokely Carion	43.1	14.3	15.8	3.8	2.3	3.3	3.1	3.9	23.9	11.9	67.4	32.6	23.0	20.4
ZMA 6862 (1105)	NA	?	Angavokely Carion	44.0	15.2	16.7	4.8	2.9	3.2	2.6	4.4	24.0	12.3	68.5	33.4	23.5	21.1
M. ambohimitombi marefo ssp. nov. (Ca20)
ZSM 762/2001 [HT]	FGMV 2001.493	M	Itremo	33.4	12.8	12.2	3.8	2.7	1.7	1.7	3.0	20.3	7.6	48.9	22.0	16.3	14.3
ZSM 761/2001 [PT]	FGMV 2001.492	F	Itremo	35.9	12.8	12.2	4.3	2.2	1.6	1.6	3.5	20.7	8.5	51.5	24.2	17.8	15.7

...Continued on thte next page TABLE 5. (Continued )

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL
MRSN A6747 [PT]	FAZC 14047	F?	Antsirakambiaty forest	32.8	11.9	12.4	4.7	3.0	2.8	2.0	3.4	17.3	9.1	50.5	23.9	17.0	15.2
ZSM 759/2001 [PT]	FGMV 2001.476	M	Itremo	34.1	12.3	12.6	3.4	2.4	2.0	1.9	3.6	19.4	8.6	51.7	23.8	17.0	15.0
ZSM 760/2001 [PT]	FGMV 2001.478	M	Itremo	30.9	12.2	11.6	3.4	1.8	1.9	1.5	3.4	20.5	8.3	52.5	24.4	16.7	15.6
ZSM 763/2001 [PT]	FGMV 2001.495	M	Itremo	34.6	13.7	13.5	3.7	2.8	1.8	1.8	3.0	19.2	8.6	52.2	23.2	16.3	14.9
M. bourgati
MNHN 1972.437 [HT]	NA	M?	38.6	15.7	15.0	4.8	3.5	2.7	2.9	4.4	23.4	11.6	64.4	29.4	20.1	19.6
ZSM 769/2001	FGMV 2001.517	F	Andringitra, Andohariana	32.3	12.5	12.6	3.4	2.3	2.9	2.7	3.6	20.0	9.0	54.0	25.6	17.5	17.2
ZSM 770/2001	FGMV 2001.520	M	Andringitra, Andohariana	33.0	13.2	NM	3.3	2.8	2.6	1.9	3.4	21.4	9.4	55.5	26.5	18.3	17.8
ZSM 771/2001	FGMV 2001.521	F	Andringitra, Andohariana	34.0	11.9	12.7	3.6	2.7	2.5	1.8	3.2	21.5	9.6	55.5	25.7	18.1	17.2
ZSM 772/2001	FGMV 2001.522	F	Andringitra, Andohariana	39.3	14.7	15.1	4.0	2.7	3.7	3.2	4.3	23.1	11.1	65.4	28.7	20.3	19.8
ZSM 773/2001	FGMV 2001.523	F	Andringitra, Andohariana	40.0	14.8	15.1	4.2	3.0	2.5	2.0	4.0	23.4	10.5	63.8	29.0	20.5	19.6
ZSM 774/2001	FGMV 2001.580	M	Andringitra, Imaitso	38.9	14.9	15.4	4.1	3.4	3.5	2.8	3.5	22.7	10.0	60.1	26.4	19.0	18.4
ZSM 775/2001	FGMV 2001.581	F	Andringitra, Imaitso	35.7	13.9	13.6	3.9	2.3	3.1	2.5	3.4	21.5	10.2	60.6	27.3	19.3	19.2
M. curtus
BMNH 1947.2.10.30 [LT*]	NA	M?	35.3	11.8	12.6	4.4	2.5	2.7	2.0	3.9	20.8	9.4	56.1	25.8	17.4	NM
BMNH 1947.2.10.28 [PLT]	NA	F	38.4	14.1	14.2	4.6	3.4	2.7	2.8	4.5	20.5	11.0	55.5	26.0	17.8	NM
ZSM 757/2001	FGMV 2001.422	F	Ibity	36.0	13.0	13.1	3.9	2.3	2.4	2.0	3.4	20.4	10.2	55.3	25.3	18.0	16.9
ZSM 758/2001	FGMV 2001.423	F	Ibity	32.5	12.4	12.2	3.5	1.9	2.5	2.2	3.4	20.3	9.3	53.0	23.5	16.6	15.4

...Continued on thte next page TABLE 5. (Continued )

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL
ZSM 766/2001	FGMV 2001.473	F	Itremo	39.0	13.5	13.9	4.1	2.6	2.4	2.2	3.4	22.6	10.3	60.1	28.0	19.4	18.7
MRSN A6748	FAZC 14046	F?	Antsirakambiaty	41.2	13.8	14.3	5.0	3.1	2.9	2.2	3.8	20.2	10.0	58.6	27.4	19.4	17.4
MRSN A6760	FAZC 13995	F?	Antsirakambiaty	33.5	11.9	12.4	4.5	2.9	2.5	2.2	2.9	18.5	9.0	51.4	23.0	16.4	15.6
MRSN A6757	FAZC 13993	M	Antsirakambiaty forest	33.7	12.5	12.3	4.3	3.4	2.4	2.1	3.8	18.3	9.5	45.4	19.3	16.9	15.2
ZSM 764/2001	FGMV 2001.462	F	Itremo	37.8	14.4	14.1	4.1	2.0	2.5	2.3	3.7	21.6	9.7	57.9	26.2	17.7	16.9
ZSM 765/2001	FGMV 2001.463	F	Itremo	34.3	13.1	12.6	3.6	1.9	2.4	1.9	3.1	21.1	9.0	59.3	27.4	19.0	17.5
ZSM 767/2001	FGMV 2001.488	F	Itremo	39.8	13.7	13.6	3.6	2.0	2.5	2.0	3.6	22.2	9.7	61.3	28.5	18.9	18.2
ZSM 768/2001	FGMV 2001.489	F	Itremo	37.9	14.0	13.9	3.6	2.5	2.2	1.8	3.3	22.1	8.8	52.9	25.7	17.9	16.5
M. madecassus
MNHN 1953.246 [LT]	NA	F?	Andringitra	27.5	9.6	9.7	3.1	1.9	1.8	2.2	3.0	16.6	7.9	47.0	22.4	15.3	NM
MNHN 1989.3592 [PLT]	NA	F?	Andringitra	31.0	9.6	10.2	3.0	2.3	1.8	2.0	3.0	16.7	7.6	43.1	21.0	14.5	NM
MNHN 1989.3594 [PLT]	NA	F?	Andringitra	29.2	10.6	10.3	3.2	2.9	2.0	2.2	3.3	16.3	7.7	43.7	17.2	14.7	NM
MNHN 1989.3591 [PLT]	NA	M?	Andringitra	25.4	8.0	8.6	2.2	1.6	1.7	1.8	3.0	12.7	6.2	36.1	21.3	12.0	NM
MNHN 1972.1182	NA	F	Andringitra	32.0	10.8	11.1	3.5	2.3	1.9	2.6	3.2	17.2	8.5	51.0	24.8	18.3	NM
MNHN 1972.1185	NA	F	Andringitra	32.4	10.9	11.0	3.3	2.2	2.1	2.7	3.1	17.1	8.7	50.5	25.2	17.7	NM
MNHN 1972.1192	NA	F	Andringitra	33.7	10.9	11.2	3.5	2.6	2.0	2.5	3.4	18.0	8.4	49.6	25.1	16.7	NM
MNHN 1972.1204	NA	F	Andringitra	29.3	9.5	10.0	3.1	2.0	1.8	2.2	3.3	16.2	7.7	44.7	25.3	13.1	NM
MNHN 1972.1206	NA	F	Andringitra	31.4	10.5	10.2	3.5	2.1	2.1	2.3	3.4	19.0	8.5	51.4	21.4	17.2	NM
MNHN 1972.1198	NA	M	Andringitra	27.0	9.4	9.9	3.3	2.0	1.7	2.4	3.0	15.5	7.8	45.4	20.1	15.0	NM
MNHN 1972.1199	NA	M	Andringitra	29.8	9.9	10.7	3.4	2.9	2.0	2.1	3.3	17.6	8.8	48.6	23.7	16.8	NM
MNHN 1972.1190	NA	M?	Andringitra	27.0	9.2	9.9	3.1	1.7	2.0	2.2	2.8	15.3	7.0	42.4	21.1	14.2	NM
ZSM 755/2001	FGMV 2001.538	?	Andringitra, Cuvette Boby	29.7	10.0	9.8	3.4	2.2	1.8	1.4	2.6	16.7	7.7	48.1	23.0	16.1	14.4

...Continued on thte next page TABLE 5. (Continued)

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL
M. pauliani
MNHN 1972.1508 [HT]	NA	M	Ankaratra	31.0	11.0	11.0	3.1	2.5	1.8	2.3	2.6	18.0	8.9	50.4	25.7	17.2	NM
MNHN 1972.1510 [PT]	NA	F	Ankaratra	33.7	11.5	10.8	4.0	2.2	1.6	2.2	3.1	18.7	8.9	48.7	24.2	17.0	NM
MNHN 1972.1511 [PT]	NA	F	Ankaratra	31.1	10.7	10.8	3.3	2.0	1.5	2.1	2.7	17.6	8.3	47.1	23.8	16.0	NM
MNHN 1972.1515 [PT]	NA	F	Ankaratra	32.9	10.6	11.1	3.4	2.2	1.6	2.0	3.1	18.2	9.6	49.1	23.8	16.7	NM
MNHN 1972.1509 [PT]	NA	F?	Ankaratra	29.7	11.2	11.7	3.1	2.8	1.9	2.1	3.1	17.4	9.0	47.5	24.4	17.0	NM
MNHN 1972.1512 [PT]	NA	F?	Ankaratra	27.7	10.0	10.5	3.4	2.2	1.5	2.1	3.0	17.5	8.7	42.1	24.2	17.0	NM
MNHN 1972.1513 [PT]	NA	F?	Ankaratra	25.8	9.6	9.7	3.4	2.1	1.7	1.9	2.7	16.6	8.4	44.9	23.1	15.8	NM
MNHN 1972.1514 [PT]	NA	M	Ankaratra	29.5	11.0	11.3	3.3	2.7	1.6	2.1	3.0	17.7	9.0	47.4	24.5	16.7	NM
MNHN 1972.1516 [PT]	NA	M?	Ankaratra	31.6	11.5	11.9	3.5	3.1	1.7	2.3	3.0	18.9	8.9	50.1	25.5	17.3	NM
ZMA 6803 (1184)	NA	F	Ankaratra	31.6	10.1	10.4	3.4	2.0	1.4	2.3	3.0	16.5	8.1	45.5	22.1	15.6	NM
ZMA 6803 (1185)	NA	F?	Ankaratra	27.5	9.2	9.3	3.4	2.3	1.4	1.7	3.1	15.7	7.8	42.8	21.8	15.2	NM
ZMA 6803 (1186)	NA	F?	Ankaratra	26.9	8.8	9.2	3.2	1.6	1.4	1.8	2.7	16.0	8.6	44.8	21.4	14.7	NM
ZMA 6803 (1187)	NA	F?	Ankaratra	24.4	8.6	9.1	3.1	1.6	1.3	1.7	2.8	15.0	7.4	41.6	20.2	14.0	NM
ZMA 6803 (1188)	NA	M?	Ankaratra	24.8	8.8	9.1	3.2	1.8	1.5	1.8	2.7	15.4	8.0	41.1	20.5	14.0	NM
ZMA 6803 (1184)	NA	?	Ankaratra, 2200 m	31.2	10.2	10.5	3.5	1.8	1.4	1.5	2.8	16.0	7.9	45.6	21.8	14.9	13.6
ZMA 6803 (1185)	NA	?	Ankaratra, 2200 m	27.8	9.1	9.2	3.4	1.8	1.4	1.7	3.1	15.5	7.9	42.0	21.1	15.4	12.9
ZMA 6803 (1186)	NA	?	Ankaratra, 2200 m	26.8	8.9	9.0	3.4	1.6	1.7	1.6	3.0	16.1	7.4	43.8	21.4	15.3	13.2
ZMA 6803 (1187)	NA	?	Ankaratra, 2200 m	24.4	8.5	9.2	3.3	1.6	1.7	1.8	2.7	15.0	7.9	41.3	20.4	14.5	13.1
ZMA 6803 (1188)	NA	?	Ankaratra, 2200 m	24.5	8.7	9.1	3.3	1.5	1.3	1.7	2.7	15.1	7.7	40.8	20.7	14.2	13.5
ZSM 756/2001	NA	?	Ankaratra	25.7	8.6	9.4	3.0	1.5	1.4	1.4	2.2	15.4	7.5	44.1	20.7	14.7	13.8
M. mahery sp. nov. (Ca14)
ZSM 23/2004 [HT]	FGZC 37	M	Isalo	30.4	11.7	12.7	4.3	4.0	3.2	1.8	3.5	17.8	8.3	45.1	21.1	15.0	13.2
ZSM 136/2006 [PT]	FGZC 940	F	Bemaraha	45.0	17.0	16.3	5.8	4.9	3.7	3.5	4.6	23.0	12.4	59.9	27.6	19.7	17.9

...Continued on thte next page TABLE 5. (Continued )

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL
ZSM 26/2004 # [PT]	FGZC 42	F	Ranohira,creek nearby	36.0	13.8	13.0	4.7	3.5	3.3	2.6	3.3	11.2	9.5	NM	NM	16.4	15.6
ZSM 567/2009 [PT]	ZCMV 11457	F	Makira (Sa- haovy)	41.6	13.9	14.8	5.2	3.6	4.0	2.1	3.7	22.8	12.1	60.9	21.9	19.5	17.5
ZSM 569/2009 [PT]	ZCMV 11484	F	Makira (Sa- haovy)	43.2	15.3	16.2	6.2	4.3	3.7	2.7	4.2	23.1	11.1	60.4	27.6	19.3	17.4
ZSM 61/2006 [PT]	FGZC 793	F	Bemaraha	48.6	17.6	18.4	6.4	4.8	4.2	2.9	4.6	24.5	12.4	64.5	30.0	21.1	18.9
ZSM 942/2003 [PT]	FGMV 2002- 1486	F	Isalo	34.2	12.8	13.3	4.4	3.8	3.2	2.7	3.5	17.9	9.0	49.0	22.5	16.1	14.6
ZSM 134/2006 [PT]	FGZC 938	M	Bemaraha	33.7	13.1	14.1	5.2	4.3	3.0	2.7	3.3	18.0	9.7	48.1	22.1	15.8	14.7
ZSM 135/2006 [PT]	FGZC 939	M	Bemaraha	33.2	12.4	12.8	4.9	4.3	3.2	2.2	3.4	17.4	8.8	44.7	20.9	15.0	13.2
ZSM 25/2004 # [PT]	FGZC 39	M	near Ranohira	31.0	12.1	11.2	4.0	3.7	2.8	1.7	3.5	9.6	8.5	NM	NM	13.8	13.4
ZSM 9/2006 [PT]	FGZC 682	M	Bemaraha	37.2	14.2	14.1	5.0	4.0	2.8	2.5	3.8	19.7	10.1	49.9	23.2	15.8	15.9
ZSM 941/2003 [PT]	FGMV 2002- 1485	M	Isalo	30.1	12.5	13.2	3.9	3.9	3.0	2.0	3.2	16.8	8.4	45.4	21.1	15.1	13.3
ZSM 927/2003 # [PT]	FGMV 2002.1421	M	Isalo	29.2	12.3	11.2	4.2	3.5	3.2	2.0	3.3	8.5	9.4	NM	NM	15.0	14.6

Synonyms .— The taxon Mantidactylus laevis Angel, 1929 was listed as a dubious species by Guibé (1978) and considered a synonym of M. alutus by Glaw and Vences (1992a) . The holotype MNHN 1929.208, collected by G. Petit from the vicinity of Antananarivo (type locality ‘environs de Tananarive’ ; SVL 32 mm according to the original description) was reported to be lost according to Guibé (1978) , and this has been confirmed by S. Grosjean (pers. comm. to A. Ohler , 22 January 2022 ). Because M. alutus is the only species of Brygoomantis occurring in the Antananarivo area, little doubts remain that this synonymy is correct. In order to stabilize synonymies and as the holotype has been lost for more than 40 years, we here designate the lectotype of Rana aluta , MZUT An 725.1 as the neotype of Mantidactylus laevis Angel, 1929 . Diagnosis .—A member of the M. curtus clade and sister to a monophyletic group of all other species of the clade. See Table 4 for a list of diagnostic morphological characters. The combination of relatively small body size of up to 31 mm , slightly granular skin with (weakly expressed) dorsolateral ridges, small tympanum diameter of ca 9% of SVL in males, absence of small white spots on flanks, presence of a light frenal stripe in most specimens, and advertisement calls as a regular series of short pulsed notes distinguishes M. alutus from species of the other clades. Within the M. curtus clade, M. ambohimitombi , M. bourgati and M. curtus have larger body sizes, and M. madecassus and M. pauliani are high-elevation endemics with usually a shorter snout and absence of dorsolateral ridges ( Table 4 ). For detailed distinction from new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. alutus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation .—Variation in measurements is given in Table 5 . See Fig. 11 for colouration in life and its variation. Some individuals have a light vertebral stripe. There is moderate sexual size dimorphism (confirmed male SVL 23.4–27.2 mm [n = 2] vs confirmed female SVL 31.0 mm [n = 2]). Males with large and distinct femoral glands (e.g. Fig. 11b ); in MNHN 1894.1 and 1894.2, FGL and FGW are 2.4 mm x 1.4 mm and 1.7 mm x 1.4 mm , respectively. The glands can be of orange/yellow colour in some individuals in life (e.g. Fig. 11b ), which may be related to the reproductive state. Natural history.— Typically found in slow-flowing parts of streams or associated swamps in Madagascar’s highlands. Often at the edge of forest or in streams devoid of forest, with some gallery vegetation only. Males call from the edge or very shallow parts of water, from concealed positions during the day or from more open positions at night. See Vences et al. (2002) for observations from Ankaratra. This species is also abundant in parks of Antananarivo , e.g. in the Tsimbazaza garden (Glaw & Vences 2007 ). Calls.— The advertisement call of M. alutus , recorded on 21 January 2003 , 17:30 h, near Antoetra, 20.5–21.0°C air temperature ( Vences et al. 2006 : CD 2, track 61, cut 1), consists of a short, regularly pulsed note, emitted in series at regular intervals ( Fig. 12 ). Notes exhibit distinct amplitude modulation, with amplitude continuously increasing from the beginning, reaching maximum call energy at the middle of the note, before continuously decreasing towards the note’s end. Numerical parameters of eight analysed calls are as follows: call duration (= note duration) 149–290 ms (234.3 ± 53.5 ms); 20–27 pulses per note (24.1 ± 2.7); pulse duration 5–8 ms (6.3 ± 1.0 ms); pulse repetition rate within notes 83.3–130.4 pulses/s (108.2 ± 17.3); dominant frequency 1040–1116 Hz (1081 ± 26 Hz); prevalent bandwidth 600–4800 Hz; call repetition rate (= note repetition rate) within regular series ca 110 calls/min. Calls recorded on 1 January 1994 , 18:20 h, at Manjakatompo, 18°C air temperature ( Vences et al. 2006 : CD 2, track 61, cut 2) generally agree in character with the calls described above. Calls of a series containing nine calls have the following parameters: call duration (= note duration) 254–310 ms (278.5 ± 16.8 ms); 21–27 pulses per note (23.7 ± 2.2); pulse duration 3–6 ms (4.9 ± 0.7 ms); pulse repetition rate within notes 66.7–120.0 pulses/s (83.2 ± 19.0); dominant frequency 1270–1378 Hz (1326 ± 45 Hz); prevalent bandwidth 800–5600 Hz; call repetition rate (= note repetition rate) within series ca 115 calls/min. Tadpoles .—The tadpoles of M. alutus were described by Blommers-Schl̂sser (1979) and Schmidt et al. (2009) . Distribution.— Endemic to the central highlands of Madagascar ( Fig. 7 ). This species is known from Antananarivo , Ankaratra, Ibity, Ambohitantely, Ranomafanakely,Antoetra, a swamp in the Alaotra region, Mantasoa, Forêt de Tampina, and Tsinjoarivo. Elevation range: 933–2090 m a.s.l. Etymology.— Probably derived from the Greek adjective ἄλουτος, meaning ‘unwashed’ or ‘speckled’, presumably in reference to the dorsal colour pattern.