Search

Amnirana parva sp. nov. Lesser White-Lipped Frog Figs. 5–7 Holotype. ZMB 88458 (field #: RG27, GenBank #: 16S: OQ400914), adult male, Liberia, Krahn-Bassa Proposed Protected Area, 5°39′02.1′′N, 8°39′05.0′′W, 28 March 2018, leg. M.-O. Rödel & J. Glos. Paratypes. Twelve males, ten females: ZMB 71273 (field #: MOR ANK 15, GenBank #: 16S: MG552248), adult female, Ghana, Ankasa National Park, 5°16′38.5212′′N, 2°38′41.8812″W, 5 August 2005, leg. A.C. Agyei & A. Hillers; ZMB 71274 (field #: MOR OWS 28, GenBank #: 16S: MG552262), adult female, Ghana, Owabi Wildlife Sanctuary, 6°44′50.5212″N, 1°42′11.6388″W, July 2005, leg. A.C. Agyei & A. Hillers; ZMB 71276 (field #: MOR LE 11, GenBank #: 16S: MG552287), adult female, Ghana, Leklebi-Agbesia, 6°56′ 1.3812″N, 0°29′1.5612″W, 19 July 2005, leg. A.C. Agyei & A. Hillers; ZMB 79187 (field #: GO14), adult male, Liberia, Gola National Forest, 7°27′10.6812″N, 10°41′31.3188″W, 28 November 2005, leg. A. Hillers; ZMB 79197 (field #: G84), adult female, Ghana, Volta Region, 22 August 2001, leg. M.-O. R ̂del; ZMB 79210 (field #: GOL 060, GenBank #: 16S: MG552310), adult male, Sierra Leone, Gola North Forest Reserve, 7°35′10.14″N, 11°1′30.9612″W, 15 September 2005, leg. A. Barrie & A. Hillers; ZMB 79253 (field #: GRE 010, GenBank #: 16S: MG552307), adult female, Liberia, Grebo National Forest, 5°24′6.48″N, 7°44′0.6612″W, 7 December 2005, leg. A. Hillers; ZMB 87105–06 (field #: P.LI 12 004–P.LI 12 005, GenBank #: 16S: MG552296 & MG552305), adult female and male, Liberia, Gba Community Managed Forest, 7°28′52.6584″N, 8°34′28.5564″W, 28 January 2012, leg. J. Penner; ZMB 87107 (field #: P.LI 12 077, GenBank #: 16S: MG552312), adult male, Liberia, Gba Community Managed Forest, 7°30′34.5636″N, 8° 41′50.7408″W, 3 February 2012, leg. J. Penner; ZMB 87108 (field #: P.LI 12 321, GenBank #: 16S: MG552315), adult male, Liberia, Sapo National Park, 5°29′42.8352″N, 8°21′55.1988″W, 5 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87109 (field #: P.LI 12 466, GenBank #: 16S: MG552323), adult male, Liberia, Mount Ghi South, 5°38′34.9944″N, 8°12′30.1428″W, 29 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87110–11 & 87116 (field #: P.LI 12 503–P.LI 12 505, GenBank #: 16S: MG552319 & MG552320), adult male, female and male, Liberia, Mount Ghi South, 5°39′11.124″N, 8°12′21.708″W, 3 October 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87112 (field #: P.LI 12 300, GenBank #: 16S: MG552322), adult male, Liberia, Sapo National Park, 5°31′20.8272″N, 8°20′46.2084″W, 14 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87113 (field #: P.LI 12 377, GenBank #: 16S: MG552317), adult male, Liberia, Sapo National Park, 5°30′18.5652″N, 8°20′14.9208″W, 18 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87114 (field #: P.LI 12 432, GenBank #: 16S: MG552316), adult female, Liberia, Mount Swa, 6°26′18.0636″N, 9°3′25.9416″W, 24 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87115 (field #: P.LI 12 279, GenBank #: 16S: MG552314), adult female, Liberia, Sapo National Park, 5°31′34.8564″N, 8°20′53.1636″W, 12 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 88438 (field #: RG07), adult female, Liberia, Krahn-Bassa Proposed Protected Area, 06°02′34.8″N, 08°20′10.3″W, 25 March 2018, leg. M.-O. R ̂del & J. Glos; ZMB 88512 (field #: RG80), adult male, Liberia, Foya Proposed Protected Area, 08°00′21.3″N, 10°25′20.2″W, 5April 2018, leg. M.-O. Rödel & J. Glos; ZMB 88549 (field #: RG118, GenBank #: 16S: OQ400915), adult male, Liberia, Foya Proposed Protected Area, 08°03′18.6″N, 10°23′04.0″W, 8 April 2018, leg. M.-O. R ̂del & J. Glos. Additional material. Ten males, ten females; ZMB 90052 (field #: N035), adult female, Guinea, Keoulenta, 7°41′20.6”N, 8°21′19.0”W, 21 September 2009, leg. J. Doumbia, L. Sandberger, K. Camara & F. Gbêmou; ZMB 90053 (field #: N064), adult female, Guinea, Foromota, 7°42′58.3”N, 8°21′41.6”W, 3 November 2009, leg. J. Doumbia, L. Sandberger, K. Camara & F. Gbêmou; ZMB 90054 (field #: S4- Dian 91), adult male, Guinea, Diandian Missidé, 10°59′10.356”N, 13°47′44.556”W, 3 October 2016, leg. J. Doumbia & K. Camara; ZMB 90055 (field #: Ban _0362), adult female, Guinea, Banco, 8°31′6.312”N, 8°56′15.864”W, 17 May 2019, leg. J. Doumbia & K. Camara; ZMB 90056 (field #: G009), adult female, Liberia, Gangra, 7°33′47.556”N, 8°38′15.144”W, 14 January 2009, leg. J. Doumbia, K. Camara & A. T. Johnson; ZMB 90057 (field #: Wata 2_0201), adult female, Guinea, Wataférédou, 8°39′15.192”N, 8°51′11.196”W, 8 May 2019, leg. J. Doumbia & K. Camara; ZMB 90059 (field #: G003), subadult female, Liberia, Gangra, 7°32′40.992”N, 8°37′35.58”W, 12 January 2009, leg. J. Doumbia, K. Camara & A. T. Johnson; ZMB 90060 (field #: G009), adult male, Liberia, Gangra, 7°33′47.556”N, 8°38′15.144”W, 14 January 2009, leg. J. Doumbia, K. Camara & A. T. Johnson; ZMB 90062 (field #: Obay _0548), adult male, Liberia, Obayanmai, 8°8′59.82”N, 9°52′49.764”W, 7 April 2019, leg. J. Doumbia, K. Camara, F. Gbêmou & F. B. Beyan; ZMB 90063 (field #: Obay _0611), adult male, Liberia, Obayanmai, 8°8′56.04”N, 9°52′26.436”W, 9 April 2019, leg. J. Doumbia, K. Camara, F. Gbêmou & F.B. Beyan; ZMB 90159 (field #: PG.L.15/040), adult female, near Bouaflé, Ivory Coast, 7°0′15.1488”N, 5°34′37.0344”W, 7 May 2015, leg. J. Penner & N.L. Gonwouo; ZMB 90161 (field #: PG.L.13 033), subadult female, Liberia, Dugbe, 5°5′15”N, 8°36′29.88”W, 26 August 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90162 (field #: PG.L.13 109), adult male, Liberia, Dugbe, 5°8′40.92”N, 8°29′57.12”W, 3 September 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90163 (field #: LI10 014), adult male, Liberia, Tokadeh, 7°26′40.704”N, 8°39′27.972”W, 11 November 2010, leg. J. Penner; ZMB 90164 (field #: PG.L.13 054), adult female, Liberia, Dugbe, 5°4′20.28”N, 8°33′23.76”W, 28 August 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90165 (field #: LI10 121), adult male, Liberia Tokadeh, 7°27′52.164”N, 8°39′53.928”W, 27 November 2010, leg. J. Penner; ZMB 90166 (field #: LI10 114), adult male, Liberia, Eastern Nimba Nature Reserve, 7°29′3.264”N, 8°34′36.264”W, 26 November 2010, leg. J. Penner; ZMB 90167 (field #: LI10 136), adult male, Liberia, Geipa, 7°28′59.196”N, 8°32′13.344”W, 3 December 2010, leg. J. Penner; ZMB 90168 (field #: PG.L.13 188, adult male, Liberia, Dugbe, 5°5′45”N, 8°29′32.64”W, 13 September 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90169 (field #: LI10 120, adult female, Liberia, Eastern Nimba Nature Reserve, 7°29′3.264”N, 8°34′36.264”W, 28 November 2010, leg. J. Penner. Diagnosis. Amnirana parva sp. nov. differs from other West African Amnirana genetically, morphologically, and acoustically. The uncorrected pairwise-divergence in 16S ribosomal RNA between the new species and its closest congener, A. fonensis, is 5.8%. The new species can be distinguished from A. fonensis by several morphological traits in male specimens. SVL of male A. parva sp. nov. (41.9–50.4 mm) is smaller than in A. fonensis (48.7–65.3 mm). Area of humeral gland is smaller in A. parva sp. nov. (3.7–14.8 mm 2) than in A. fonensis (13.5–26.4 mm 2) as well. Males of A. occidentalis and A. galamensis reach larger SVL (67 and up to 80 mm, respectively; females of these species reach 96 and 86 mm, respectively), have differing color pattern and completely smooth dorsal skin (R̂del & Bangoura 2004; Channing & R̂del 2019). Most males of A. parva sp. nov. have dark spots on the dorsum. Breeding males of A. fonensis differ by a yellow dorsal coloration. The ranges of two Central African Amnirana species that are morphologically similar to A. parva sp. nov., A. albolabris (comprising various lineages, see Jongsma 2018) and A. asperrima (Perret, 1977), do not overlap with the new species. Nominotypical A. albolabris from Central Africa show a larger body size in males (Perret 1977 reports 44–57 mm SVL for A. albolabris from Cameroon). Amnirana asperrima can also be distinguished by both its larger adult body size (SVL of males: 46.7–60.1 mm) and the texture of its dorsal skin, which is densely covered with tiny spines. Other Central African Amnirana species show distinct morphological traits and cannot be confused with Amnirana parva sp. nov.: A. amnicola (Perret, 1977) adults are larger (SVL of males and females: 53–72 mm) and have a more pointed snout; and A. lepus (Andersson, 1903) adults are much larger (SVL: 70–100 mm), have finely granular skin, fully webbed toes, and much longer hind limbs. Last, the new species also differs from populations previously referred to A. longipes (Perret, 1960) that are much larger (SVL 77–87 mm) and have a more robust body and longer hind limbs. Although we follow Jongsma (2018) in recognizing A. longipes as a junior synonym of A. albolabris, we include it here in anticipation that this may be recognized as a valid taxon if one or several lineages of Central African A. albolabris are recognized as distinct species. Description of the holotype. Figs. 5–6; measurements in mm. Adult male with long and slender body, slightly dorsoventrally flattened; snout–vent length 42.9; snout acuminate with rounded tip in dorsal view, narrowly rounded in lateral view; head length 15.8, approximately 27% of SVL; head width 13.6; eye–snout distance 6.7; nostrils directed laterally; nostril closer to the snout (1.7) than to eye (4.7); eye diameter 6.1, slightly larger than tympanum diameter (5.2); interorbital distance 4.3; approximately one third of eye projecting above dorsal margin of head in lateral view; canthus rostralis distinct and sharply protruding; loreal region concave; anterocranial part of upper arm with large protruding gland (3.6 long, 2.0 wide, surface: 7.2 mm 2); long and slender fingers, tips broader than subarticular tubercles, forming discs, reaching 130–200% of finger width, inner metacarapal tubercle large and elongated, outer metacarpal tubercle smaller and more rounded; manual webbing absent; finger formula IIVariation. Measurements (in mm) of all individuals are summarized in Table 5. Overall, the male paratypes were similar to the holotype in external appearance and coloration. Male SVL ranged from 41.9–50.4 (mean ± sd: 45.9 ± 2.9, N= 13). The humeral gland area spanned between 3.7–14.8 mm 2 (9.4 ± 2.7 mm 2, N= 13). Female SVL ranged from 53.6–75.6 (63.9 ± 7.8, N= 10) thus markedly exceeding male SVL (Fig. 7). Seven of 13 showed slightly more extended (1-1), and a single male paratype showed slightly less extended webbing (2-2) of the fourth toe. In female paratypes seven of 10 individuals exhibited a webbing formula with slightly more extended webbing on toe IV(1-1). The dorsal skin was covered with small tubercles in all individuals, but females had fewer, smaller, and less spiny tubercles. In some individuals, the skin appeared almost smooth at first sight, but tiny granules were always present upon detailed inspection. The dorsolateral ridge was interrupted in the majority of specimens: females (six of 10) and males (nine of 13). Females showed a brighter brown dorsum and few to no dark spots (see Figs. 6b, 7b, 8e, g, 11c). On the dorsum of males, the ground color varied from chocolate-brown to dark brown and all males had at least some dark spots. The ventral coloration varied greatly among males. Among the paratypes, some had a plain white throat (e.g., ZMB 88549, 87112), similar to the holotype, whereas others exhibited dusty brown speckling covering almost the entire throat (e.g., ZMB 87113). The belly of all males were white to cream, and some individuals had irregular brown spots (e.g., ZMB 87110) that were densest near the legs, throat, or flanks, but were absent at the center of the venter. Females showed some color variation on the venter. In ZMB 87114, the belly and throat were white, whereas ZMB 79253 and ZMB 79197 had a cream-colored belly and a dusty brown speckled throat. Concerning life coloration (Figs. 8, 11), the dorsum exhibited various shades of brown, sometimes with darker spots (especially in males, most females without darker spots). The flanks in males were greenish yellow (Fig. 8f, h) to chocolate brown dorsally (Fig. 8b), usually turning lighter ventrally (but see Fig. 8a). The flanks in females were olive green, turning paler ventrally (Figs. 8c, e, g, 11c), and becoming almost cream-colored towards the belly (then often with some greenish or brownish mottling). The tympanum was brown in both sexes. Males had a narrow green line surrounding the eyes. In both sexes the iris was golden, sometimes with a reddish tinge, with the lower half of the iris being darker. The limbs and dorsum were a similar shade of brown in both sexes. The limbs sometimes were somewhat lighter, but with darker patterning. The number of conspicuous dark transverse bars on the hind limbs varied among individuals (e.g. compare Figs. 8b & 8h). The venter was beige or cream. Both sexes had a light cream or white upper lip, and rarely the anterior part of the lip was brownish (Fig. 8h). Juveniles showed a more contrasting pattern compared to adults. It consisted of a greenish brown dorsum with black spots and an almost black loreal region and flank. The ventral part of the flank and the belly were white with black dots. The juveniles had more conspicuous dark cross bands on the hind limbs and blackish dots on the forelegs; a bright white upper lip, and a bright red iris (uppermost part golden) (Fig. 11f). Acoustics. The advertisement call of Amnirana parva sp. nov. consists of a whiny-voiced croaking sound with ascending frequency modulation from 932 to 1092 Hz (mean ± sd: 1030 ± 86, N= 3). The sound consists of up to five harmonics of which the second-lowest is the dominant one in the first half, the lowest one in the second half (Fig. 4a). The call duration ranges from 0.26– 0.78 s (0.52 ± 0.26, N= 3). Dominant frequency at the beginning of the call ranges from 948–1034 Hz (1004 ± 49, N= 3). All call characteristics are summarized in Table 4. A call description by Schiøtz (1964c) from central Sierra Leone seems to refer to our new species. In that paper, Schiøtz describes three different sections of the call (such a clear distinction of sections was not detected by us). Call duration and complexity, however, seem to increase with the motivation of the frogs (MOR pers. obs.). Schiøtz (1964c) also mentions that these calls were different from those he recorded in Cameroon for A. albolabris, though he did not provide further details. Distribution. Confirmed A. parva sp. nov. records so far are known from western Sierra Leone to southerncentral Ghana, all within the belt of the (former) rainforest zone, bordering the southern humid Guinean savanna (Fig. 9). So far, no confirmed genetic records exist for Ivory Coast. However, morphologically we could assign frogs from the Taï National Park, the Azagny National Park, the Banco National Park, the Haute Dodo, Cavally and Tanoé forest reserves, as well as the Ivorian parts of Mount Nimba to the new species. Other formerly published records of A. albolabris (see literature cited below) from this country from habitats that include primary lowland evergreen and humid but semi-deciduous forest cannot be unambiguously assigned to either A parva sp. nov. or A. fonensis (see below). Life history. It is difficult in many cases to assign previous field observations and literature records to A. parva sp. nov. or A. fonensis. We summarize observations based on confirmed records (morphology and genetics of vouchers, and/or acoustics) as well as records, which originate from primary humid-evergreen rainforest. The latter decision was based on the results of the environmental niche modelling (see below). All observed and collected individuals of A. parva sp. nov. were encountered in closed primary or slightly degraded lowland rainforest, on the banks of or near to small to medium-sized streams (Fig. 10). In Banco National Park, southeastern Ivory Coast, the species was recorded in the forest, forest patches dominated by bamboos, and in man-made ponds of a fish-farm in a forest clearing (Fig. 11a, b). Although we often heard small choruses, Published as part of Griesbaum, Frederic, Jongsma, Gregory F. M., Penner, Johannes, Kouamé, N'Goran Germain, Doumbia, Joseph, Gonwouo, Nono L., Hillers, Annika, Glos, Julian, Blackburn, David C. & Rödel, Mark-Oliver, 2023, The smallest of its kind: Description of a new cryptic Amnirana species (Amphibia, Anura, Ranidae) from West African rainforests, pp. 301-339 in Zootaxa 5254 (3) on pages 309-319, DOI: 10.11646/zootaxa.5254.3.1, http://zenodo.org/record/7727536, {"references":["R ˆ del, M. - O., Bangoura, M. A. & B ˆ hme, W. (2004) The amphibians of south-eastern Republic of Guinea (Amphibia: Gymnophiona, Anura). Herpetozoa, 17, 99 - 118.","Jongsma, G. F., Barej, M. F., Barratt, C. D., Burger, M., Conradie, W., Ernst, R., Greenbaum, E., Hirschfeld, M., Leache, A. D., Penner, J., Portik, D. M., Zassi-Boulou, A. - G., R ˆ del, M. - O. & Blackburn, D. C. (2018) Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saharan Africa. Molecular Phylogenetics and Evolution, 120, 274 - 285. https: // doi. org / 10.1016 / j. ympev. 2017.12.006","Perret, J. - L. (1977) Les Hylarana (Amphibiens, Ranides) du Cameroun. Revue suisse de Zoologie, 84, 841 - 868. https: // doi. org / 10.5962 / bhl. part. 91429","Schiotz, A. (1964 c) The voices of some West African amphibians. Videnskabelige Meddelelser fra dansk Naturhistorik Forening, 127, 35 - 83.","Lamotte, M., Lauwarier, G. & Perret, J. - L. (1957) Contribution a l'etude des batraciens de l'Ouest Africain V. - Le developpement larvaires de Rana (Hylarana) albolabris. Bulletin de l'Institut fondamental d'Afrique Noire, Ser. A, 19, 1312 - 1327.","Guibe, J. & Lamotte, M. (1958) La reserve naturelle integrale du Mont Nimba. XII. Batraciens (sauf Arthroleptis, Phrynobatrachus et Hyperolius). Memoires de l'Institut Francais d'Afrique Noire, 53, 241 - 273.","Channing, A., R ˆ del, M. - O. & Channing J. (2012) Tadpoles of Africa. The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, Germany, 402 pp.","McIntyre, P. (1999) Hylarana albolabris, predation. Herpetological Review, 30, 223."]}

Redescription of A. fonensis (Rödel & Bangoura, 2004) Figs. 12–14 Holotype. SMNS 11788, adult male, Guinea, Simandou Range, northwest flank of Pic de Fon, 8°36′N, 8°51′W; 6 November 2002, leg. M.A. Bangoura. Paratypes. SMNS 11790, adult male, Guinea, Simandou Range, village situated to the northwest of Pic de Fon, 3 November 2002, leg. M.A. Bangoura; ZFMK 64481–64482, adult males, Guinea, Park National du Haut Niger, November 1996 – February 1997, leg. G. Nikolaus. Referenced specimens. Six males, one female; ZMB 79185 (field #: TI 60, GenBank #: 16S: MG552336), adult female, Sierra Leone, Tingi Hills, 8°52′2.1″N, 10°47′32.1612″W, 4 June 2007, leg. J. Johnny & A. Hillers; ZMB 79203 & 84733 (field #: SI 032–033, GenBank #: 16S: MG552334 & MG552340), two adult males, Guinea, Korombadou, 9°16′49.62″N, 9°6′ 52.0812″W, 5 October 2008, leg. J. Doumbia & A. Hillers; ZMB 79208 (field #: TI 020, GenBank #: 16S: MG552335), adult male, Sierra Leone, Tingi Hills, 8°53′25.44″N, 10°47′24.7812″W, 1 June 2007, leg. J. Johnny & A. Hillers; ZMB 90058 (field #: N044, 09-609), adult male, Guinea, N’Zérékore, 7°35′33.3″N, 8°28′5.4″W, 24 October 2009, leg. J. Doumbia, L. Sandberger, K. Camara & F. Gbêmou; ZMB 90158 (field #: LI10 088), adult male, Liberia, Tokadeh, 7°26′40.704″N, 8°39′27.972″W; 22 November 2010, leg. J. Penner & N.L. Gonwouo; ZMB 90160 (field #: PG.L. 15/041), adult male, Ivory Coast, 7°0′15.1488″N, 5°34′37.0344″W, 7 May 2015, leg. J. Penner & N.L. Gonwouo. For diagnostic differences to A. parva sp. nov., please see the diagnosis of the new species above. Description (based on the type series and seven additional specimens; measurements in mm, see Table 5). Medium-sized, slender frog, SVL of males range from 48.7 to 65.3 (mean ± sd: 57.2 ± 5.2, N= 10); a female measured 68.0; head slightly longer than broad, approximately one third of SVL; snout acuminate with rounded tip in dorsal view, narrowly rounded in lateral view; tympanum diameter around 85% of eye diameter, in two type specimens (SMNS 11788 & 11790) tympanum slightly larger than eye; inter-orbital distance slightly smaller than eye diameter; nostril closer to snout than to eye; canthus rostralis distinct and rounded; loreal region concave; dorsolateral ridge broad and protruding, only two male individuals show slight discontinuities in the posterior third of the ridge; males with large humeral glands on anterocraniad part of upper arms; area of humeral gland spans between 13.5–26.4 mm 2 (20.5 ± 4.86, N= 10); fingers long and thin, without webbing; tips rounded, forming small discs; finger formula: II– d), dark spots on dorsum and bands on hind limbs visible but paler; limbs and yellowish brown; flanks yellow, lighter towards cream belly; lip and throat bright yellow; dorsolateral ridge yellow to reddish brown; tympanum of breeding males with yellow center bordered by pale brown ring; tympanum of females dark or reddish brown (Fig. 13e, f). Acoustics. The advertisement call of male A. fonensis consists of a whiny-voiced croaking sound with an ascending frequency modulation of 548–723 Hz (mean ± sd: 633 ± 87, N= 3). The sound comprises up to five harmonics of which the lowest is the dominant one (see Fig. 4b) and lasts 0.55– 0.92 s (0.7 ± 0.19, N = 3). All call characteristics are summarized in Table 4. Distribution. Amnirana fonensis was described from the Pic de Fon, Simandou Range, and from the Haut Niger National Park, both located in Guinea. Our new records add further localities, including the first records from eastern Sierra Leone, the Nimba Mountains in northern Liberia, and central Ivory Coast. However, we cannot at this point clarify which previous West African records of Amnirana albolabris refer to either A. fonensis or A. parva sp. nov. (but see below). Life history. The observed and collected individuals were found near forest streams. These forests were less humid than in confirmed A. parva sp. nov. records and comprise semi-deciduous and degraded forest fragments (Fig. 14d) surrounded by savanna. Amnirana fonensis males differ from breeding A. parva sp. nov. males, by a bright yellow breeding coloration. Similar yellow breeding color has been reported from many, unrelated frogs across the tropics. In West Africa such color change has been observed in Phrynobatrachus alleni Parker, 1936 (R̂del 2003; Kanga 2021), Sclerophrys maculata (Hallowell, 1854) (Hillers & R̂del 2007) and S. togoensis (Ahl, 1924) (R̂del & Bangoura 2004). Amnirana fonensis breeds in water bodies of different sizes but seems to prefer stagnant or slow-flowing water. In a savanna-like habitat (Foma), males called from inundated patches, well concealed between grasses and reeds. In degraded gallery forests at Wataférédou, males called while floating on the water surface (Fig. 14a), well concealed between roots of trees and shrubs, or between low vegetation at the water edge. One to two males initiated calling and many other males then joined the chorus. Amnirana fonensis called at night as well as during the day. For instance, between 20:35 GMT and 23:00 GMT, we encountered many calling males (N> 100); the peak of calling activity was between 20:40 GMT and 21:30 GMT. Likewise between 14:45 and 16:00 GMT, we observed numerous calling males (N> 70); the peak of calling activity was at 14:57 GMT. In Foma, females attached eggs to grasses emerging from flowing water. After a few hours these eggs became submerged. In Wataférédou, the eggs were deposited between roots of trees and shrubs in the forest streams. The dark brown and yellow eggs were deposited in large clumps (Fig. 14b, c). We counted 809, 1074, and 2017 eggs in three clutches. The egg diameter ranged from 0.9‒2.4 mm (mean ± sd: 2.1 ± 0.42 mm mm; N= 100). We observed aquatic heteropterans (family Nepidae) preying on the eggs. It is likely that the tadpoles described by Lamotte (1957) from the Simandou range, can actually be assigned to A. fonensis. At Wataférédou, where A. fonensis was breeding, the forest stream was regularly polluted by people washing laundry. The edges of this gallery forest were degraded by subsistence farming, and cattle, and cleared for charcoal production and fuelwood., Published as part of Griesbaum, Frederic, Jongsma, Gregory F. M., Penner, Johannes, Kouamé, N'Goran Germain, Doumbia, Joseph, Gonwouo, Nono L., Hillers, Annika, Glos, Julian, Blackburn, David C. & Rödel, Mark-Oliver, 2023, The smallest of its kind: Description of a new cryptic Amnirana species (Amphibia, Anura, Ranidae) from West African rainforests, pp. 301-339 in Zootaxa 5254 (3) on pages 319-322, DOI: 10.11646/zootaxa.5254.3.1, http://zenodo.org/record/7727536, {"references":["Rodel, M. - O. & Bangoura, M. A. (2004) A conservation assessment of amphibians in the Foret Classee du Pic de Fon, Simandou Range, southeastern Republic of Guinea, with the description of a new Amnirana species (Amphibia Anura Ranidae). Tropical Zoology, 17, 201 - 232. https: // doi. org / 10.1080 / 03946975.2004.10531206","R ˆ del, M. - O. (2003) The amphibians of Mont Sangbe National Park, Ivory Coast. Salamandra, 39, 91 - 110.","Kanga, K. P., Kouame, N. G., Zogbasse, P., Gongomin, B. A. - I., Agoh, K. L., Kouame, A. M., Konan, J. C. B. Y. N., Adepo-Gourene, A. B., Gourene, G. & R ˆ del, M. - O. (2021) Amphibian diversity of a West African biodiversity hotspot: an assessment and commented checklist of the batrachofauna of the Ivorian part of the Nimba Mountains. Amphibian and Reptile Conservation, 15 (1) [General Section]: 71 - 107 (e 275).","R ˆ del, M. - O., Bangoura, M. A. & B ˆ hme, W. (2004) The amphibians of south-eastern Republic of Guinea (Amphibia: Gymnophiona, Anura). Herpetozoa, 17, 99 - 118.","Lamotte, M., Lauwarier, G. & Perret, J. - L. (1957) Contribution a l'etude des batraciens de l'Ouest Africain V. - Le developpement larvaires de Rana (Hylarana) albolabris. Bulletin de l'Institut fondamental d'Afrique Noire, Ser. A, 19, 1312 - 1327."]}

Heterixalus carbonei The description of the external morphology is based on a specimen in stage 31, field number 61HiwaB7.3.01, from Kirindy Forest (TL 22.56 mm, BL 7.63 mm; Table 1-2, Fig. 2). External morphology. In the dorsal view, the body has an ovoid form. Its widest point lies in the distal half to three quarters of the body. The snout is broadly rounded. In profile, the body is slightly depressed (BW 113% of BH). Its maximal height is in its distal half shortly behind mid-body. The snout is pointed. The eyes are moderately sized (ED 14% of BL) in a lateral position with dorsolateral orientation, and visible in the ventral view. Nares are located dorsolaterally, closer to the snout than to the eyes. They have a dark spot on their back and open both dorso- and anterolaterally in a round opening form. The distance between nostrils is about one quarter of the inter-orbital distance (NN 26% of IOD). The spiracle is sinistral and visible in the dorsal view. Its opening is located at maximum body width, shortly below the level of the proximal end of the tail myotomes axis. The spiracle opens posterodorsally, and its inner wall is free from the body. The vent tube is dextral, very short, and detached from the ventral fin. The coiled gut is visible. The tail is twice as long as the body (TAL 196% of BL). The tail muscle is narrow (TMW 32% of BW) and low (TMH 49% of BH). It rapidly converges (TMHM 22% of THM) until it reaches the pointed tail tip. Dorsal and ventral fins originate pointed at the body tail junction. The dorsal fin diverges to maximum height in the proximal quarter to half of the tail and starts converging towards its pointed tail tip in the distal quarter to half. The ventral fin reaches the comparably lower maximum height in the proximal quarter. Fin forms result in a tail that is at its maximum, which is in the proximal quarter to half of the tail (THM 67% of MTH), as high as the body (MTH 97% of BH). The oral disc type is generalized without emargination and of moderate size (ODW 41% of BW). Its position and orientation are anteroventrally. On the lower labium, there is a single row of very few marginal papillae. Ventrally and laterally, there are some submarginal papillae. The longish and rounded papillae form a moderately sized dorsal gap (DG 55% of ODW). The LTRF is 1/3(1), P3 is much shorter than P1 and P2. Labial teeth have three to four cusps. The upper tooth row has at least half the width of the oral disc (A1 51% of ODW). The same is true for the jaw sheaths (JL 59% of ODW). The jaws sheaths are half keratinized with pointed serrations on their edges. The upper jaw sheath has a broad, arched form, while the lower one is V-shaped. Coloration in preservative. Laterally, the tadpole is faintly pigmented in the anterior half and strongly pigmented in the posterior half. The coiled gut is visible. Dorsally, there is a broad strip of pigmentation that forks anteriorly. The fins and the lateral tail muscle show sparse pigmentation with a web of fine brown lines on the muscle and the dorsal fin. The limb buds are without pigmentation as well as the spiracle, which is transparent. The eyes are black. Ventrally, there is hardly any pigmentation, including the oral disc, which has dark brown pigments solely on the keratinized part of the jaws sheaths and the labial teeth. Variation. The external morphology and general appearance was very similar between specimens. The ratios taken on five tadpoles in stages 27 to 31 vary in the following proportions: BH 104-133% of BW; ED 12-14% of BL; NN 24-46% of IOD; TAL 168-196% of BL; TMW 22-31% of BW; TMH 31-49% of BH; TMHM 21-28% of THM; THM 71-89% of MTH; MTH 68-97% of BH; DF 121-196% of TMHM; VF 126-177% of TMHM; DF 95 116% of VF; ODW 31-41% of BW; DG 39-55% of ODW; A1 51-61% of ODW; JL 51-59% of ODW., Published as part of Griep, Stine & Glos, Julian, 2020, Description of tadpoles of the frogs Heterixalus tricolor, H. carbonei and H luteostriatus (Anura: Hyperoliidae) from western Madagascar, pp. 332-344 in Zootaxa 4767 (2) on pages 338-340, DOI: 10.11646/zootaxa.4767.2.8, http://zenodo.org/record/3771530

Heterixalus tricolor The description of external morphology is based on a specimen in stage 30, field number 56#25, from Kirindy Forest (TL 23.23 mm, BL 8.14 mm; Tables 1-2, Fig. 1). External morphology. In the dorsal view, the body form is ovoid. Its widest point lies in the distal half to three-quarters of the body. The snout is broadly rounded. In profile, the body is slightly depressed (BW 110% of BH). Its maximal height is in its distal half shortly behind mid-body. The snout is pointed. The eyes are moderately sized (ED 14% of BL). Their position is lateral with a dorsolateral orientation, visible in the ventral view. Nares are located dorsolaterally, nearer to the snout than to the eyes. They have a dark spot on their back and open both dorso- and anterolaterally. Their opening form is round. The distance between nostrils is about one third of the distance between orbitals (NN 35% of IOD). The spiracle is sinistral and visible in the dorsal view. Its opening is located at maximum body width, shortly below the level of the proximal end of the tail myotomes axis. Its inner wall is free from the body, formed in a way that the aperture opens more laterally than posterodorsally. The vent tube is dextral, very short, and not attached to the ventral fin. The coiled gut is visible. The tail is nearly twice as long as the body (TAL 185% of BL). The tail muscle is narrow (TMW 34% of BW) and low (TMH 41% of BH, TMHM 32% of THM). At the highest point, which lies in its proximal quarter to half, the tail is nearly as high as the body (MTH 87% of BH). From that point, the tail height continuously lowers towards the pointed tail tip (THM 77% of MTH), which is reached by the tail muscle. The dorsal fin originates proximal to the body tail junction. The fin is pointed in its origin and widens towards the point of maximum tail height. From that point, dorsal fin height remains mostly constant until it starts lowering in the distal half of the tail towards the fin���s pointed end at the tail tip. The ventral fin originates at the body tail junction. It reaches its maximum height, which is lower than the maximum height of the dorsal fin, in the proximal quarter of the tail. The height remains mostly constant until it starts lowering in the distal half of the tail towards the fin���s pointed tail tip. The oral disc is small (ODW32% of BW), and its type is generalized without emargination. Its position and orientation are anteroventral. Ventrally and laterally, on the lower labium, there is a single row of very few marginal papillae and some submarginal papillae. The papillae are rounded and form a moderately sized dorsal gap (DG 47% of ODW). On the upper labium, the labial teeth form one long row (A1 is 64% of ODW). The LTRF is 1/3(1), P3 is much shorter than P1 and P2. Labial teeth have three to four cusps. The jaws sheaths are wide (JL 54% of ODW) with pointed serrations at the edges. Jaws sheaths are half keratinized. The upper jaw sheath is arched, the lower one is V-shaped. Closed, the upper jaw sheath covers the lower one. Coloration in preservative. The body of the tadpole is pigmented dorsally and on the anterior part of the sides. The pigmentation is most prominent around the lower halves of the eyes. There is hardly any pigmentation at the abdomen and on the lower, distal sides of the body. The coiled gut is visible. Pigmentation is weak on the spiracle and nonexistent on the limb buds. The eyes are black. The dorsal fin and the tail muscle have only sparse pigmentation. The ventral fin is mostly without pigmentation. There is a dark brown line on parts of the tail myotomes axis. The oral disc has no pigments except for the dark brown keratinized parts of the jaws sheaths and the labial teeth. Variation. The external morphology and general appearance was very similar between specimens. The ratios taken on five tadpoles in stages 30 to 35 vary in the following proportions: BH 108-118% of BW; ED 14-15% of BL; NN 26-37% of IOD; TAL 176-188% of BL; TMW 25-34% of BW; TMH 37-42% of BH; TMHM 25-32% of THM; THM 77-90% of MTH; MTH 80-91% of BH; DF 87-162% of TMHM; VF 122-135% of TMHM; DF 67- 125% of VF; ODW 28-32% of BW; DG 45-57% of ODW; A1 62-69% of ODW; JL 52-60% of ODW., Published as part of Griep, Stine & Glos, Julian, 2020, Description of tadpoles of the frogs Heterixalus tricolor, H. carbonei and H luteostriatus (Anura: Hyperoliidae) from western Madagascar, pp. 332-344 in Zootaxa 4767 (2) on pages 337-338, DOI: 10.11646/zootaxa.4767.2.8, http://zenodo.org/record/3771530

Heterixalus luteostriatus The description of the external morphology is based on a specimen in stage 30, field number 2_Kirindy, from Kirindy Forest (TL 21.58 mm, BL 8.64 mm; Table 1-2, Fig. 3). External morphology. In the dorsal view, the body has an ovoid form with its widest point in the distal half to three-quarters of the body. The snout is broadly rounded. In profile, the body is depressed (BW 126% of BH), with its highest point in the distal half shortly behind mid-body. The snout is pointed. Moderately sized eyes (ED 13% of BL) are in a lateral position with a dorsolateral orientation. They are not visible in the ventral view. Nares are externally invisible. The spiracle is sinistral and visible in the dorsal view. It opens posterodorsally, at the broadest part of the body, slightly below the level of the proximal end of the tail myotomes axis. Its inner wall is free from the body. The vent tube is dextral, very short, and not attached to the ventral fin. The coiled gut is visible in an irregular shape. The tail reaches a length of about one and a half body lengths (TAL is 150% of BL). The tail muscle, which reaches the pointed tail tip, is narrow (TMW 36% of BW) and low both in the origin (TMH 46% of BH) and at mid-tail (TMHM 36% of THM). At its maximum height, which lies in the proximal quarter to half of the tail, the tail has a similar height as the body (MTH 98% of BH). More distal, at mid-tail, the tail height remains comparably high (THM 97% of MTH). Dorsal and ventral fins have their pointed origin at the body tail junction. Both fins reach their maximum height, which is lower in the ventral fin than in the dorsal fin, in the proximal quarter to half of the tail. The height remains mostly constant until both fins start converging in the distal half of the tail to a pointed tail tip. The oral disc is small and generalized without emargination (ODW 29% of BW). It has an anteroventral position and orientation. On the lower labium, there is a single row of very few marginal papillae. Ventrally and laterally, there are some submarginal papillae. The papillae are rounded and form a moderately sized dorsal gap (DG 60% of ODW). The LTRF is 1/3(1), P3 is much shorter than P1 and P2. The upper tooth row is about half as wide as the oral disc (A1 53% of ODW). Serrated labial teeth have two to three cusps. The wide jaws sheaths (JL 71% of ODW) have edges with pointed serrations. The upper jaw sheath has a broad, round form while the lower one is V-shaped. Both jaws sheaths are half keratinized. Coloration in preservative. Dorsally, the body of the tadpole shows uniform pigmentation. Laterally, the pigmentation pattern is fading. Pigmentation is particularly sparse in the area between the pigmented spiracle and the tail. There are parts of light brown pigmentation on the transparent abdomen, the coiled gut is visible. The limb buds are without pigmentation. The eyes are black. A pattern of uniformly pigmented patches covers the tail. Scattered spots of pigmentation are visible in the papillae of the oral disc. The teeth and the keratinized parts of the jaws sheaths are blackish-brown. Variation. The external morphology and general appearance were very similar between specimens. The ratios taken on five tadpoles in stages 25 to 34 vary in the following proportions: BH 116-127% of BW; ED 12-13% of BL; TAL 150-168% of BL; TMW 21-36% of BW; TMH 29-49% of BH; TMHM 31-36% of THM; THM 83-97% of MTH; MTH 82-98% of BH; DF 103-131% of TMHM; VF 67-95% of TMHM; DF 122-164% of VF; ODW 25-29% of BW; DG 44-68% of ODW; A1 50-69% of ODW; JL 54-71% of ODW. Morphometric differences between species When performing an NMDS, all five individuals within a species formed clear clusters that separated mainly along axis 1 (Fig. 4). The stress value (0.16) of the final two-dimensional solution indicated a reasonable preservation of ordering relationships of the multidimensional dissimilarities. Morphometry along this axis differed significantly between species (PerManova: TSS = 0.81, F = 6.69, p, Published as part of Griep, Stine & Glos, Julian, 2020, Description of tadpoles of the frogs Heterixalus tricolor, H. carbonei and H luteostriatus (Anura: Hyperoliidae) from western Madagascar, pp. 332-344 in Zootaxa 4767 (2) on pages 340-341, DOI: 10.11646/zootaxa.4767.2.8, http://zenodo.org/record/3771530

Genus-level taxonomy of the Cophylinae Mini adds a ninth genus to the Cophylinae for a unique clade of miniaturised frogs that falls sister to the large-bodied Plethodontohyla (Fig 1). Although body size is the most obvious character that differentiates these two sister genera, they are also distinguished by a number of osteological features, and can be identified without skeletal analysis by their digital reduction (present in Mini, absent in Plethodontohyla) and vomerine teeth (absent in Mini, present in Plethodontohyla). Despite these differences and consistent recovery of the two genera as being reciprocally monophyletic in genetic analyses, the uncorrected p-distances between these genera in the 3��� fragment of the 16S rRNA mitochondrial gene analysed here are at first glance surprisingly small at 8.3���13.3% (these distances would be distinctly higher if insertions and deletions would be considered in their calculation). Nevertheless, we consider the differences between these clades sufficiently great and robust that we regard them as constituting separate genera. Aside from their morphological and osteological differentiation, a further argument for their classification in distinct genera comes from the strength of support for their sistergroup relationship; while the two clades here seen as genera Plethodontohyla and Mini have been placed sister to each other in most molecular analyses so far, support values for this grouping often were low, and typically lower than the respective support for each of the two clades. The clade stability criterion [36] is therefore better served considering both clades as separate genera. The relationship of Mini to Plethodontohyla is analogous to the relationship of Stumpffia to Rhombophryne: a genus-level sister clade of miniaturised frogs (although Stumpffia also contains several non-miniaturised species), recovered in robust genetic phylogenies as reciprocally monophyletic, and distinguished by several diagnostic characters [14, 15, 23]. Peloso et al. [19] argued for the lumping of Stumpffia, Rhombophryne, and later also Anilany [22] into a single genus, Rhombophryne. In response, we showed that the initial argument for lumping was based on misidentified specimens [15], and subsequently incorrectly coded morphology, ignoring various unique diagnostic features of Anilany, and the relationships of particularly unstable taxa (most notably Stumpffia tridactyla) [23]. We revised the taxonomy of the genus Stumpffia, describing 26 new species, and providing a more robust phylogeny that resolved the phylogenetic position of Stumpffia tridactyla [14]. Despite this progress, the Amphibian Species of the World database (ASW) currently continues to use the lumped taxonomy, in contrast to AmphibiaWeb and other researchers that have adopted our proposed taxonomy (e.g. [20, 53]). The newly described Rhombophryne proportionalis, which is the only miniaturised Rhombophryne so far known, is highly distinct from Stumpffia, lacking, for example, the externally obvious digital reduction that is present in all miniaturised species of Stumpffia, and differing in body shape and proportions [14]. This demonstrates that even miniaturised Rhombophryne species can be distinguished by external morphology from Stumpffia species, providing still further support for the recognition of Rhombophryne, Stumpffia, and Anilany as separate genera., Published as part of Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., R��del, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel & Glaw, Frank, 2019, Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera, pp. 1-45 in PLoS ONE 213314 on pages 37-38, DOI: 10.1371/journal.pone.0213314, http://zenodo.org/record/2614083, {"references":["36. Vences M, Guayasamin JM, Miralles A, de la Riva I. To name or not to name: Criteria to promote economy of change in Linnaean classification schemes. Zootaxa. 2013; 3636 (2): 201 - 44. https: // doi. org / 10. 11646 / zootaxa. 363 6.2. 1","14. Rakotoarison A, Scherz MD, Glaw F, Kohler J, Andreone F, Franzen M, et al. Describing the smaller majority: Integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar. Vertebrate Zoology. 2017; 67 (3): 271 - 398.","15. Scherz MD, Vences M, Rakotoarison A, Andreone F, Kohler J, Glaw F, et al. Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae). Molecular Phylogenetics and Evolution. 2016; 100: 372 - 81. https: // doi. org / 10.1016 / j. ympev. 20 16.04. 0 19 PMID: 270856 71","23. Scherz MD, Vences M, Rakotoarison A, Andreone F, Kohler J, Glaw F, et al. Lumping or splitting in the Cophylinae (Anura: Microhylidae) and the need for a parsimony of taxonomic changes: a response to Peloso et al. (2017). Salamandra. 2017; 53 (3): 479 - 83.","19. Peloso PLV, Frost DR, Richards SJ, Rodrigues MT, Donnellan S, Matsui M, et al. The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in narrow-mouthed frogs (Anura, Microhylidae). Cladistics. 2016; 32 (2): 113 - 40. https: // doi. org / 10.1111 / cla. 12118","22. Peloso PLV, Raxworthy CJ, Wheeler WC, Frost DR. Nomenclatural stability does not justify recognition of paraphyletic taxa: A response to Scherz et al. (2016). Molecular Phylogenetics and Evolution. 2017; 111 (2017): 56 - 64. https: // doi. org / 10.1016 / j. ympev. 2017.03. 0 16 PMID: 28336422","20. Tu N, Yang M, Liang D, Zhang P. A large-scale phylogeny of Microhylidae inferred from a combined dataset of 121 genes and 427 taxa. Molecular Phylogenetics and Evolution. 2018; 126: 85 - 91. https: // doi. org / 10.1016 / j. ympev. 201 8.03. 0 36 PMID: 2964958 1","53. Feng Y-J, Blackburn DC, Liang D, Hillis DM, Wake DB, Cannatella DC, et al. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary. Proceedings of the National Academy of Sciences of the USA. 2017; 114 (29): E 5864 - E 70. https: // doi. org / 10.1073 / pnas. 17046 32114 PMID: 286739 70"]}

Anodonthyla eximia sp. nov. urn:lsid:zoobank.org:act: 2C419E74-C13D-447D-BDCF-312324515EAE (Figs 1, 2, 3, 5, 12 and 13, Tables 1 and 2) Remark. This species was previously listed as A. sp. Ranomafana (Maharira) [24], A. sp. Ca04 Ranomafana (ZCMV 204) [15] and Anodonthyla sp. 4 Ranomafana [20]. Holotype. ZMA 20246 (ZCMV 204, GenBank accession number GU177052 and FJ559111 for the 5��� and 3��� fragments of the 16S rRNA gene, respectively, and GU177063 for the cox1 gene), an adult male specimen (vocal sac inflated when collected) collected from a campsite at the base of Maharira mountain (21.3258&ring;S, 47.4025&ring;E, approx. 1248 m a.s.l.) in Ranomafana National Park, Vatovavy-Fitovinany Region, former Fianarantsoa province, southeastern Madagascar on 25 January 2004 by M. Vences, I. de la Riva, and E. Rajeriarison. Diagnosis. An extremely miniaturised frog assigned to the genus Anodonthyla on the basis of the possession of a large, cultriform prepollex, T-shaped terminal phalanges, absence of postchoanal vomer, and by its genetic affinities. It is separated by uncorrected p-distances of 9.3���17.0% in the analysed 3��� fragment of the 16S rRNA gene from all other members of the genus Anodonthyla. Anodonthyla eximia sp. nov. is characterised by the unique combination of the following characters (n = 1 male specimen): (1) male SVL 11.3 mm; (2) ED/HL 0.45; (3) HW/SVL 0.31; (4) FARL /SVL 0.30; (5) TIBL/SVL 0.39; (6) HIL/SVL 1.34; (7) finger 1 highly reduced, other fingers small; (8) toe 1 absent, toes 2 and 5 quite reduced; (9) maxillary and premaxillary teeth absent; (10) vomerine teeth absent; (11) lateral colour border absent; (12) black inguinal spots absent; (13) postchoanal vomer absent; (14) nasal subrectangular with an acuminate maxillary process, not displaced; (15) quadratojugal-maxilla contact absent; (16) zygomatic ramus of squamosal short, thin, and anterodorsally oriented; (17) clavicles present, curving with simple lateral articulations, medially bulbous; (18) prepollex cultriform, longer than first metatarsal; (19) carpal 2 present; (20) finger phalangeal formula 1-2-3-3; (21) toe phalangeal formula 2-2- 3-4-3; (22) single-note, unpulsed calls, not emitted in series; (23) weakly frequency modulated calls; (24) call dominant frequency 8406 �� 78 Hz (n = 5); (25) call duration 59.6 �� 6.5 ms (n = 5); (26) inter-call interval 3749.0 �� 1149.9 ms (n = 4). This species is considerably smaller than all other Anodonthyla species (11.3 mm vs 15���34 mm), and as such is only possible to confuse with juveniles of its congeners. In addition to its small size, it can be distinguished from all other Anodonthyla species by the absence of flared crests on the humerus in adult males. Among similarly sized adult frogs in Madagascar, it can be distinguished by the possession of a large inner metacarpal tubercle with a large, cultriform prepollex (present only in male Anodonthyla, much smaller in all other species except Anilany, where it is broad and triangular instead of cultriform), and a laterally displaced neopalatine not in contact with the sphenethmoid or vomer (neopalatine either in contact with sphenethmoid or in contact with vomer in all other species; in some species of Stumpffia, the neopalatine is lost, unpublished data). Holotype description. Specimen in a good state of preservation, a piece of the right thigh removed as a tissue sample. Body oblong; head wider than long, narrower than the body width; snout rounded in dorsal and lateral view; nostrils directed laterally, not protuberant, closer to eye than to tip of snout; canthus rostralis rounded, straight; loreal region concave, vertical; tympanum indistinct, round, about 46% of eye diameter; supratympanic fold weak, not raised, straight from posterior corner of eye to axilla; tongue long, thin, attached anteriorly, not notched; maxillary teeth absent, vomerine teeth absent; choanae oblong, very small. Forelimbs relatively broad; subarticular tubercles indistinct, single; outer metacarpal tubercle small, indistinct, single; inner metacarpal tubercle large and distinct, bulging outward strongly, underlain by cultriform prepollex; hand without webbing; all fingers short, first highly reduced and only marginally longer than the prepollex; relative length of fingers 1 < 2 = 4 < 3, fourth finger equal in length to second; tip of third finger marginally expanded, other fingers not expanded into discs. Hind limbs robust; TIBL 39% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle rounded, indistinguishable from first toe; outer metatarsal tubercle absent; no webbing between toes; first toe practically absent, second and fifth toes shortened; relative length of toes 2 < 5 < 3 < 4; fifth toe distinctly shorter than third. Skin on dorsum smooth, without distinct dorsolateral folds. Ventral skin smooth. After 13 years in 70% ethanol, the dorsum is pale brown with a faint darker brown chevron over the scapular region (Fig 2). The dorsal legs have faint dark brown crossbands, especially on the shanks. The dorsal colouration fades over the shanks to cream, which is continuous on the venter. The lateral head is dark brown, with a distinct border to the dorsum and flank formed by the supratympanic fold. The chin is a lighter brown than the dorsum but distinctly not the cream of the abdomen. The ventral legs are translucent cream. Colour in life as in preservative but more vibrant; the venter was slate grey with blue-cream flecks. Bioacoustics. Calls recorded from an unknown specimen by M. Vences at 08h45, on 26 January 2004 at Maharira in Ranomafana National Park (21.3258&ring;S, 047.4025&ring;E, 1248 m a.s.l.) in the leaf litter of primary rainforest, referred to this species (Fig 4D, Table 2). The holotype and single known specimen of this species was collected during careful searches of the leaf litter from the exact spot where similar calls were heard, and upon capture, had an apparent partially inflated vocal sac. No other calls assignable to a miniature frog were heard in the area at this time in the morning. Assignment of the calls to this species therefore is the most likely hypothesis but remains tentative. The call bears remarkable resemblance to that of Stumpffia miery (Table 2), which is also known from lower elevations in Ranomafana National Park, and detailed future field study will be required to confirm its assignment to Anodonthyla eximia sp. nov. Calls were emitted as part of a large chorus in the early hours of the morning following cyclonic rainfall after the retreat of major flooding of the area. Calls consisted of a single note and were emitted at regular intervals without defined call series. Calls were weakly frequency modulated with an increase in pitch, but recording quality is too poor for detailed analysis. For approximate call parameters, see Table 2. Osteology (Fig 13). Based on ZMA 20246 (figured). Cranium (Fig 13 D���13G). Shape and proportions. Skull short and rounded, longer than wide, widest at the bowing of the quadratojugal roughly in line with the anterior face of the prootic. Braincase proportionally broad, with an extremely short rostrum. Neurocranium. Moderately ossified, otic capsules partly ossified. Sphenethmoid unossified. Prootic in dorsal contact with lateral flange of frontoparietal, ventral contact with parasphenoid alae, not approaching contralateral. Septomaxilla miniscule, very tightly curled, not further discussed due to low ossification and insufficient resolution. Columella (stapes) well ossified, pars media plectra (stylus) long and slightly curved, posteriorly and dorsally oriented toward the broadened, somewhat posteriorly oriented pars interna plectra (baseplate). Nasal narrow, retaining the shape of larger cophylines: subrectangular with an elongated, acuminate maxillary process that does not closely approach the maxillary pars facialis; displaced laterally, broadly separated from contralateral. Frontoparietal with rounded anterior edge, laterally rather straight-edged, with short lateral flange covering prootic, posteriorly strongly connected to exoccipital, lacking any dorsal process, separated from contralateral by a narrow gap with a small rhomboid facet at the level of the prootics, which may represent a pineal foramen. Parasphenoid with narrow, rather straight-edged cultriform process and roughly equally broad perpendicular alae, considerably shorter than frontoparietals, in contact with exoccipitals posterodorsally, prootics dorsally along the edges of the alae, anteroventrally free; posteromedial process long but not participating in foramen magnum. Vomer lacking postchoanal portion (typical of Anodonthyla); prechoanal portion strongly curved, bearing a small lateral ramus. Neopalatine simple and very thin, laterally displaced, not contacting any other ossified elements. Maxillary arcade gracile, maxilla and premaxilla edentate, anterior extension of maxilla not exceeding lateral extent of premaxilla. Premaxilla with a narrow dorsal alary process rising laterally, pars palatina shallowly divided into a narrow palatine process and broad lateral process. Maxilla practically lacking a pars facialis and bearing a narrow pars palatina, its posterior tip acuminate and not contacting the quadratojugal, the lingual surface of the pars palatina contacting the anterior ramus of the pterygoid, which has taken over the articulation. Pterygoid with a short medial ramus, long and strongly curved anterior ramus, and broad posterior ramus, posteriorly calcified to the quadratojugal complex. Quadratojugal bowed laterally, rather short, broadly connected to the ventral ramus of the squamosal, bearing a small posteroventral knob, anteriorly dividing and with decreasing ossification, not connected to the maxilla; the articulation of the mandible is apparently somewhat fortified by the mineralisation of the posterior ramus of the pterygoid+squamosal+quadratojugal posteroventral knob. Squamosal with a slender, rather straight ventral ramus, thin, posterodorsally oriented otic ramus, and short, thin, anterodorsally oriented zygomatic ramus. Mandible slim and edentate, largely unremarkable, with a weakly raised coronoid process on the angulosplenial. Mentomeckelians separated from the dentary, with small, poorly ossified hooked ventrolateral projections. Posteromedial processes of hyoid proximally rounded without an obvious medial crista. Postcranial skeleton (Fig 13 A���13C, 13H and 13I). Eight procoelous presacrals, all much broader than long, lacking neural spines, with round posterior articular processes, presacral I with a complete neural arch, presacrals II���IV with thicker and longer transverse processes than V���VIII. Sacrum with flared diapophyses, the leading and trailing edges roughly equally curved, the articulation type IIB sensu Emerson [42]. Urostyle bicondylar, long, broadening gently posteriorly, with a somewhat flared head and a low dorsal ridge. Pectoral girdle without ossified prezonal or postzonal elements, with ossified clavicles. Clavicle thin and curved, with a simple lateral junction, equal in length to the coracoid, medially bulbous. Coracoid broad, flared laterally and strongly flared medially with a curving medial articular surface with the contralateral. Scapula robust, with a broad pars acromialis, the cleithral border straight. Cleithrum ossified for three-quarters the width of the scapular border, thickened anteriorly. Suprascapula unossified. Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna slender with a distinct sulcus intermedius. Carpals composed of radiale, ulnare, element Y, prepollex, carpal 2, and a large post-axial element formed by carpals 3���5. Prepollex extremely long and acuminate, longer than first metacarpal. Finger phalangeal formula is reduced (1-2- 3-3), and the terminal phalanx of the first finger small and columnar, others bearing T-shaped knobs. Pubis ossified; iliac shafts passing ventral to and beyond sacrum, oblong in cross-section, with a weak dorsal crest, a distinct dorsal prominence, and a shallow oblique groove. Femur weakly sigmoid, bearing a distinct posterior crest. Tibiofibula slightly shorter than femur in length, with a sulcus intermedius. Tibiale and fibulare fused proximally and distally. T1 and T2 +3 tarsals present, T1 considerably smaller than T2+3. Centrale present, larger than other tarsals. Prehallux elongated, half the length of first metatarsal. Phalangeal formula standard (2-2- 3-4-3). Terminal phalanx of toe 1 a small round element, those of toes 2���5 with T-shaped knobs. Etymology. The species epithet eximia is the feminine form of the Latin adjective eximius meaning ���remarkable��� or ���special���, in reference to the surprisingly small body size and terrestrial habits of this Anodonthyla species. Distribution, natural history, and conservation status. Anodonthyla eximia sp. nov. is known only from Maharira in Ranomafana National Park (Fig 6). It is a terrestrial species. Nothing else is known of its natural history. It is likely that this species should be classified as Vulnerable like other species from Maharira (e.g. Gephyromantis runewsweeki), but as we know almost nothing of its range and ecology, we instead recommend that it be considered Data Deficient until more data are available., Published as part of Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., R��del, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel & Glaw, Frank, 2019, Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera, pp. 1-45 in PLoS ONE 213314 on pages 33-37, DOI: 10.1371/journal.pone.0213314, http://zenodo.org/record/2614083, {"references":["24. Vences M, Glaw F, Kohler J, Wollenberg KC. Molecular phylogeny, morphology and bioacoustics reveal five additional species of arboreal microhylid frogs of the genus Anodonthyla from Madagascar. Contributions to Zoology. 2010; 79 (1): 1 - 32.","15. Scherz MD, Vences M, Rakotoarison A, Andreone F, Kohler J, Glaw F, et al. Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae). Molecular Phylogenetics and Evolution. 2016; 100: 372 - 81. https: // doi. org / 10.1016 / j. ympev. 20 16.04. 0 19 PMID: 270856 71","20. Tu N, Yang M, Liang D, Zhang P. A large-scale phylogeny of Microhylidae inferred from a combined dataset of 121 genes and 427 taxa. Molecular Phylogenetics and Evolution. 2018; 126: 85 - 91. https: // doi. org / 10.1016 / j. ympev. 201 8.03. 0 36 PMID: 2964958 1","42. Emerson SB. The ilio-sacral articulation in frogs: form and function. Biological Journal of the Linnean Society. 1979; 11: 153 - 68. https: // doi. org / 10.1111 / j. 1095 - 8312.1979. tb 00032. x"]}

Mini gen. nov. urn:lsid:zoobank.org:act: 67171236-00A2-428B-8194-584BF52E84E6 (Figs 2–9, Table 1) Type species. Mini mum sp. nov. Contents. Mini mum sp. nov., M. scule sp. nov., and M. ature sp. nov. Etymology. The genus name is derived from English prefix ‘mini-’, denoting a small version of an object. We treat this name as an arbitrary combination of letters in the sense of the International Code of Zoological Nomenclature Articles 30.1.4.1 and 30.2.2, and we assign it the feminine gender. We have searched all available taxonomic databases and could not find any evidence that this name has ever been used to refer to a genus of animals, and we therefore conclude that it is available. Diagnosis. Diminutive terrestrial frogs (adult SVL 8.2–14.9 mm), assigned to the Madagascar-endemic subfamily Cophylinae on the basis of divided vomers, procoelous vertebral column, divided sphenethmoids, and genetic affinities. Skin smooth to slightly granular, occasionally iridescent. A lateral colour border is present but varies in intensity among species. Highly reduced fingers and toes, fusion or loss of carpal 2, and paedomorphic skull morphology: laterally displaced narrow nasals, teeth absent from vomer, in some species present on the maxilla and premaxilla, otic capsule sometimes dorsally ossified, brain case comprising most of the skull’s length and width. All members of the genus Mini gen. nov. resemble miniaturised to extremely miniaturised members of the genus Stumpffia. However, all species can be distinguished from Stumpffia on the basis of their curving clavicles and a fused or lost carpal 2. In the species accounts below, we provide detailed distinctions from Stumpffia relevant to each species. Justification. The erection of the genus Mini is justified by significant genetic differentiation from all other major cophyline lineages (see Fig 1), by the fact that it does not form a monophyletic group with the genus Stumpffia, and furthermore by the strong morphological differences (including but not restricted to the much smaller size) to all species of its sister clade, Plethodontohyla. The following characters distinguish the genus from all Plethodontohyla species, including juveniles: digital reduction of the fingers and toes (vs no reduction), laterally displaced and reduced nasals (vs large nasals situated anterior to frontal), parasphenoid cultriform process shorter than frontoparietals (vs roughly equal in length to the frontoparietals) and considerably narrower than alary processes (vs as wide or wider), vomerine teeth absent (vs present), carpal 2 absent (vs present). Uncorrected p-distances between Mini and Plethodontohyla range from 8.3–13.3% in the 3’ fragment of 16S rRNA analysed here, and they have been found to be sister to Plethodontohyla in all phylogenetic analysis since their first inclusion in genetic datasets [15, 17, 20, 22] (except the DNA barcoding study of Vieites et al. [40], where they were placed at the base of Rhombophryne + Stumpffia + Anilany, but that study lacks any resolution at deep nodes, and was not intended to provide phylogenetic hypotheses at deep levels). Distribution. The genus Mini is apparently endemic to low-elevation habitats (0–350 m a. s. l.) of southeastern Madagascar (Fig 3).

Rhombophryne proportionalis sp. nov. urn:lsid:zoobank.org:act: 1 A79607 B-E9D4-4214-BF47-9BC0B4320E5F (Figs 1, 2, 3, 5, 10 and 11, Tables 1 and 2) Remark. This species was previously referred to as Stumpffia sp. Ca34 [15, 46] and as Stumpffia sp. 39 MV-2012 (KC 351481) [20]. Holotype. ZSM 1826 /2010 (ZCMV 12404, GenBank accession number KC351380 and KU 937808 for 5��� and 3��� fragments of the 16S rRNA gene, respectively, and KF611640 for the cox1 gene), an adult male specimen (seen calling, not recorded) collected at Bepia Campsite on the Tsaratanana massif (Camp 3; 14.1182&ring;S, 48.9782&ring;E, 2294 m a.s.l.), Diana Region, former Antsiranana province, northern Madagascar on 16 June 2010 by M. Vences, D.R. Vieites, R.D. Randrianiaina, S. Rasamison, and E. Rajeriarison. Paratypes. ZSM 1840 /2010 (ZCMV 12405, GenBank accession number KC351481 and MK 459317 for 5��� and 3��� fragments of the 16S rRNA gene, respectively), adult male specimen with the same collection data as the holotype; and ZSM 636 /2014 (DRV 6224), an adult presumed male specimen collected at Andranomadio Campsite in Tsaratanana (Camp 4; 14.0801&ring;S, 48.9854&ring;E, 2503 m a.s.l.) on 16 June 2010 by the same collectors. Diagnosis. A diminutive frog assigned to the genus Rhombophryne on the basis of absence of clavicles, presence of vomerine, maxillary, and premaxillary teeth, short and broad skull, and genetic affinities. It is separated by uncorrected p-distances of 7.0���12.9% in the analysed 3��� fragment of the 16S rRNA gene from other members of the genus Rhombophryne. Rhombophryne proportionalis sp. nov. is characterised by the unique combination of the following characters (n = 3 male specimens): (1) male SVL 11.0��� 12.3 mm; (2) ED/HL 0.40���0.48; (3) HW/SVL 0.33���0.37; (4) FARL /SVL 0.33���0.35; (5) TIBL/SVL 0.34���0.36; (6) HIL/SVL 0.21��� 1.32; (7) finger 1 reduced, 2 and 4 short; (8) toe 1 highly reduced, 2 somewhat reduced; (9) maxillary and premaxillary teeth present; (10) vomerine teeth present; (11) lateral colour bor- der absent; (12) black inguinal spots sometimes present; (13) postchoanal vomer present, spatulate, medially fused to parasphenoid; (14) nasal broad and not laterally displaced; (15) quadratojugal-maxilla contact strong and broad; (16) zygomatic ramus of squamosal long, thick, curved, and horizontal; (17) clavicles absent; (18) prepollex short and triangular; (19) carpal 2 present; (20) finger phalangeal formula 2-2-3-3; (21) toe phalangeal formula 2-2-3-4- 3; (22) unpulsed calls emitted in series of 9���17 calls at irregular intervals; (23) non-frequency modulated calls; (24) call dominant frequency 5460 �� 117 Hz (n = 79); (25) call duration 45.4 �� 8.2 ms (n = 79); (26) inter-call interval 63.0 �� 9.0 ms (n = 73). Among extremely miniaturized cophylines, this species is unique in possessing vomerine teeth. It can be distinguished from almost all other miniaturised species in lacking clavicles (also absent in S. contumelia, S. obscoena, S. davidattenboroughi, S. makira, S. achillei, and S. analanjirofo, and some specimens of S. tridactyla, unpublished data). It is also characterised by a dark colouration of the lateral surface of the head with a distinct colour border, and less reduced fingers and toes. Confusion with juvenile Rhombophryne species is still possible, but these have much larger teeth proportional to their skull size, and most lack the distinct lateral head colouration and possess clavicles. The call is unique among the frogs of Madagascar and is instantly distinctive in being emitted as a rapid, high-pitched series of tonal notes. Holotype description. Specimen in a good state of preservation, part of the right thigh removed as a tissue sample. Body somewhat rhomboid; head wider than long, narrower than body; snout rounded in dorsal view, squared in lateral view; nostrils directed laterally, not protuberant, closer to eye than to tip of snout; canthus rostralis rounded, concave; loreal region flat, vertical; tympanum indistinct, round, about 57% of eye diameter; supratympanic fold distinct, weakly raised, curving slightly from posterior corner of eye over tympanum toward axilla; tongue very broad, disc-like, posteriorly free, unlobed; maxillary teeth present; vomerine teeth present in two tiny patches either side of the midline; choanae rounded. Forelimbs slen- der; subarticular tubercles faint, single; outer metacarpal tubercle faint, paired; inner metacarpal tubercle distinct, elongated; hand without webbing; first finger reduced, second and fourth short; relative length of fingers 1 < 2 = 4 < 3; finger tips not expanded. Hind limbs robust; TIBL 34% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle thin and indistinct; outer metatarsal tubercle small and indistinct; no webbing between toes; first toe highly reduced, second toe short; relative length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter than third. Skin on dorsum smooth in preservative, without distinct dorsolateral folds. In life, the dorsal skin was smooth was scattered tubercles, and distinct ridges above the scapular region (Fig 10). Ventral skin smooth in preservative, granular in life. After eight years in 70% ethanol, the dorsum is chocolate brown in colour, darker over the head, with a faint dark brown line running from the inguinal region anteriorly toward the eye. The dorsal leg has a dark brown crossband on the shank. The lateral head has a distinct colour border to the dorsum, being a much darker brown, its border defined by the supratympanic fold. The flank has an indistinct colour border to the venter. The venter is brown with cream flecks, slightly darker and less flecked on the chin. The ventral legs and arms are as the ventral abdomen in colour. The bottom of the foot is dark brown along the medial half. Colour in life as in preservative but more vibrant. Variation. For measurements, see Table 1. The paratypes are in general very similar to the holotype in morphology. ZSM 636/2014 is slightly smaller than the other specimens, and has a slightly shorter, more rounded head. Its supratympanic fold is also less curved than those of the other specimens, being rather more straight from the eye to above the arm. ZSM 1840/ 2010 has a more massive body than the others. The colouration of the specimens is relatively consistent, with the whole venter of ZSM 1840/2010 being darker than those of the other two specimens. The dorsolateral lines of the holotype are present in ZSM 1840/2010, but not in ZSM 636/2014, instead being broken in that specimen into spots above the suprascapular region and lines in the inguinal region. The crossbands of the shanks are less distinct in ZSM 1840/2010 than the other two specimens. Bioacoustics. Specimens called only during the day, from open, shrubby landscape between dense vegetation, on the ground. In some areas, numerous specimens could be heard calling in a chorus. Calls were recorded from an uncollected specimen by M. Vences (Fig 5C, Table 2) at around 11h40 on 15 June 2010 in Camp Bepia (14.11822&ring;S, 048.97822&ring;E, 2294 m a.s.l.), and further calls were heard but not recorded at Camp Andranomadio (14.0801&ring;S, 048.9854&ring;E, 2503 m a.s.l.). A precise description of the call structure is difficult; the vocalization is a series of short tonal units that cannot readily be assigned to units. In past studies, we have described roughly comparable structures differently: in Stumpffia psologlossa, where the units have a very short duration, as a single call composed of pulses [14], in Rhombophryne mangabensis as a series of notes [52], and in R. minuta two closely spaced units were seen as components of a single note [52]. To allow comparison within Rhombophryne we here define the tonal units in the vocalizations of the new species as notes, and the entire series as a call, but emphasize that it also would be possible to define each unit as call and the entire series as call series, or to consider each unit as a pulse as their duration falls within the 5���50 ms range for which the pulse category was recommended by K��hler et al. [33]. Calls are rapid series of high-pitched slightly frequency modulated notes. Each call consists of a series of 9���17 notes, and calls are repeated at long and irregular intervals. For detailed call parameters, see Table 2. Osteology (Fig 11). Based on ZSM 1826/2010 (figured) and ZSM 636/2014 (not figured). Cranium (Fig 11 C���11F). Shape and proportions. Skull short and stout, relatively wide, widest at quadratojugal anterior to the otic region. Short rostrum. Braincase broad. Neurocranium. Well ossified except the weakly calcified otic capsules. Sphenethmoid well ossified, in contact with the frontoparietal in ZSM 1826/2010 but not in ZSM 636/2014, with a calcified anterior medial cone; broadly separated from prootic. Prootic in dorsal contact with lateral flange of frontoparietal, ventral contact with parasphenoid alae, not approaching contralateral. Septomaxilla miniscule, roughly spiralled, similar in shape to those of other Rhombophryne (see [25]). Columella (stapes) well ossified, pars media plectra (stylus) steeply sloping upwards proximally to its broad, bilobed, flattened pars interna plectra (baseplate). Nasal broad, curved over the nasal capsule, triangular, acuminate maxillary process not closely approaching maxillary pars facialis, broadly separated from contralateral, situated anterior to frontoparietal. Frontoparietal with slanted anterior edge, laterally bulging, with short lateral flange covering prootic, posteriorly weakly connected to exoccipital, anteroventrally contacting sphenethmoid, lacking any dorsal process (suggestions of such processes in ZSM 1826/ 2010). Parasphenoid with broad, squared cultriform process and broad perpendicular alae, shorter than frontoparietals, posteromedial process not participating in foramen magnum, anteroventrally in contact with postchoanal vomer and not in contact with neopalatine. Vomer divided into pre- and postchoanal portions; prechoanal portion narrow, arcuate, triradiate, with a short lateral and anterior ramus and long posterior ramus; postchoanal portion spatulate bearing a single vomerine tooth or a pair thereof, separated from contralateral by a narrow space, in dorsal contact with the parasphenoid proximally and the neopalatine distally, lacking an anterior projection. Neopalatine laminar, broader than lateral postchoanal portion of vomer, laterally approaching but not contacting the maxilla, exceeding the lateral-most point of the postchoanal vomer to approach the anterior tip of the parasphenoid but not in contact with it, such that the vomer contacts the neopalatine near its midpoint and not at its terminus. Maxillary arcade quite robust, maxilla and premaxilla bearing numerous diminutive teeth, premaxilla and maxilla in narrow contact anteriorly. Premaxilla with a broad acuminate dorsal alary process rising vertically, pars palatina shallowly divided into a narrow palatine process and broad lateral process. Maxilla with a low triangular pars facialis and a narrow pars palatina, its posterior tip acuminate and broadly overlapping the quadratojugal, the lingual surface of the pars palatina in broad contact with the anterior ramus of the pterygoid. Pterygoid with a short medial ramus, long anterior ramus with broad contact with the maxilla, leaving a channel for the pterygoid cartilage, posterior ramus broad and posterolaterally calcified to the quadratojugal complex. Quadratojugal bowed laterally, broadly connected to the ventral ramus of the squamosal, and with a broad articular surface with the maxilla, bearing a large posteroventral knob. Squamosal with a broadened ventral ramus and narrow otic and zygomatic rami, the otic ramus long and thin and oriented dorsally, the zygomatic ramus short and thin, oriented anteriorly. Mandible slim and edentate, largely unremarkable, with a low coronoid process on the angulosplenial. Mentomeckelians strongly connected to the dentary, with unusual, flat ventrolateral projections. Posteromedial processes of hyoid proximally pointed with a broad medial crista. Postcranial skeleton (Fig 11A, 11B, 11G and 11H). Eight procoelous unfused presacrals, much broader than long, lacking neural spines, with round posterior articular processes, presacral I with a mostly complete neural arch, presacrals II���IV with thicker and longer transverse processes than V���VIII. Sacrum with expanded diapophyses, the leading and trailing edges roughly equally angled, the articulation type IIB sensu Emerson [42]. Urostyle bicondylar, long, broadening posteriorly, without lateral processes and with a low dorsal ridge. Pectoral girdle without ossified prezonal or postzonal elements, lacking ossified clavicles. Coracoid broad, strongly flared with a large medial articular surface with the contralateral. Scapula also robust, with a broad pars acromialis distinct from the pars glenoidalis. Cleithrum ossified for half the width of the scapular border, acuminate, thickened anteriorly. Suprascapula unossified. Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna broad with a distinct sulcus intermedius. Carpals well ossified in ZSM 1826/2010 and poorly ossified in ZSM 636/2014, composed of radiale, ulnare, prepollical element, element Y, carpal 2, and large post-axial element formed by carpals 3���5. Finger phalangeal formula is standard (2-2-3-3). Small distal knobs on terminal phalanges of the fingers. A very small prepollex is present in ZSM 1826/2010 but is not visible in ZSM 636/2014. Pubis partly calcified, iliac shafts passing ventral to and beyond sacrum, nearly cylindrical, without a dorsal crest and with a weak dorsal prominence and shallow oblique groove. Femur weakly sigmoid with a low posterior crest. Tibiofibula shorter than femur, with a sulcus intermedius. Tibiale and fibulare weakly fused proximally and distally. T1 and T2+3 tarsals present, T1 considerably smaller than T2+3. Centrale present but not large. Prehallux unossified. Phalangeal formula standard (2-2-3-4-3). Distribution, natural history, and conservation status. Rhombophryne proportionalis sp. nov. is known from two localities (Bepia and Andranomadio) on the Tsaratanana massif in northern Madagascar. Both of these sites fall within the Tsaratanana National Park (formerly a Strict Nature Reserve). It is a terrestrial species that lives among the leaf litter. Two other cophyline frogs, Platypelis alticola and P. tsaratananaensis, are also known from these locations, and both are currently listed as Endangered, due to their small distribution < 2000 km 2, presence in a single threat-defined location, and potentially on-going decline in habitat quality. We therefore suggest R. proportionalis sp. nov. also to be Endangered, following the same rationale. Etymology. The species epithet is the Latin adjective proportionalis meaning ���proportional���, in reference to the comparatively proportional dwarfism that this species has apparently undergone (see discussion). It is a feminine adjective in the nominative singular., Published as part of Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., R��del, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel & Glaw, Frank, 2019, Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera, pp. 1-45 in PLoS ONE 213314 on pages 28-33, DOI: 10.1371/journal.pone.0213314, http://zenodo.org/record/2614083, {"references":["15. Scherz MD, Vences M, Rakotoarison A, Andreone F, Kohler J, Glaw F, et al. Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae). Molecular Phylogenetics and Evolution. 2016; 100: 372 - 81. https: // doi. org / 10.1016 / j. ympev. 20 16.04. 0 19 PMID: 270856 71","46. Perl RGB, Nagy ZT, Sonet G, Glaw F, Wollenberg KC, Vences M. DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia. 2014; 35: 197 - 206. https: // doi. org / 10.1163 / 15685381 - 0 0 0 0 2942","20. Tu N, Yang M, Liang D, Zhang P. A large-scale phylogeny of Microhylidae inferred from a combined dataset of 121 genes and 427 taxa. Molecular Phylogenetics and Evolution. 2018; 126: 85 - 91. https: // doi. org / 10.1016 / j. ympev. 201 8.03. 0 36 PMID: 2964958 1","14. Rakotoarison A, Scherz MD, Glaw F, Kohler J, Andreone F, Franzen M, et al. Describing the smaller majority: Integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar. Vertebrate Zoology. 2017; 67 (3): 271 - 398.","52. Glaw F, Kohler J, Vences M. A new fossorial frog, genus Rhombophryne, from Nosy Mangabe Special Reserve, Madagascar. Zoosystematics and Evolution. 2010; 86 (2): 235 - 43. https: // doi. org / 10.1002 / zoos. 20100 0 0 0 6","33. Kohler J, Jansen M, Rodriguez A, Kok PJR, Toledo LF, Emmrich M, et al. The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa. 2017; 4251 (1): 1 - 124. https: // doi. org / 10.11646 / zootaxa. 425 1.1. 1 PMID: 28609991","25. Scherz MD, Hawlitschek O, Andreone F, Rakotoarison A, Vences M, Glaw F. A review of the taxonomy and osteology of the Rhombophryne serratopalpebrosa species group (Anura: Microhylidae) from Madagascar, with comments on the value of volume rendering of micro-CT data to taxonomists. Zootaxa. 2017; 4273 (3): 301 - 40. https: // doi. org / 10.11646 / zootaxa. 427 3.3. 1 PMID: 28610237","42. Emerson SB. The ilio-sacral articulation in frogs: form and function. Biological Journal of the Linnean Society. 1979; 11: 153 - 68. https: // doi. org / 10.1111 / j. 1095 - 8312.1979. tb 00032. x"]}

Mini ature sp. nov. urn:lsid:zoobank.org:act: 0C1C4CE2-419A-4030-86D1-B1B89793697D (Figs 1–3 and 9, Table 1) Remark. This species was previously listed as Stumpffia sp. Ca53 MV 2017(MF 867231) by Tu et al. [20], though with incorrect accession number (correct number is MF 768231). Holotype (Figs 2 and 9). ZSM 86 /2004 (FGZC 0 151, GenBank accession numbers MF768231 and MK 459307 for 5’ and 3’ fragments of the 16S rRNA gene, respectively, and MF768147 for cox1 gene), a presumed adult specimen collected in Andohahela National Park between Isaka and Eminiminy above ‘ Camp 1’ (ca. 24.75&ring;S, 46.85&ring;E, ca. 350 m) between 29 and 31 January 2004 by F. Glaw, M. Puente, M. Teschke (née Thomas), and R.D. Randrianiaina. Diagnosis. A highly miniaturised frog assigned to Mini gen. nov. on the basis of its small size, curved clavicles, laterally displaced and reduced nasals, and fusion or loss of carpal 2. This assignment is supported by its genetic affinities (Fig 1; [20]). It is separated by uncorrected pdistances of 10.0–10.6% in the analysed 3’ fragment of the 16S rRNA gene from other members of the genus Mini gen. nov., and 10.8–13.7% from all members of the genus Plethodontohyla. Mini ature sp. nov. is characterised by the unique combination of the following characters (n = 1 specimen): (1) SVL 14.9 mm; (2) ED/HL 0.38; (3) HW/SVL 37.2; (4) FARL /SVL 0.32; (5) TIBL/SVL 0.34; (6) HIL/SVL 1.18; (7) finger 1 strongly reduced, 2 and 4 reduced; (8) toe 1 absent, toes 2 and 5 quite reduced; (9) maxillary and premaxillary teeth present; (10) vomerine teeth absent; (11) lateral colour border absent; (12) black inguinal spots present; (13) postchoanal vomer present, spatulate, medially fused to parasphenoid; (14) nasal cultriform and laterally displaced; (15) quadratojugal-maxilla contact strong and broad; (16) zygomatic ramus of squamosal long, thin, curved, and horizontal; (17) clavicles present, curving with simple lateral articulations, medially not bulbous; (18) prepollex thin and cultriform; (19) carpal 2 absent or fused to post-axial carpal 3–5 element; (20) finger phalangeal formula 1-2-3-3; (21) toe phalangeal formula 1-2-3-4-3; (22–26) calls unknown. Within the genus Mini gen. nov., the new species can be distinguished by its distinctly larger body size (14.9 mm vs 8.2–11.3 mm), shorter relative hindlimb length (HIL /SVL 1.18 vs 1.41–1.72) and phalangeal formula of fingers (1-2-3-3 vs 1-2-3- 2 in M. mum sp. nov. and 0-2- 3- 2 in M. scule sp. nov.). Additionally, it can be distinguished from M. mum sp. nov. by the presence of maxillary and premaxillary teeth (vs absence), and less distinct lateral colour border, and M. scule sp. nov. by proportionally smaller nasals and braincase, broader quadratojugal-maxillary contact, and vertical dorsal process of premaxilla (vs anterior). This species is particularly similar to some highly miniaturised Stumpffia species, but it can be distinguished from all Stumpffia based on the condition of the carpals and the presence of curved clavicles, and most Stumpffia by the presence of maxillary and premaxillary teeth. Holotype description. Specimen in a moderately good state of preservation, the left arm removed as a tissue sample, the whole body somewhat dorsoventrally flattened in preservative. Body oblong; head wider than long, narrower than body width; snout slightly pointed in dorsal view, pointed in lateral view; nostrils directed laterally, not protuberant, equidistant between tip of snout and eye; canthus rostralis rounded, indistinct, slightly concave; loreal region flat, vertical; tympanum indistinct, round, ~48% of eye diameter; supratympanic fold absent; tongue long, broadening posteriorly, attached anteriorly, not notched; maxillary teeth present; vomerine teeth present; choanae small and round. Forelimb slender; subarticular tubercles single, elongated; outer/palmar metacarpal tubercle small and round; inner metacarpal tubercle slightly smaller than outer/palmar; hand without webbing; first finger strongly reduced, second and fourth fingers reduced, third finger basally broadened; relative lengths of fingers 1 < 2 = 4 < 3, fourth and second finger equal in length; finger tips not expanded into discs. Hindlimbs stocky; TIBL 34% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle indistinguishable from first toe; outer metatarsal tubercle absent; no webbing between toes, second and fifth toes reduced; relative lengths of toes 2 < 5 < 3 < 4, fifth toe distinctly shorter than third; toe tips slightly pointed distally. Skin on dorsum smooth without a distinct dorsolateral fold, but with a distinct colour border, see below. Ventral skin smooth. After 14 years in 70% ethanol, the dorsum is light brown, paler—almost beige—laterally, and slightly translucent, with a thin beige vertebral stripe and a darkened area on the posterior head. The skin above the eyes is translucent and dark in colour through the presence of the eyes beneath. The dorsal forelimb is beige flecked with brown, the hand is lighter medially, and the fingers have faint cream annuli. The dorsal hindlimb is beige in base colour with several brown crossbands on the thigh and shank that line up when the leg is folded together. A trapezoid of brown is present around the vent. The foot is dorsally as the forelimb with a light annulus before each distal phalanx. A distinct colour border that is not straight is present laterally, running along the canthus rostralis from the nostril through the eye, through the supratympanic region along the torso to the inguinal region. Small oblong dark brown spots are present in the inguinal region. The side of the head is dark brown. Ventral to this colour bor- der the frog is mocha speckled with beige, lightening ventrally to beige with loose cream speckles. The ventral skin is translucent, and some of the organs can be seen through it. The chin is not differently coloured than the rest of the ventral body. This pattern continues onto the ventral limbs. No data on life colouration are available. Variation. This species is currently known from a single specimen only. Osteology (Fig 9). Based on ZSM 86/2004 (figured). Cranium (Fig 9 D–9G). Shape and proportions. Skull almost equilateral, roughly as wide as long, widest at quadratojugal-squamosal junction, lateral to the otic region. Braincase moderately broad, rostrum not shortened. Neurocranium. Well ossified, including the otic capsules. The anterior cone of the sphenethmoid is ossified and contacts the frontoparietal dorsally but is not in contact with any other bones; small isolated lateral mineralisations of this bone are also present anterodorsal to the anterior tip of the cultriform process of the parasphenoid. Prootic in dorsal contact with lateral flange of frontoparietal, ventral contact with parasphenoid alae, not approaching contralateral ventrally. Septomaxilla miniscule, relatively poorly mineralised, and therefore not further discussed here. Columella (stapes) well ossified, pars media plectra (stylus) long and nearly straight, weakly posteriorly and dorsally oriented toward the reniform, dorsally elongated pars interna plectra (baseplate). Nasal narrow and cultriform, laterally displaced, curved downward laterally, acuminate maxillary process not closely approaching maxillary pars facialis, broadly separated from contralateral. Frontoparietal with rounded anterior edge, laterally bulging, with short lateral flange covering prootic, posteriorly strongly connected to exoccipital, anteroventrally contacting sphenethmoid, lacking any dorsal process. Parasphenoid with narrow, rather straight-edged cultriform process and broad perpendicular alae, considerably shorter than frontoparietals, in contact with exoccipitals posterodorsally, prootics dorsally along the edges of the alae, anteroventrally in contact with postchoanal vomer and not in contact with neopalatine; posteromedial process excluded from participating in foramen magnum by exoccipitals. Vomer divided into pre- and postchoanal portions; prechoanal portion narrow, arcuate, triradiate, with a short lateral and anterior ramus and long, curving posterior ramus; postchoanal portion spatulate and edentate, contacting its contralateral on the midline, in dorsal contact with the parasphenoid proximally and the neopalatine distally, lacking an anterior projection. Neopalatine simple, straight, almost indistinguishable from lateral postchoanal portion of vomer, laterally broadly separated from the maxilla, not exceeding the lateral-most point of the postchoanal vomer. Maxillary arcade fairly slight, maxilla and premaxilla bearing numerous diminutive teeth, anterior extension of maxilla exceeding lateral extent of premaxilla but not in contact with it. Premaxilla with a broad acuminate dorsal alary process rising laterally, pars palatina shallowly divided into a narrow palatine process and broad lateral process. Maxilla with a low triangular pars facialis and a narrow pars palatina, its posterior tip acuminate and broadly overlapping the quadratojugal, the lingual surface of the pars palatina running parallel to but not touching the anterior ramus of the pterygoid. Pterygoid with a short medial ramus, long anterior ramus, and broad posterior ramus and posterolaterally calcified to the quadratojugal complex. Quadratojugal weakly bowed laterally, broadly connected to the ventral ramus of the squamosal, bearing a small posteroventral knob; the articulation of the mandible is apparently fortified by the mineralisation of the posterior ramus of the pterygoid+squamosal+quadratojugal posteroventral knob. Squamosal with a slender ventral ramus, broadened otic ramus, and long, thin zygomatic ramus, the otic ramus oriented slightly dorsally and posteriorly, the zygomatic ramus horizontal and strongly curved. Mandible slim and edentate, largely unremarkable, with a low coronoid process on the angulosplenial. Mentomeckelians separated from the dentary, with slightly bulbous, almost hooked ventrolateral projections. Posteromedial processes of hyoid proximally rounded with a broad medial crista. Postcranial skeleton (Fig 9 A–9C, 9H and 9I). Eight procoelous unfused presacrals, much broader than long, lacking neural spines, with round posterior articular processes, presacral I with a complete neural arch, presacrals II–IV with thicker and longer transverse processes than V–VIII. Sacrum with expanded diapophyses, the leading and trailing edges roughly equally angled, the articulation type IIB sensu Emerson [42]. Urostyle bicondylar, long, not broadening posteriorly, without lateral processes and with a low dorsal ridge. Pectoral girdle without ossified prezonal or postzonal elements, with ossified clavicles. Clavicle thin and curved, with a simple lateral junction, shorter than the coracoid. Coracoid fairly narrow, not flared laterally, strongly flared medially with a large medial articular surface with the contralateral. Scapula slender, with a thin pars acromialis, the cleithral border concave. Cleithrum ossified for two thirds the width of the scapular border, thickened anteriorly. Suprascapula unossified. Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna slender with a distinct sulcus intermedius. Carpals poorly ossified, composed of radiale, ulnare, prepollical element, element Y, and large post-axial element formed by carpals 3–5. Carpal 2 has either been lost or fused to the latter element. Finger phalangeal formula is reduced (1-2-3- 3), and the terminal phalanx of the first finger is a small, round element. Small distal knobs on terminal phalanx of finger 3. Prepollex thin and cultriform and extending only to the base of the first metacarpal. Pubis calcified, iliac shafts passing ventral to and beyond sacrum, nearly cylindrical, without a dorsal crest and with a weak dorsal prominence and shallow oblique groove. Femur weakly sigmoid, almost lacking a posterior crest. Tibiofibula shorter than femur, with a sulcus intermedius. Tibiale and fibulare fused proximally and distally. T1 and T2+3 tarsals present, T1 considerably smaller than T2+3, plus a small additional ossification (possibly a sesamoid) between the bases of metatarsals 2 and 3. Centrale present, roughly the size of T2+3. Prehallux subtriangular. Phalangeal formula reduced (1-2-3-4-3). Terminal phalanges of toes 2–4 with almost T-shaped knobs. Distribution, natural history, and conservation status. This species is known only from a single specimen from Andohahela National Park in southeast Madagascar (Fig 6). The species is larger than the other members of the genus Mini, and its ecology may differ accordingly. Advertisement calls were not recorded. At present it is not possible to estimate its distribution or population status, and we prefer to suggest this species be considered Data Deficient until more information is available. Etymology. We use the specific epithet ‘ature’ as an arbitrary combination of letters, in order to form a pun on ‘miniature’ from the name in apposition, in reference to the small size of this species. It is to be regarded as an invariable noun.