Search

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.

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"]}

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 Köhler 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.

Ndriantsoa, Serge H., Riemann, Jana C., Vences, Miguel, Klages, Johannes, Raminosoa, Noromalala R., Rödel, Mark-Oliver, Glos, Julian (2013): A new Stumpffia (Amphibia: Anura: Microhylidae) from the Ranomafana region, south-eastern Madagascar. Zootaxa 3636 (4): 575-589, DOI: http://dx.doi.org/10.11646/zootaxa.3636.4.5