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

TABLE 4. Summary of the most important diagnostic characters of species in the Mantidactylus subgenus Brygoomantis. SVL is given only based on reliably sexed specimens, summarised and rounded to the nearest mm from original measurements in Tables 5‒11. Dorsolateral ridges are always weakly expressed in Brygoomantis, but clearly visible in some species while completely absent in others. Dorsal skin rugosity is estimated from pictures in life and can appear different in preservative: 0, smooth; 1, finely granular; 2, coarsely granular / tubercular. Webbing at toe 5 is given as 0, half or less than half of one phalanx free of web, not counting terminal phalanx (corresponding to numbers 0–0.5 in webbing formula); 1, about one phalanx free of web (0.75–1.25); 2, more than one phalanx free of web (1.5–2). (Yes) in parentheses indicates that the respective character state is found in some individuals of the respective species, and often only weakly expressed, while it is missing in others. Lat.met., state of the lateral metatarsalia, either separated (sep.) by webbing or connected (conn.) by tissue. NM, no measurements available. *For the advertisement calls of M. katae sp. nov. and M. fergusoni sp. nov., homology of call components remains unclear. Here, each unit is interpreted as single–pulse call (= single-pulse note) arranged in series (under this definition the provided pulse rate reflects call rate), but it would also be possible to consider the entire vocalization as single-note multi-pulsed call with very long duration, containing very widely spaced pulses. Values for pulse rate (pulses/second) are rounded. We here evaluate the presence versus absence of regular call series (last column), where regular series are considered as such, if calls are emitted at really regular intervals in series of similar duration containing a more or less defined number of calls. # In M. ambohimitombi, several unsexed individuals of the type series, probably females, reach sizes of up to 65 mm, being the largest of all Brygoomantis known. †One female of M. biporus of 19.3 mm SVL is probably a not fully mature specimen; other females of the species measure 31–36 mm SVL.

	SVL (M)	SVL (F)	(END	HTD/	FGW/SVL	SVL/	Dorsolat.	Dorsal	Lat. met.	Webbing	White	White	Yellow	Pulses	Pulses /	Regular
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Species	[mm]	[mm]	+ NSD) / SVL [%]	SVL (males) [%]	(males) [%]	HIL [%]	ridges	skin		toe 5	spots on flanks	marking on snout tip	inguinal marking	/ call	second	call series
M. curtus clade
M. curtus	34	33–40	11–14	10	8	58–74	No	0	sep.	0	No	No	No	NM	NM	NM
M. alutus	25	27–31	13–17	9	6	60–67	Yes	1	sep.	1	(Yes)	No	No	20–27	67–130	Yes
M. amb. ambo- himitombi	33–38	34–51 (65#)	13–16	8–11	8–10	52–69	(Yes)	1	sep.	0	(Yes)	No	(Yes)	9–14	56–69	No
M. amb. miloko ssp. nov.	42	49	13–16	10	7	54–66	No	1	sep.	1	(Yes)	No	No	NM	NM	NM
M. amb. marefo ssp. nov.	31–35	36	9–15	6–8	NM	59–70	No	0–1	sep.	0	No	No	No	NM	NM	NM
M. bourgati	33-39	32–40	11–18	NM	NM	59–65	(Yes)	1	sep.	0	No	No	No	36–50	46–54	No
M. madecassus	27–30	29–34	11–16	7–10	NM	59–72	No	0	sep.	0	No	No	No	NM	NM	NM
M. pauliani	30–31	31–33	9–14	8–9	NM	57–69	No	0–1	sep.	0	No	No	No	NM	NM	NM
M. mahery sp. nov.	29–37	34–49	14–18	11–13	7–9	66–75	(Yes)	1	sep.	0	No	No	No	21–26	108–667	Yes
M. ulcerosus clade
M. ulcerosus	29–36	33–45	15–19	11–14	6–12	66–71	(Yes)	2	sep.	0	No	No	No	45–65	50–220	Yes
M. bellyi	32–41	33–46	12–19	10–17	6–9	66–70	No	1–2	sep.	0	No	No	No	41–46	64–78	No
M. schulzi	21–24	25–29	14–19	14–15	11	61–72	(Yes)	1	sep.	0	No	Yes	No	6–73	55–130	(Yes)
Species	[mm]	[mm]	+ NSD) / SVL [%]	SVL (males) [%]	(males) [%]	HIL [%]	ridges	skin		toe 5	spots on flanks	marking on snout tip	inguinal marking	/ call	second	call series
M. steinfartzi sp. nov.	17–22	22–28	12–18	15–17	10–13	63–76	No	0–1	sep.	0	Yes	(Yes)	No	40–54	70–115	(Yes)
M. betsileanus clade
M. betsileanus	22–29	30–37	15–21	10–16	5–9	50–62	Yes	1	sep.	1	No	Yes	No	107– 201	40–105	No
M. noralottae	33–36	37–40	15–19	10–12	5–7	59	Yes	0–1	sep.	0	(Yes)	No	No	92–108	37–41	No
M. tripunctatus	26–27	33–35	15–20	14–15	8–9	59–72	Yes	1	sep.	1	No	Yes	No	70–80	30–48	No
M. incognitus sp. nov.	27	26	13–16	14	10	54–61	Yes	1	sep.	1	No	Yes	No	NM	NM	NM
M. jonasi sp. nov.	22–25	NM	17–19	14–18	7–10	56–65	Yes	1–2	sep.	1	No	Yes	No	19–72	14–30	No
M. katae sp. nov.	22–27	26–36	15–20	12–15	9–13	55–62	Yes	1	sep.	0	No	Yes	No	(1)*	(10–16)*	(Yes)*
M. kortei sp. nov.	27	35–37	15–17	13	9	60–67	Yes	1	sep.	0	(Yes)	(Yes)	No	12–27	29–69	No
M. riparius sp. nov.	27	22-27	15–18	13	8	61–63	(Yes)	1	sep.	0	(Yes)	(Yes)	No	15–41	49–114	No
M. fergusoni clade
M. fergusoni sp. nov.	25–30	36–42	16–20	10–14	8–9	61–66	(Yes)	2	sep.	0	No	No	No	(1)*	(2–5)*	(Yes)*
M. georgei sp. nov.	28–31	NM	17–18	13–15	8–12	61–66	(Yes)	1–2	sep.	0	No	(Yes)	No	26–85	13–33	No
M. jahnarum sp. nov.	29–30	30–34	15–17	11	10–12	59–70	(Yes)	1–2	sep.	0	No	(Yes)	No	16–45	25–35	Yes
M. marintsoai sp. nov.	29	35–39	15–18	13	NM	61–65	(Yes)	1–2	sep.	1	No	No	No	NM	NM	NM
M. tricinctus clade
M. tricinctus	17–19	18–23	16–21	12–14	9–13	59–72	No	1	conn.	2	No	Yes	Yes	8–19	147–214	Yes
M. grubenmanni sp. nov.	17–18	19–20	15–20	12–16	10–11	61–64	No	1–2	conn.	2	(Yes)	Yes	Yes	3–7	48–182	Yes
Species	[mm]	[mm]	+ NSD) / SVL [%]	SVL (males) [%]	(males) [%]	HIL [%]	ridges	skin		toe 5	spots on flanks	marking on snout tip	inguinal marking	/ call	second	call series
M. gudrunae sp. nov.	20–25	23–29	17–20	11–15	8–13	62–68	No	1	(sep.)	1	No	Yes	Yes	NM	NM	NM
M. biporus clade
M. biporus	28–32	(19†)/31– 36	13–22	10–12	6–9	66–74	No	0–1	sep.	1	Yes	No	No	15–21	130–211	Yes
M. augustini sp. nov.	24	21–25	13–18	13	8	60–65	(Yes)	0–1	sep.	1	Yes	No	No	9–15	44–71	Yes
M. bletzae sp. nov.	NM	26–27	13–15	NM	NM	65–67	Yes	1	sep.	0	Yes	No	No	NM	NM	NM
M. brevirostris sp. nov.	23	28	14–17	12	7	66–70	No	0–1	sep.	2	Yes	No	No	NM	NM	NM
M. eulenbergeri sp. nov.	20–23	25–28	14–17	12–14	7–11	67–75	(Yes)	0	sep.	1	(Yes)	No	No	NM	NM	NM
M. glosi sp. nov.	21	25	15–17	13	7	73–74	Yes	1	sep.	1	(Yes)	No	No	18–23	91–115	Yes
M. stelliger clade
M. stelliger sp. nov.	21–23	31	16–17	12	9–10	65–68	No	1	sep.	2	Yes	No	No	NM	NM	NM
M. inaudax clade
M. inaudax	22–30	27–33	13–16	12–16	6–10	67–76	No	0–1	sep.	1	(Yes)	No	No	17–28	61–143	Yes
M. m. manerana sp. nov.	25–28	28–29	15–17	9–12	7–9	65–69	Yes	1	sep.	1	Yes	(Yes)	No	26–31	63–116	(Yes)
M. m. fotaka ssp. nov.	26	29	14–16	13	9	71–74	Yes	1	sep.	1	Yes	(Yes)	No	NM	NM	NM
M. m. antsanga ssp. nov.	NM	38	14	NM	NM	71	Yes	NM	sep.	0	NM	NM	No	NM	NM	NM