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 10.11646/megataxa.7.2.1 2703-3090 7441023 2FD8C310-6486-4592-92F6-5EB894EBD6AC Mantidactylus gudrunae sp. nov. Identity and justification .—This lineage has been considered as confirmed candidate species M. sp. 7 by Vieites et al. (2009) and M. sp. Ca7 by Perl et al. (2014) . It is a member of the M. tricinctus clade, and strongly differs from the two other lineages in the clade ( M. tricinctus and M. grubenmanni ) by concordant strong divergence in 16S and Rag-1 sequences. Furthermore, it also differs in various morphological features (see Diagnosis below). We are therefore confident that this lineage represents a distinct, evolutionarily isolated separate species. Holotype .— ZSM 146/2004 (field number FGZC 274), adult male , collected by F. Glaw , M. Puente , R.D. Randrianiaina , and M. Teschke (née Thomas) on 7 February 2004 at Manantantely ( 24.983°S , 046.917°E , 20–150 m a.s.l.), Anosy Region , Madagascar . 16S and cox1 barcode sequences of the holotype are available from GenBank (accessions AY848141 and JN133257 ). Paratypes .—A total of six paratypes : ZSM 136/2004 ( FGZC 250 ), ZSM 138/2004 ( FGZC 259 ), two adult males, and ZSM 154/2004 ( FGZC 286 ), adult female , with the same collection data as the holotype ( 7–8 February 2004 ); ZSM 68/2004 ( FGZC 115 ), adult female , collected by F. Glaw , M. Puente , M. Teschke (née Thomas), and R. Randrianiaina on 29–31 January 2004 at ‘Camp 1’ , between Isaka and Eminiminy , Andohahela National Park ( 24.7586°S , 046.8542°E , 247 m a.s.l.); ZSM 95/2004 ( FGZC 167 ), adult male , and ZSM 96/2004 ( FGZC 168 ), adult female , collected by F. Glaw , M. Puente , M. Teschke (née Thomas), and R. Randrianiaina on 31 January 2004 above ‘Camp 1’ , between Isaka and Eminiminy , Andohahela National Park (ca 24.750°S , ca 046.850°E , ca 350 m a.s.l. ) . Additional material .— The following specimens belong to genetically divergent populations and therefore are not included in the paratype series: ZSM 196/2005 ( FGZC 2594 ), adult female , collected by F. Glaw , and P. Bora on 4 February 2005 in the forest at the QMM Climate Station , Sainte Luce ( 24.7798°S , 047.1713°E , 23 m a.s.l. ); ZSM 181/2021 ( ACZCV 375 , extraction ACP 3589, tissue ACZC 8514), ZSM 182/2021 ( ACZCV 376, ACP 3590, ACZC 8515), ZSM 183/2021 ( ACZCV 377, ACP 3591, ACZC 8516), collected by S. Hyde Roberts at Sainte Luce (S9) on 10 October 2016 ; MRSN A7044 ( FAZC 15282 , ACP 0997 , ACZC 4429 ), collected by F. Andreone and G.M. Rosa on 21 February 2012 at Sainte Luce ; MRSN A7045 ( FAZC 15419 , ACP 1053 , ACZC 4485 ) and MRSN A7046 ( FAZC 15427 , ACP 1057 , ACZC 4489 ), one male and one female , collected by F. Andreone and G.M. Rosa on 29 February 2012 at Tsitongambarika , Ivohibe . Diagnosis .— Mantidactylus gudrunae sp. nov. is a member of the M. tricinctus clade as revealed by the phylogenomic analysis, and sister to a monophyletic group comprising M. tricinctus and M. grubenmanni . See Table 4 for a list of diagnostic morphological characters. The combination of small body size (male SVL 20–25 mm , female SVL 23–29 mm ), presence of a whitish marking on snout tip and of a yellow inguinal marking, and absence of white spots on flanks, distinguishes M. gudrunae sp. nov. from members of other Brygoomantis clades ( Table 4 ). Within the M. tricinctus clade, it differs from both M. tricinctus and M. grubenmanni by a slightly larger body size (male SVL 20–25 mm vs < 20 mm ), more strongly expressed webbing on foot, and lateral metatarsalia only partly connected or separated by webbing (vs connected). For detailed distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. gudrunae sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. TABLE 9. Morphometric measurements (all in mm) of voucher specimens of the Mantidactylus tricinctus clade. Type status is given in square brackets after voucher number: HT, holotype; PT, paratype; LT, lectotype; PLT, paralectotype. For abbreviations of measurements, see Materials and Methods. NM, not measured; NA, not applicable.

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL	FGL	FGW
M. tricinctus
MNHN 1931.26 [LT]	NA	F	Befotaka	18.0	6.9	7.8	2.2	1.5	1.7	1.3	2.2	12.6	5.5	30.3	NM	9.1	NM	NA	NA
MNHN 1931.27 [PLT]	NA	M	Vondrozo	17.1	6.3	7.3	2.2	2.3	1.7	1.2	2.4	9.6	4.8	26.6	NM	8.2	NM	2.1	1.6
MNHN 1994.611 [PLT]	NA	M	Befotaka	17.1	6.4	7.1	2.2	2.2	1.6	1.4	2.2	10.0	5.0	26.1	NM	7.1	NM	2.3	1.5
MNHN 1994.612 [PLT]	NA	M	Befotaka	19.2	7.2	7.6	2.3	2.3	1.9	1.7	2.7	10.5	5.4	26.7	NM	8.7	NM	2.7	1.8
MNHN 1994.613 [PLT]	NA	M	Vondrozo	18.1	6.7	7.6	2.7	2.5	2.0	1.7	2.5	11.4	4.8	NM	NM	NM	NM	2.2	1.8
ZSM 177/2006	NA	F	Befotaka- Midongy	19.7	7.0	8.6	3.2	1.9	1.8	1.3	2.2	11.7	5.5	30.5	14.4	9.4	9.4	NA	NA
ZSM 2377/2007	ZCMV 5444	F	Manombo	23.4	8.1	10.2	3.3	2.4	2.9	1.9	2.7	14.4	6.6	38.8	17.8	10.9	11.9	NA	NA
ZSM 2415/2007	ZCMV 5420	M	Manombo	19.1	7.2	8.4	3.4	2.5	2.4	1.5	2.8	12.6	5.6	29.4	14.0	9.0	9.6	2.8	2.0
ZSM 176/2006	BOR 1066	M	Befotaka- Midongy	16.8	6.2	6.9	3.0	2.2	1.6	1.4	2.3	9.9	4.8	25.3	11.7	7.7	7.7	2.9	2.2
M. grubenmanni sp. nov.
ZSM 375/2006 [HT]	ZCMV 1404	M	An’Ala	17.9	6.5	7.6	3.0	2.8	2.4	1.2	2.0	11.4	4.8	28.3	13.3	8.8	8.7	3.0	1.9
ZFMK 62252 [PT]	NA	F	An’Ala	20.3	7.0	8.2	2.6	2.1	1.8	1.3	2.3	13.5	6.0	32.7	NM	10.0	NM	NA	NA
ZFMK 62253 [PT]	NA	F	An’Ala	19.2	6.5	7.7	2.3	2.0	1.8	1.3	2.5	12.7	5.5	30.8	NM	8.7	NM	NA	NA

...Continued on the next page TABLE 9. (Continued)

Voucher	Field number	Sex	Locality	SVL	HW	HL	ED	HTD	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIBL	FGL	FGW
ZFMK 62254 [PT]	NA	F	An’Ala	19.7	6.9	7.9	2.8	1.9	2.2	1.4	2.5	13.1	6.1	32.0	NM	10.4	NM	NA	NA
ZFMK 62251 [PT]	NA	M	An’Ala	16.7	6.0	6.7	2.5	2.2	1.8	1.4	2.0	11.1	4.8	27.4	NM	8.4	NM	2.5	1.8
ZFMK 62255 [PT]	NA	M	An’Ala	17.6	6.3	7.4	2.7	2.5	1.9	1.4	2.1	11.5	5.5	27.5	NM	9.0	NM	2.9	1.7
ZFMK 62256 [PT]	NA	M	An’Ala	17.8	6.5	7.3	2.6	2.4	1.6	1.3	2.1	11.7	5.5	29.4	NM	8.6	NM	2.6	1.7
ZFMK 62257 [PT]	NA	M	An’Ala	17.8	6.5	7.2	2.6	2.1	2.1	1.4	2.2	11.6	5.3	28.0	NM	8.6	NM	2.5	1.8
M. gudrunae sp. nov.
ZSM 146/2004 [HT]	FGZC 274	M	Manantantely	21.1	NM	8.8	3.2	2.8	2.5	1.7	2.6	13.0	5.6	32.9	14.4	9.7	10.0	4.4	2.5
ZSM 183/2021	ACZC 8516	F	Sainte Luce (S9)	28.6	11.1	11.7	4.4	2.9	2.8	2.0	3.3	17.8	8.2	44.9	20.2	13.6	13.4	NA	NA
ZSM 154/2004 [PT]	FGZC 286	F	Manantantely	24.4	10.1	8.9	3.8	2.5	2.5	1.9	3.1	8.5	6.6	NM	NM	10.8	12.5	1.4	1.1
ZSM 196/2005	FGZC 2594	F	Ste. Luce	26.3	10.4	9.5	4.2	2.6	2.7	1.7	2.6	10.6	7.5	NM	NM	12.6	13.7	NA	NA
ZSM 68/2004 [PT]	FGZC 115	F	Andohahela	25.3	10.5	10.4	3.8	2.4	2.5	2.0	2.9	9.8	8.7	NM	NM	12.9	14.3	1.5	1.1
ZSM 96/2004 [PT]	FGZC 168	F	Andohahela	26.5	10.8	10.1	3.9	3.1	2.9	2.2	2.9	10.7	8.2	NM	NM	12.9	14.0	1.5	1.0
MRSN A7046	FAZC 15427	F	Tsitongambarika	22.6	8.0	9.6	3.2	2.2	2.1	2.0	2.8	13.4	6.7	36.3	16.5	11.0	10.7	NA	NA
ZSM 182/2021	ACZC 8515	M	Sainte Luce (S9)	24.4	9.0	10.1	2.7	2.6	2.7	1.6	2.9	14.4	6.8	36.3	16.3	11.1	11.3	3.2	2.5
ZSM 136/2004 [PT]	FGZC 250	M	Manantantely	21.8	8.6	9.4	4.0	3.3	2.4	1.4	2.7	14.6	6.8	34.9	16.4	10.9	10.8	3.7	2.8
ZSM 138/2004 [PT]	FGZC 259	M	Manantantely	20.5	7.9	9.0	3.8	2.9	2.0	1.4	2.6	13.7	5.8	31.8	14.2	9.8	9.3	4.2	2.5
ZSM 95/2004 [PT]	FGZC 167	M	Andohahela	20.2	8.6	7.8	2.9	2.7	2.1	1.7	2.3	8.5	6.2	NM	NM	9.7	10.5	2.3	1.6
MRSN A7044	FAZC 15282	M	Sainte Luce	23.0	8.0	9.6	3.6	2.7	2.6	1.4	2.9	13.6	6.5	33.7	15.2	10.4	11.0	3.2	2.1
MRSN A7045	FAZC 15419	M	Tsitongambarika	21.9	8.0	9.2	3.7	3.0	2.4	1.8	2.8	12.6	6.4	34.5	15.4	10.0	10.2	2.5	2.3

FIGURE 61. Mantidactylus gudrunae sp. nov. in life, in dorsolateral and ventral view. (a,b) Adult male from Manantantely, photographed in 2004. (c,d) Adult male (holotype ZSM 146/2004 = FGZC 274) from Manantantely, photographed in 2004. (e) Unsexed specimen (ACZC 9918 = FAZC 15246) from Sainte Luce. (f) Unsexed specimen (ACZC 4418 = FAZC 15257) from Sainte Luce. (g) Unsexed specimen (ACZC 4447 = FAZC 15324) from Tsitongambarika. (h) Unsexed specimen (ACZC 4489 = FAZC 15427) from Tsitongambarika. Description of the holotype . —Adult male in good to moderate state of preservation ( Fig. 56 ). Tongue removed as tissue sample; femoral glands partly detached for examination in internal view. Body relatively slender. Head as wide as body. Snout rounded in dorsal view, truncate in lateral view. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave. Tympanum distinct and large, rounded, horizontal diameter of tympanum 88% of horizontal eye diameter. Supratympanic fold in its first part almost identical to tympanum edge, thereafter distinct, running rather straight from behind eye and bending about 70° close to posterior edge of tympanum towards forelimb insertion. Maxillary teeth present. Vomerine teeth form two elongated aggregations, directed posteromedially from choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I<II<IV<III. Finger discs slightly enlarged. Nuptial pads absent. Foot slightly shorter than tibia (97%). Lateral metatarsalia separated. Inner metatarsal tubercle present. Outer metatarsal tubercle indistinct but recognisable. Webbing formula: 1(no web), 2i(no web), 2e(traces), 3i(2), 3e(1), 4i(2.5), 4e(2.25), 5(1). Relative length of toes: I<II<V<III<IV. Skin on the upper surface smooth, some granules on flanks (in life, also tubercular on dorsum, without dorsolateral folds or longitudinal ridges). Ventral side smooth. Femoral glands with a large and distinct distal ulcerous macrogland; and a small proximal granular gland field apparently made up of only a few gland granules. Colour in preservative: dorsally almost uniformly brown, with a dark band between the eyes bordering on a lighter colour on the anterior head surface. Some white spots and markings laterally on head. Limbs with dark crossbands. Ventrally, beige on limbs, brown with distinct white pattern on throat, chest and anterior belly. Larger white spots arranged to form a median intrreupted white line on throat. Lower lip ventrally with alternating white/ dark brown pattern. In life, colourarion was similar but more contrasted. A small yellowish marking was present in the inguinal region. The light ventral pattern was bright silvery white. FIGURE 62. Preserved holotype specimens of newly named species in the Mantidactylus biporus clade and of the M. stelliger clade. Scale bars equal 5 mm. Variation .—Variation in measurements is given in Table 9 . See Fig. 61 for colouration in life and its variation. There is weak sexual size dimorphism (confirmed male SVL 20.2–24.4 mm [n = 7] vs confirmed female SVL 22.6–28.6 mm [n = 6]). Males have a larger tympanum than females (HTD/ED ratio is 62–79% in females, 75– 96% in males). Femoral glands of males in life distinct and coloured with a conspicuous yellowish shade; a large and distinct distal ulcerous macrogland is clearly visible, as is a smaller proximal granular gland field. Natural history .—Specimens have been found along slow running water bodies in coastal rainforest. They are active during the night and call from water. Their call is rarely heard. The colouration of this species is quite variable, with some specimens showing an orangish colouration on the arms or on the dorsal stripe. Sometimes reminiscent of the colouration of species in the subgenus Ochthomantis (e.g. Fig. 61f ). Calls .—The calls of this species have not yet been recorded. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to the South East of Madagascar ( Fig. 7 ). This species is known from Andohahela, Manantantely, Sainte Luce, and Tsitongambarika. Elevation range: 23–415 m a.s.l. Etymology .—We dedicate this species to Gudrun Grubenmann from Z̧rich. Together with her husband Moritz, she has been travelling in Madagascar for many decades and has supported our research with important observations of Malagasy amphibians and reptiles.