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Rakotoarison, Andolalao, Scherz, Mark D., Bletz, Molly C., Razafindraibe, Jary H., Glaw, Frank, Vences, Miguel (2019): Description of the lucky Cophyla (Microhylidae, Cophylinae), a new arboreal frog from Marojejy National Park in north-eastern Madagascar. Zootaxa 4651 (2): 271-288, DOI: https://doi.org/10.11646/zootaxa.4651.2.4

Taxonomy is the cornerstone of extinction risk assessments. Currently, the IUCN Red List treats species complexes either under a single overarching species name-resulting in an unhelpfully broad circumscription and underestimated threat assessment that does not apply to any one species lineage-or omits them altogether-resulting in the omission of species that should be assessed. We argue that taxonomic uncertainty alone, as in species complexes, should be grounds for assessment as Data Deficient (DD). Yet, use of the DD category is currently discouraged, resulting in assessments based on poor data quality and dismissal of the importance of taxonomic confidence in conservation. This policy may be leading to volatile and unwarranted assessments of hundreds of species across the world, and needs to be revised. To illustrate this point, we here present a partial taxonomic revision of torrent frogs from eastern Madagascar in the Mantidactylus subgenus Hylobatrachus. Two named species, Mantidactylus (Hylobatrachus) lugubris and M. (H.) cowanii, and several undescribed candidate species are recognised, but the application of the available names has been somewhat ambiguous. In a recent re-assessment of its conservation status, M. (H.) lugubris was assessed including all complex members except M. (H.) cowanii within its distribution, giving it a status of Least Concern and distribution over most of eastern Madagascar. After describing two of the unnamed lineages as Mantidactylus (Hylobatrachus) atsimo sp. nov. (from southeastern Madagascar) and Mantidactylus (Hylobatrachus) petakorona sp. nov. (from the Marojejy Massif in northeastern Madagascar), we show that Mantidactylus (Hylobatrachus) lugubris is restricted to the central east of Madagascar, highlighting the inaccuracy of its current Red List assessment. We propose to re-assess its status under a more restrictive definition that omits well-defined candidate species, thus representing the actual species to which its assessment refers, to the best of current knowledge. We recommend that for species complexes in general, (1) nominal lineages that can be confidently restricted should be assessed under the strict definition, (2) non-nominal species-level lineages and ambiguous names should be prioritised for taxonomic research, and (3) ambiguous names should be assessed as DD to highlight the deficiency in data on their taxonomic status, which is an impediment to their conservation. This would reduce ambiguity and underestimation of threats involved in assessing species complexes, and place the appropriate emphasis on the importance of taxonomy in anchoring conservation.

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.

Uroplatus finaritra sp. nov. Figs. 1–5, Table 1 Uroplatus sp. G—Raxworthy et al. (2008) Uroplatus sp. Ca9— Ratsoavina et al. (2013) LSID: urn:lsid:zoobank.org:act: E6A1407B-7C9E-46F6-8FEC-D120A111A344 Holotype. UADBA-R 70489 (field number APR 12522), adult female, collected on the eastern slope of Marojejy by a tributary of the Ampanasatongotra River, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.4559°S, 49.7749°E (780 m a.s.l.) on the night of 15 May 2016 by A. P. Raselimanana. Paratypes. UADBA-R 70493 (field number APR 12591), adult male, UADBA-R 70490 (APR 12590), adult male, and UADBA-R 70491 (APR 12589), adult female, three specimens collected in the same locality as the holotype (750–810 m a.s.l.) on 18 May 2016 by A. P. Raselimanana; UADBA-R 70492 (APR 12691), adult male, and UADBA-R 70494 (APR 12692), juvenile female, two specimens collected on the western slope of Marojejy by Tsiasisa River, 2.2 km east of Antanimbaribe, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.5074°S, 49.6153°E (845 m a.s.l.) on the night of 28 May 2016 by A. P. Raselimanana; UADBA-R 70501 (MSZC 0253), subadult female, ZSM 458 /2016 (MSZC 0271), subadult female, two specimens collected near the path below Camp 1 (Camp Mantella) of Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.438°S, 49.776°E (450 m a.s.l.) on 21–23 November 2016 by M. D. Scherz, M. Bletz, A. Rakotoarison, J. Razafindraibe, A. Razafimanantsoa, and M. Vences. Etymology. Finaritra is a Malagasy word for special greetings but also means healthy and happy. We refer to our delight in describing this splendid and exceptionally large species from a clade of generally small-sized leaftailed geckos. The name is an invariable noun in apposition. Diagnosis. Uroplatus finaritra sp. nov. is assigned to the Uroplatus ebenaui group based on its relatively small size and its triangular head with supraocular spines, and overall leaf-mimicking aspect. It is characterised by (1) a dark red oral mucosa, (2) a comparatively large body size (82.5–95.3 mm SVL in adults), (3) 7–8 lamellae (exceptionally 6) under the third finger and toe, and (4) a long (length 53–65% of SVL), wide (width 16–18% of SVL), leaf-shaped tail. It differs from all members of the U. fimbriatus group (U. fimbriatus, U. giganteus, U. henkeli, U. sikorae, and U. sameiti) and U. lineatus by its smaller size (SVL 82.5–95.3 mm vs. 85–200 mm) and the lack of lateral dermal fringes on any part of the body (vs. presence except in U. lineatus). Also, Uroplatus finaritra sp. nov. diverges from the species of the U. fimbriatus group by their lack of lateral body compression. The prominent triangular head, supraocular spines, and rather smooth skin distinguish Uroplatus finaritra sp. nov. from U. alluaudi, U. guentheri, U. malahelo and U. pietschmanni. Amongst the U. ebenaui group, the rather long tail (TAL/SVL 0.53–0.68) differentiates Uroplatus finaritra sp. nov. from U. ebenaui, U. fiera, U. fotsivava, and U. kelirambo (TAL/SVL 0.22–0.44), and furthermore it differs from U. fiera and U. fotsivava by a pigmented oral mucosa (vs. unpigmented). The new species differs from U. finiavana by a wider tail (TAW/SVL 0.16–0.18 vs. 0.05–0.14), and by its pigmented oral mucosa (vs. unpigmented). By its large tail and pigmented oral mucosa, the new species is most similar to U. malama and U. phantasticus. It differs from U. phantasticus by larger size (SVL 83–95 mm vs. 52–76 mm), a generally shorter tail (TAL/SVL 0.53–0.65 vs. 0.62–0.76), a typically higher number of lamellae under the third finger and toe (6–8 vs. 5–7), and the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black). It differs from U. malama by a spinier integument in some adult males, by the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black), and probably by a narrower tail (TAW/SVL 0.16–0.18 vs. 0.24). In the mitochondrial phylogeny, the new species occupies an isolated position within the U. ebenaui group, without affinities to the morphologically similar U. phantasticus and U. malama (which are not each other's closest relatives either). The nuclear gene c-mos reveals haplotype sharing of the new species with U. kelirambo and two undescribed candidate species (U. sp. Ca3 and Ca4) occurring in the same region of Madagascar, but morphologically very different by their short tails. TABLE 1. Morphological measurements and scale counts of Uroplatus finaritra sp. nov. For abbreviations used, refer to Methods; additional abbreviations: M, male; F, female; sa, subadult; j, juvenile; HT, holotype; PT, paratype; na, not applicable (due to missing original tail); nm, not measured. All measurements in mm. Description of the holotype. Adult female in fair condition, with a fully intact tail, both of its hind limbs fractured (Fig. 2). SVL 95.3 mm, tail length 50.9 mm, maximum tail width 17.0 mm; for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis indistinct and concave; snout sloping weakly downwards anteriorly; snout weakly depressed, fairly long (1.7 times longer than eye diameter); eyes large (eye diameter 6.5 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.1 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in ventral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and very few, miniscule spines on the hind limbs (one on the knee, as well as a small flap of skin), and two miniscule spines on the forelimb (on the elbow and near the wrist); forelimb reaches the canthus rostralis when adpressed forward and midbody when adpressed backwards along body (forelimb length/axilla–groin distance 33.4/ 50.2 mm = 67%), hindlimb reaches ¾ up the body when adpressed forward along body (hindlimb length/axilla–groin distance 41.4/ 50.2 mm = 83%); original tail length 53% of snout–vent length, membranous borders of the tail symmetrical, without any emarginations, long and leafshaped, with a thin, borderless spatulate tip. Toes bear small claws, with the distal phalange not much wider than the rest of the digit; the third finger bears 7 lamellae, the third toe 8 lamellae. Nares separated from each other by at least 10 small granular scales, from the first supralabial by 3 scales, and from the rostral scale by 5 scales; first supralabial not taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 25/26 (right/left) supralabials and 25/25 infralabials; no enlarged postmental scales or chin shields; dorsal and ventral scales of head, neck, body, limbs, and tail small, granular, juxtaposed and largely of uniform size, except on the posterior ventral abdomen, where they are slightly larger than the rest of the body, and arranged almost uniformly. Two curved lines (rows of slightly enlarged scales) extending from the posterolateral parts of the head (nuchal region) converge on the neck forming a V-shaped pattern (neck triangular line). A similar, moderately distinct, fairly straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Another, less distinct line is present connecting the anterior angle of the eyes over the snout in a bowed line. The body possesses very few dermal spines—a prominent pointed supraocular spine, a single spine at the posterior angle of the head, and a small number of spines on each limb (described above). No axillary pits present. Colouration. After two years of preservation in 70% ethanol the colour is slightly faded (Fig. 2). All dorsal surfaces of the specimen are a ruddy brown, fading to a salmon brown in the neck region and at the insertion of the hind limbs. A fine network of blackish lines is present across the body, with the following notable features: a somewhat straight line connecting the suprocular spines (described above); a v-shaped marking on the neck, which extends into a vertebral stripe that reaches all the way to the tail tip; a number of posteriorly oriented chevrons from the mid-dorsum to the belly, becoming thinner posteroventrally, three of which are most distinct. The exterior of the shank and outermost two toes are grey-black, whereas the inner foot is brown. The dorsal snout is darker than the rest of the head. The tail is not different in colour or pattern from the dorsal body. A distinct lateral line runs along the ventrolateral edges of the abdomen from the insertion of the leg to the axilla. The ventral colouration is uniformly orange-brown, with a faint, symmetrical pattern of dark lines on the chin. The soles of the hands and feet are grey. The ventral tail is the colour of the trunk mottled with grey splotches. A faint cream tear-like marking is present at the posterior corner of the eye above the supralabial. The oral mucosa is dark red, with a pink tongue and cream under the eyes and along the lower jaw. In life, the iris colour was silver externally and rusty around the pupil (Fig. 3d). Variation. Molecular data are also available from a previous paper (Raxworthy et al. 2008) for one specimen (RAN 42274) from Marojejy as Uroplatus sp. G, included for 12S and cytochrome b as U. sp. Ca 9 in Ratsoavina et al. (2013). The available sequence for 12S agrees with those of our specimens. The paratypes were sexed as adult males by the presence of everted hemipenes (UADBA-R 70493) or presence of distinctly enlarged tail base (UADBA-R 70490 and 70492), and distinctly notched tail edges; or as females based on the absence of hemipenes or hemipenial bulges, presence of visible oviduct or ovary structures after dissection, and generally smooth tail edges. In the smallest specimen (UADBA-R 70494) female inner reproductive organs were observed but were only weakly developed, and we therefore consider this specimen as juvenile. The sexual maturity of two further specimens (UADBA-R 70501 and ZSM 458/2016) is unclear, as they are considerably smaller than the large adult females (UADBA-R 70491 and 70489) but have similar gonad development. We tentatively consider these to be subadults. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2–9, and is more in males than in females, as males are more spiny than females in general, and have more heterogeneous scalation. Forelimbs can bear a minimum of two spines, or as many as eight, with a small spine present on the elbow. Hind limbs can bear 2–15 spines, and the dermal knee flap almost always bears at least a small spine, which is smaller in females than males. In males the tail is moderately emarginated, with semicircular emarginations that tend to be asymmetrical. In females the tail is without emarginations and has smooth edges. Internarial scales range from nine to ten, supralabials from 25–29, and infralabials from 21–25. The third finger bears 7–8 lamellae (6 in one juvenile specimen, UADBA-R 70494), the third toe bears 7–8 lamellae. The colouration is varied, as is commonly the case in Uroplatus species (Figs. 3–4). The line on the head connecting the supraocular spines is generally bowed, but can be slightly pointed posteriorly, and in UADBA-R 70492, it is irregular at its midpoint. The dark vertebral stripe is only present in the holotype, UADBA-R 70491 and 70494, and in other specimens is either absent, or is cream and not dark (in UADBA-R 70492 and 70490). Overall dorsal colouration differs dramatically, from almost completely reddish brown (UADBA-R 70501) to highly mottled greys and blacks (UADBA-R 70493). Irregular flecks can be present or absent on any part of the body. The different colouration of the outer shank and foot is always present, and a cream tear-like marking below the eye is almost always present (absent in UADBA-R 70494 and 70501). The ventrolateral stripe is always present but not always distinct (indistinct in UADBA-R 70493). Ventral colouration is as variable as dorsal colouration. UADBA-R 70491 is extremely similar to the holotype, and to a lesser degree 70492, 70494, and 70501 resemble the holotype ventrally. UADBA-R 70490 and 70493 differ dramatically in being darker in colour, having irregular flecks of cream and reddish brown, and lacking the symmetrical markings on the chin. In life, the colouration of the iris is typically silvery but can be red (Figs. 3c, f), and the oral mucosa was consistently dark red (Fig. 5). Distribution. At present this species has only been collected on the Marojejy Massif, at altitudes from 450–845 m a.s.l. Natural History. The holotype was collected active at night on a stem 0.5 m above the forest floor in nearly closed canopy humid forest at the bottom of a slope, in the vicinity of a Uroplatus lineatus specimen. UADBA-R 70490, 70491, and 70493 were collected on small branches and leaves between 1.5 and 2.5 m above the ground in similar forest. UADBA-R 70492 and 70494 were also active on branches 3–4.5 m from the forest floor in nearly closed-canopy humid forest with a thick understory and ferns. UADBA-R 70501 and ZSM 458/2016 were collected active at night on thin branches (up to 4 m above the ground) in closed-canopy, low-altitude rainforest. We consider the various collection localities to represent a single Threat-Defined Location in the sense of the International Union for the Conservation of Nature (IUCN 2012), as we have recently done for several frog species from low- to mid-elevation forests in Marojejy (Scherz et al. 2016, Rakotoarison et al. 2017). In line with these assessments, we recognise that there is on-going habitat degradation due to illegal logging activity, especially at low altitudes in and around Marojejy National Park. The Extent of Occurrence and Area of Occupancy of the new species is certainly greater than 10 km 2, but is likely less than 500 km 2, and as such we recommend a status of Endangered for it under criterion B1ab(iii).

Guibemantis albomaculatus sp. nov. urn:lsid:zoobank.org:act: D33E461D-331B-4FE4-8808-DB476F9FB2AA Figs 1–4 Suggested common name: White-spotted Malagasy Pandanus Frog. Diagnosis Assigned to the subgenus Pandanusicola of the genus Guibemantis based on its small body size, phytotelm-breeding habitats (in Pandanus plants), moderate webbing between toes, connected lateral metatarsalia, the presence of both inner and outer metatarsal tubercles, type 2 femoral glands in males, and DNA sequence similarities (Glaw & Vences 2006). The new species is characterized by the unique combination of the following characters: (1) male SVL 18.9–22.9 mm, female SVL 20.9–26.7 mm, (2) two conspicuous parallel light-colored dorsolateral lines on the dorsum, (3) small white spots on the darkly colored arms, legs and flanks, (4) dark ventral surfaces to hands and feet, (5) distinct rostral stripe and (6) orange femoral glands. Etymology The specific epithet is used as a descriptive adjective and refers to the small white spots present on the arms, legs and flanks. Material examined Holotype MADAGASCAR: adult ³, Montagne d’Ambre National Park, northern Madagascar [geographical coordinates not taken, but most likely from a Pandanus plant very close to the Gîte d’étape (12.5270° S, 49.1720° E, 1055 m a.s.l.) or less likely from the Voie des mille arbres (ca 12.5200° S, 49.1756° E, 1052 m a.s.l.)], 18 Feb. 2003, F. Glaw, R.D. Randrianiaina and A. Razafimanantsoa leg. [ZSM 0895 /2003 (FGMV 2002.905)] (Fig. 3). Paratypes (n = 19) MADAGASCAR: 5 specs, Manongarivo Special Reserve, 20 Feb. 1992, C.J. Raxworthy, A. Raselimanana and J.B. Ramanamanjato leg. (UMMZ 212592–212596); 1 spec., Montagne d’Ambre National Park, Antomboka River, 1150 m a.s.l., 15 Nov. 1991, C.J. Raxworthy, A. Raselimanana and J.B. Ramanamanjato leg. (UMMZ 212585); 3 specs Montagne d’Ambre National Park, Antomboka River, 950 m a.s.l., 21 Nov. 1991, C.J. Raxworthy, A. Raselimanana and J.B. Ramanamanjato leg. (UMMZ 212587, UMMZ 212588, UMMZ 212591); 2 specs Manongarivo Special Reserve, “Camp 0”, 13.9756° S, 48.4267° E, 688 m a.s.l., 1–2 Feb. 2003, F. Glaw, R.D. Randrianiaina and M. Vences leg. [ZSM 0816/2003 (FGMV 2002.735), ZMA 19582 (FGMV 2002.733)]; 1 spec., same collecting data as for preceding (UADBA uncatalogued, FGMV 2002.736); 1 spec., Montagne d’Ambre National Park, 12.5275° S, 49.1725° E, 1010 m a.s.l., 19 Mar. 2000, F. Glaw, K. Schmidt and M. Vences leg. [ZSM 496/2000 (FGMV 2000.356)]; 4 specs, Montagne d’Ambre National Park, same collection data as for holotype [ZSM 0893/2003 (FGMV 2002.0901), ZSM 0894/2003 (FGMV 2002.0904), UADBA uncatalogued (FGMV 2002.902), and ZMA 19660 (FGMV 2002.997)]; 1 &female; [UADBA uncatalogued (FGZC 1010)] and 1 juv. [UADBA uncatalogued (FGZC 1012)], Montagne d’Ambre National Park, Voie des mille arbres, 12.5167° S, 49.1767° E, 1050 m a.s.l., 23 Feb. 2007, F. Glaw, P. Bora, H. Enting, J. Köhler and A. Knoll legs. The six paratypes from UADBA and ZMA were not available for morphometric comparison, but were sequenced and attributed genetically to G. albomaculatus sp. nov. (see Fig. 1). Description of the holotype Adult male. Head longer than wide and slightly wider than body; snout rounded in dorsal, ventral and lateral views; canthus rostralis rounded; nostrils much nearer to tip of snout than to eye, internarial distance 83% of interorbital distance; tympanum distinct 42% of horizontal eye diameter; dark supratympanic fold present from posterior edge of eye around the tympanum nearly to the arm insertion; one small round patch of vomerine teeth medial between eye and choanae on either side of head; tongue damaged, shape of tip unknown. Arms thin, lower arm is 91% of hand length; relative finger length 1 After seven years in preservative, the dorsal background coloration is medium brown which contrasts with the darker brown flanks. Two prominent light dorsolateral lines separate the medium brown color of the dorsum from the dark brown of the flanks. A single, relatively thick rostral stripe is present. Dorsal color of arms, legs, hands and feet is medium brown with abundant small white spots throughout. Vague light annuli are found on some digits just proximal to the finger and toe discs but are not prominent. Toe and finger discs have a similar dark coloration as the dorsal surface of the hands and feet, except for the discs of fingers I and II, which have a much lighter coloration. Venter is cream with abundant brown spots along jawline, chin and especially on the ventral surface of the hindlimbs. The pupil appears black and the iris is white. Coloration in life The life coloration of the holotype is unknown. However, based on photographs of live individuals from Manongarivo and Montagne d’Ambre (Fig. 4), there appear to be a number of coloration differences compared to preserved material. In life, the iris color is gold or bronze (white or gray in preservative). Portions of the arms and fingers as well as lateral areas of the head have a yellow or yellow-green color in life (beige in preservative). The more lightly colored toe and finger discs are a dull yellow (beige or whitish in preservative). The ventral skin is largely translucent such that chest and belly musculature and bones in the limbs are easily visible (not apparent in preservative). In males, femoral glands (type 2, as defined by Glaw et al. 2000) appear orange in life. Sexually mature males also possess a distinct white coloration on the lateral portions of the throat. A color photograph of a specimen of this species from Manongarivo is shown in Glaw & Vences (2007: 203, fig. 4). Variation The new species is similarly sized to most other Pandanusicola (overall mean SVL 22.8 ± 2.1 mm, n = 15). Females appear to be larger on average than males (female mean SVL 24.0 ± 1.7 mm, n = 9; male mean SVL 21.3 ± 1.5 mm, n = 6). While all individuals seem to possess a dark rostral stripe, some individuals have an additional gold colored one medial to the first. The small white spots on the dorsal surface of the arms, legs and flanks also vary in coverage and density among individuals. Some individuals only possess them in abundance on the hindlimbs and sparingly elsewhere. Other individuals have an abundance of these spots not only on both limbs and the flanks, but also on the back and head as well. While yellowish dorsal coloration is present in all known individuals, the dorsal base color intermixed with it differs among individuals. Most individuals have a dorsal base color of medium to dark brown but one individual was notably lighter than other specimens examined (Fig. 4B). The prominence and length of the two parallel dorsal lines also varies among individuals, with those from Manongarivo typically being more distinct. Some individuals possess a small but distinct light-colored spot between the eyes, other individuals lack this feature. Many individuals possess no obvious markings on the dorsum but a few have several vague dark splotches. On the ventral surface, most individuals possess fine brown spots on the jawline and on the undersurface on the limbs. Other individuals have these spots on the throat and chest as well. No variation was observed in the webbing on the hand. However, webbing on the foot was variable for some digits (range, where variable, given in parentheses): 1(1) 2i(1) 2e(1) 3i(2) 3e(1–2) 4i(3) 4e(2.75–3) 5(1–2). The distinct light-colored annuli present before the terminal phalange of each digit, that are prominent in some Pandanusicola, are usually only weakly developed but can be conspicuous in some individuals. Differential diagnosis Among Pandanusicola, the new species is distinguished from G. annulatus Lehtinen, Glaw & Vences, 2011, G. bicalcaratus (Boettger, 1913), G. flavobrunneus, G. liber, G. pulcher, G. punctatus and G. tasifotsy by the presence of two conspicuous parallel light-colored dorsolateral lines, by the presence of small white spots on more darkly colored arms, legs and flanks and by large genetic differences. By comparison, G. pulcher is a characteristic bright green in color with large irregular dark dorsal spots; G. bicalcaratus is yellowish or brownish dorsally with usually relatively few irregularly shaped dark spots; G. flavobrunneus is much larger (up to 38 mm SVL; Glaw & Vences 2007) and is primarily yellow with extensive brown dorsal markings; G. liber is rather variable in coloration but tends to be brownish overall, the entire surface of the throat of males is bright white, and this species does not breed in Pandanus plants; G. tasifotsy does not breed in Pandanus either and has a copper metallic coloration on the nares and/or eye orbits and prominent white lateral blotches; G. annulatus and G. punctatus are cream or olive-colored (respectively) and have abundant small dark spots; G. methueni (Angel, 1929) and G. wattersoni can have parallel dorsolateral light colored lines on the dorsum but most individuals lack this feature and it is usually poorly developed in those individuals that possess it. Furthermore, G. wattersoni is primarily a yellow or light brownish frog dorsally and the flanks and legs in G. wattersoni are always lightly colored whereas they are typically very dark in the new species. In G. methueni, the ventral surfaces of the hands and feet are very lightly colored, whereas they are dark in the new species. Guibemantis albolineatus (Blommers-Schlösser & Blanc, 1991) lacks both vomerine teeth and a rostral stripe (both present in the new species), has a distinctly shaped snout that is square in dorsal and ventral view and is notably smaller in overall body size (see Table 1). The recently described G. milingilingy differs by lacking white spots (present in the new species), by having banded forelimbs (unbanded in the new species), by having yellow femoral glands in males (orange in the new species) and by having a mid-dorsal stripe (absent or poorly developed in the new species). No other described species of Pandanusicola has the combination of light colored parallel dorsal lines with small white spots on the dark unbanded arms, legs and flanks. Distribution At present G. albomaculatus sp. nov. is known from rainforest habitats in two areas: the Manongarivo Special Reserve (type locality) and Montagne d’Ambre National Park, both in northern Madagascar (Fig. 2). The elevational range so far documented is from 950–1150 m. Based on the limited knowledge of the distribution, we suggest a conservation status of "Data Deficient". Natural history Little is known of the natural history of this species. Specimens have only been collected from Pandanus plants and are probably obligate Pandanus breeders, like most other species in this group. Vocalizations have not been recorded and eggs and larvae have yet to be documented. Available names and comparisons The only available synonyms that could possibly be applied to this species are Gephyromantis albogularis Guibé, 1947 and Gephyromantis variabilis Millot & Guibé, 1951, both of which were confidently assigned to Guibemantis liber by Lehtinen et al. (2011). Males of Guibemantis liber have a throat that is bright white over its entire surface. The whitish coloration on the throat of males in G. albomaculatus sp. nov. does not cover the entire surface nor is it as brightly white as in G. liber. Remark This species has been previously listed as Guibemantis sp. aff. albolineatus “Manongarivo” in Glaw & Vences (2007: 202–203), Guibemantis bicalcaratus (Manongarivo) in Lehtinen et al. (2007) and Guibemantis sp. Ca 12 in Vieites et al. (2009) and Perl et al. (2014).

Guibemantis woosteri sp. nov. urn:lsid:zoobank.org:act: BB4E147F-EB7A-41F9-9E1B-94B92521EE7F Figs 1���3, 5 Suggested common name: Wooster���s Malagasy Pandanus Frog. Diagnosis Assigned to the subgenus Pandanusicola of the genus Guibemantis based on its small body size, phytotelm-breeding habitats (in Pandanus plants), moderate webbing between toes, connected lateral metatarsalia, the presence of both inner and outer metatarsal tubercles, type 2 femoral glands in males, and DNA sequence similarities (Glaw & Vences 2006). The new species is characterized by the unique combination of the following characters: (1) male SVL 19.4���21.8 mm, (2) two fairly conspicuous parallel light-colored dorsolateral lines on the dorsum, (3) well-defined dark bars on the forearms and sometimes on the legs, (4) a conspicuous light area near the forearm insertion, (5) dark ventral surfaces to hands and feet, (6) thick, single dark rostral stripe, (7) distinctly reddish color dorsally surrounding the eyes, (8) white spots of the dorsum and limbs and (9) gray femoral glands in males. Etymology This new species is dedicated to the College of Wooster and its faculty, staff, students and alumni in celebration of 150 years of educational excellence. Material examined Holotype MADAGASCAR: adult &female;, Marojejy National Park at ���Camp Simpona���, 14.437�� S, 49.743�� E, 1326 m a.s.l., 18 Feb. 2005, F. Glaw, M. Vences and R.D. Randrianiaina leg. [ZSM 5063 /2005 (ZCMV 2044)] (Fig. 3). Paratypes (n = 15) MADAGASCAR: 8 adults, Manantenina River in Marojejy National Park, 700���1300 m a.s.l., Nov. 1992, R.A. Nussbaum, C.J. Raxworthy, A. Razafimanantsoa and A. Razafimanantsoa leg. (UMMZ 212601��� 212602, UMMZ 212604���212606, UMMZ 212613, UMMZ 212615, UMMZ 212617); adult ��, same data as for holotype [ZSM 5062/2005 (ZCMV 2011)]; 1 &female;, Marojejy National Park, Camp 3 ���Simpona���, 14.4366�� S, 49.7434�� E, 1325 m a.s.l., 17 Nov. 2017, M.D. Scherz, A. Rakotoarison, M. Bletz, M. Vences and J. Razafindraibe leg. [ZSM 425/2016 (ZCMV 15185)]; 1 �� and 1 juv., Marojejy, between Camp 2 ���Marojejia��� and Camp 3 ���Simpona���, no coordinates taken, 18 Nov. 2016, M.D. Scherz, A. Rakotoarison, M. Bletz, M. Vences and J. Razafindraibe leg. [ZSM 426/2016 (ZCMV 15198), UADBA uncatalogued (ZCMV 15200)]; 1 spec., Marojejy National Park, same data as for holotype, but collected on 16 Feb. 2005 [(FGZC 2878), UADBA uncatalogued]; 1 spec., same data as holotype (ZCMV 2045); 1 spec., same data as holotype, but collected above Camp Marojejia (ZCMV 2046). The paratypes from UADBA were not available for morphometric comparison, but were sequenced and attributed genetically to G. woosteri sp. nov. (see Fig. 1). Referred specimens Based on similarity in coloration (no genetic data available), we also tentatively assign to this species the series UMMZ 191339���191340 and UMMZ 191343���191344; adults from Ambatovaky Special Reserve (600���800 m), collected by C.J. Raxworthy on 21 February 1990. Description of the holotype Adult female. Head longer than wide and slightly wider than body; snout rounded in dorsal, ventral and lateral views; canthus rostralis rounded; nostrils very near to the tip of snout, internarial distance 79% of interorbital distance; tympanum distinct 38% of horizontal eye diameter; very dark and prominent supratympanic fold present from posterior edge of eye around the tympanum nearly to the arm insertion; one small round patch of vomerine teeth medial between eye and choanae on either side of head; tongue stout and strongly bifid. Arms thin, lower arm is 93% of hand length; relative finger length 1et al. 2000) appear gray in life with large granules in the gland, each of which has a white spot (Fig. 5C). Sexually mature males also possess a white coloration on portions of the throat. Variation Like most Pandanusicola, this new species is small (overall mean 22.1 �� 1.6 mm SVL, n = 14). Females are larger (mean 23.8 �� 1.0 mm SVL, n = 5) than males (mean 21.1 �� 0.8 mm SVL, n = 9) on average. Some individuals have relatively few small white dorsal spots (primarily on the hindlimbs), while others have dense spots over much of the dorsal surfaces (contrast Fig. 5A and 5B). The light dorsolateral lines on the dorsum are conspicuous in some individuals but much less so in others. Dark spotting on the dorsum also varies from none to many. Dark bars on legs vary from reasonably distinct (Fig. 5A) to inconspicuous (Fig. 5B). Head coloration can be variable also, with several specimens having a distinctly lighter coloration anterior to the eyes than posterior and one specimen possessed a distinct interocular bar. Many, but not all, specimens had a distinct light spot in the middle of the snout. The color of the digits and toepads also varies. Some individuals have distinct dark bars on the digits and this seems to be an extension of the bars on the forearms. The color of the toepads is similarly variable with the pads on some digits being dark and other digits being light. A few individuals had indications of faint annuli on the digits proximal to the toepads or a second rostral line parallel to the first (as in G. annulatus and G. wattersoni), however, these are usually not well developed. Ventrally, coloration varies from having dark markings only along the jawline to individuals with dark spotting on the throat, chest and/or thighs. The ventral surfaces of the hands and feet are apparently always dark. One individual (UMMZ 212604) had especially rough granular patches on the belly, reminiscent of those recently described from G. tasifotsy. No variation was observed in the webbing on the hand. However, webbing on the foot was variable for some digits (range, where variable, given in parentheses): 1(1) 2i(1) 2e(1) 3i(2) 3e(1���2) 4i(2.5���3) 4e(2.5���3) 5(1���2). Differential diagnosis Among Pandanusicola, the new species is distinguished from G. annulatus, G. bicalcaratus, G. flavobrunneus, G. liber, G. pulcher, G. punctatus and G. tasifotsy by the presence of two conspicuous parallel dorsolateral light-colored lines on the dorsum, by the presence of small white spots on more darkly colored arms, legs and flanks and by large genetic differences. By comparison, G. pulcher is a characteristic bright green in color with large irregular dark dorsal spots; G. bicalcaratus, is yellowish or brownish dorsally with usually relatively few irregularly shaped dark spots; G. flavobrunneus is much larger (up to 38 mm SVL; Glaw & Vences 2007) and is primarily yellow with extensive brown dorsal markings; G. liber is rather variable in coloration but tends to be brownish overall, the entire surface of the throat of males is bright white, and this species does not breed in Pandanus plants; G. tasifotsy does not breed in Pandanus either, and has a copper metallic coloration on the nares and/ or eye orbits and prominent white lateral blotches; G. annulatus and G. punctatus are cream or olivecolored, respectively, and have abundant small, well-defined dark spots; G. methueni and G. wattersoni can have parallel light-colored dorsolateral lines on the dorsum but most individuals lack this feature and, if present, it is usually poorly developed. Furthermore, G. wattersoni is a yellow or light brownish frog dorsally and its flanks and legs are always lightly colored whereas they are typically dark in the new species. In G. methueni, the ventral surfaces of the hands and feet are very lightly colored, whereas they are dark in the new species. Guibemantis albolineatus lacks both vomerine teeth and a rostral stripe (both present in the new species), has a distinctly shaped snout that is square in dorsal and ventral view and is notably smaller in overall body size (see Table 1). The recently described G. milingilingy differs by lacking the characteristic white spots on the dorsum and limbs (versus presence), by lacking the reddish colored skin on the eye orbits (versus presence) and by having yellow femoral glands in mature males (gray in the new species). Finally, G. woosteri sp. nov. differs from G. albomaculatus sp. nov. in having well-defined dark bars on the forearms and sometimes on the legs, a conspicuous light area near the forearm insertion (both of which G. albomaculatus sp. nov. lacks) and gray femoral glands in males with distinct white spots (femoral glands are orange in male G. albomaculatus sp. nov.). No other described species of Pandanusicola has the combination of light colored parallel dorsolateral lines with small white spots on the legs and flanks and well-defined dark bars on the forearms. Distribution Currently, G. woosteri sp. nov. is reliably known only from mid-elevation rainforests in Marojejy National Park, the type locality, in the North-East of Madagascar (Fig. 2), from roughly 700���1325 m a.s.l. In addition, we here consider specimens from the Ambatovaky Special Reserve in the Northern Central East of the island as possibly belonging to this species (Fig. 2), but this identification requires confirmation by molecular data. Based on the limited knowledge of the distribution, we suggest a conservation status of "Data Deficient". Natural history Only known from Pandanus plants. One adult female attributed to this species (UMMZ 212617) had four visible eggs in the body cavity (egg diameter 0.8 mm) and another (UMMZ 191340; from Ambatovaky) had six (egg diameter 1.8 mm) possibly suggesting a small clutch size. Tadpoles and breeding vocalizations are unknown. Available names and comparisons The only available synonyms that could possibly be applied to this species are Gephyromantis albogularis Guib��, 1947 and Gephyromantis variabilis Millot & Guib��, 1951, both of which were confidently assigned to Guibemantis liber by Lehtinen et al. (2011). Guibemantis liber males have a throat that is bright white over its entire surface. The whitish coloration on the throat of males in G. woosteri sp. nov. does not cover the entire surface nor is it as brightly white as in G. liber. Remark In Vieites et al. (2009) and Perl et al. (2014), the holotype of G. woosteri sp. nov. (ZSM 5063/2005 / ZCMV 2044) and one paratype (ZSM 5062/2005 / ZCMV 2011) were listed as two different candidate species (Ca15 and Ca17, respectively). However, the 16S sequence data reported here show them to be nearly identical genetically (Fig. 1) and they are also morphologically similar (Table 1). Thus, we consider them to be conspecific. In another earlier molecular analysis, Lehtinen et al. (2007) found the holotype of G. woosteri sp. nov. (ZSM 5063/2005) to be phylogenetically related to UMMZ 212597 (also from Marojejy) and UMMZ 212922 from Cap Est (in the same region, but further east on the coast). However, these forms are 7.3% and 8.1% different in the 12S and 16S mitochondrial genes (Lehtinen et al. 2007) and are morphologically distinct from G. woosteri sp. nov. We have, therefore, not included these specimens in the definition of G. woosteri sp. nov. as they likely represent additional undescribed taxa., Published as part of Lehtinen, Richard M., Glaw, Frank, Vences, Miguel, Rakotoarison, Andolalao & Scherz, Mark D., 2018, Two new Pandanus frogs (Guibemantis: Mantellidae: Anura) from northern Madagascar, pp. 1-20 in European Journal of Taxonomy 451 on pages 13-17, DOI: 10.5852/ejt.2018.451, http://zenodo.org/record/3814542, {"references":["Glaw F. & Vences M. 2006. Phylogeny and genus-level classification of mantellid frogs (Amphibia, Anura). Organisms Diversity and Evolution 6: 236 - 253. https: // doi. org / 10.1016 / j. ode. 2005.12.001","Glaw F., Vences M. & Gossmann, V. 2000. A new species of Mantidactylus (subgenus Guibemantis) from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae). Journal of Natural History 34: 1135 - 1154. https: // doi. org / 10.1080 / 00222930050020140","Glaw F. & Vences M. 2007. A Field Guide to the Amphibians and Reptiles of Madagascar, 3 rd ed. Vences and Glaw Verlag, Cologne.","Lehtinen R. M., Glaw F. & Vences M. 2011. Two new plant-breeding frog species (Anura: Mantellidae, Guibemantis) from southeastern Madagascar. Herpetological Journal 21: 95 - 112.","Vieites D. R., Wollenberg K. C., Andreone F., Kohler J., Glaw F. & Vences M. 2009. Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Science of the USA 106: 8267 - 8272. https: // doi. org / 10.1073 / pnas. 0810821106","Perl R. G. B., Nagy Z. T., Sonet G., Glaw F., Wollenberg K. C. & Vences M. 2014. DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia 35: 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Lehtinen R. M., Nussbaum R. A., Richards C. M., Cannatella D. C. & Vences M. 2007. Mitochondrial genes reveal cryptic diversity in plant-breeding frogs from Madagascar (Anura, Mantellidae, Guibemantis). Molecular Phylogenetics and Evolution 44: 1121 - 1129. https: // doi. org / 10.1016 / j. ympev. 2007.05.020"]}

Rhombophryne guentherpetersi (Guib��, 1974) Common name: Tsaratanana saw-browed diamond frog (modified from Frank & Ramus 1995) (Figs 4���7, S1���S2) Mantipus guentherpetersi Guib��, 1974: 1181 ���1182 Plethodontohyla guentherpetersi ��� Blommers-Schl��sser & Blanc, 1991: 56 ���57. First depicted in life under this name in Stuart et al. (2008). Rhombophryne guentherpetersi ��� Glaw & Vences, 2007: 118. The photo provided on page 119 of this book is misattributed and depicts an undescribed species (Scherz et al. unpubl. data). Holotype. MNHN 1953.165, an adult female specimen captured by an unknown collector on an unknown date on the Tsaratanana Massif at 2600 m a.s.l. (Figs 4, 5, S2) Paratypes. MNHN 1953.165A and MNHN 1953.165B, a juvenile and adult female with the same collection data as the holotype. MNHN 1973.592 and MNHN 1973.593, a subadult female and juvenile collected by Charles P. Blanc in 1966 on Mission ORSTOM from the same location as the holotype. Referred specimens. ZSM 606/2014 (DRV 6220), a female, ZSM 607/2014 (DRV 6223), a male, and ZSM 608/2014 (DRV 6231), a female (Fig. S1), three adult specimens collected by D. Vieites, M. Vences, R.D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, F. Randrianasolo, F. Randrianasolo and T. Rajoafiarison, ZSM 606 /2014 and 607/2014 on the 16th of June 2010 at Andranomadio (camp 4), Tsaratanana, 14.0801��S, 48.9854��E, 2503 m a. s.l.; ZSM 608 /2014 on the 18th of June 2010 at a site on the Tsaratanana mountain 14.0665��S, 48.9832��E, 2732 m a.s.l. A further three specimens from Andranomadio (with same collecting data as ZSM specimens from this site) were deposited in the UADBA collection: UADBA-A 60775 (DRV 6210) and UADBA-A 60776 (ZCMV 12401), adults; UADBA-A 60782 (ZCMV 12435), subadult. Diagnosis and comparisons. Rhombophryne guentherpetersi is a member of the genus Rhombophryne on the basis of molecular phylogenetic affinities (Fig. 3) and the possession of a clavicle combined with the absence of T- or Y-shaped terminal phalanges (vs. either absence of a clavicle or possession of a clavicle combined with T- or Yshaped terminal phalanges in the morphologically similar Plethodontohyla Boulenger, 1882). Within the genus Rhombophryne, it is assigned to the R. serratopalpebrosa species group on the basis of possessing superciliary spines and of molecular phylogenetic affinities (Fig. 3, Table 1). Rhombophryne guentherpetersi is distinguished from all other described congeners by the following unique suite of characters: adult SVL 27.3���35.7 mm; TDH/ED = 0.4 5���0.66; a weak supratympanic fold running from the rear corner of the eye to curve slightly over and behind the tympanum toward the axilla; two or three small superciliary spines; distinct, raised dorsolateral glands, and bulbous tibial glands. The pectoral girdle is distinctly narrower than that seen in its congeners (see Fig. 7; 8.2���9.4% of SVL). Furthermore, R. guentherpetersi is separated from all other Rhombophryne species except R. ornata and R. serratopalpebrosa by uncorrected pairwise distances of at least 4.4% in a segment of the 16S rRNA mitochondrial gene (see Table 1; no molecular data available for R. serratopalpebrosa)���for distinction from R. ornata, see below. Rhombophryne guentherpetersi is distinguished from all Rhombophryne species except the R. serratopalpebrosa group by the possession of superciliary spines, and from all members of this group, and indeed all cophyline microhylids, by the possession of bulbous tibial glands. Genetically, R. guentherpetersi is very closely related to R. ornata ���they are separated by just 2.4���3.4% in one fragment of the 16S gene (see Table 1), which is lower than our standard threshold for candidate species recognition (Vieites et al. 2009). However, the two species are highly morphologically distinct, differing not just in the presence or absence of tibial glands, but also in the shorter relative hindlimb length of R. guentherpetersi (HIL/SVL 1.33���1.45 vs. 1.46���1.64), tibiotarsal articulation reaching the axilla or tympanum (vs. between the tympanum and the eye), absence (vs. presence) of red colouration in the inguinal region, partially ossified limb epiphyses and carpals (vs. unossified), and much narrower pectoral girdle (length of coracoid 12%; see Fig. 7). A differential diagnosis from all other cophyline microhylids is unnecessary because the tibial glands allow easy unambiguous distinction. These are noticeable, if not well developed, even in subadults. Re-description of the holotype. (Figs 4, 5) An adult female specimen in moderately good state of preservation. Ventral incision and left lateral incision present. Body robust. Head wider than long (HW/HL = 1.35). Pupils horizontally oval. Snout rounded in dorsal and lateral views. Canthus rostralis concave. Loreal region concave. Nostrils nearer to snout tip than to eye (END/NSD = 1.24), directed laterally, slightly protuberant. Tympanum distinct, TDH/ED = 0.47. Weak supratympanic fold, from middle rear of eye curving slightly over and behind tympanum toward the axilla. Two small superciliary spines above each eye. Vomerine teeth present, curved, not meeting at the midline. Arms robust. Fingers without webbing; relative lengths 1Variation. The type series is composed of two juveniles, one subadult, and one poorly preserved adult, in addition to the holotype. Our newly added material consists of three well-preserved adults, including the first male specimen. All specimens, including the juveniles, possess superciliary spines, but they are small and variable in number, and in poorly preserved specimens cannot be well distinguished (but can be distinguished in life; see Fig. 6). ZSM 606/2014 has three superciliary spines, whereas all other specimens have two. The paratype MNHN 1953.165B is darker than the rest of the type series and has much rougher skin (but is also particularly poorly preserved). All specimens, including the juveniles, have tibial glands, but these are less bulbous in the juveniles and subadult, becoming large only in the adults (confirming that all members of the type series are indeed conspecifics). The one male specimen is smaller than the adult females, but is in all other aspects highly similar in morphology (see Appendix 1). The tibiotarsal articulation reaches either the axilla or the tympanum. A prepollex is well developed in the one available male specimen, ZSM 607/2014. The species may have some degree of sexual size dimorphism, as the only male specimen is slightly smaller than all adult female specimens available (27.3 vs. 28.9���35.7 mm). Colouration is relatively homogeneous, but shade and pattern differ somewhat (Fig. 6). The newly collected specimens are much darker brown in colour than the type series, but generally have commonalities to their colour patterns: a pair of blackish oblong markings above the suprascapular region (Fig. 6). These markings are present only in one member of the type series (MNHN 1973.592). The lateral body has several dark-brown flecks. These are present in all of the paratypes but not in the holotype. The venter is mottled dark and light brown in some specimens but is almost solid dark brown in others. A light brown interocular bar is sometimes present, anterior to which the snout is typically lighter brown than the dorsum. The dorsolateral glands can also be this light brown, or they can be continuous with the rest of the dorsum. Remarks. The following inconsistencies exist between our re-description and the original description of this species by Guib�� (1974): Superciliary spines are present; these were apparently overlooked in his description. Tibiotarsal articulation was given as reaching the eye. This could not be verified in any member of the type series except the poorly preserved paratype MNHN 1973.592, the vertebral column of which is inverted (i.e. convex instead of concave), possibly misleading tibiotarsal articulation. The holotype is 10 mm longer than described (34.6 mm vs. 25 mm). Guib�� (1974, 1978) stated that the prepollex was developed in males, but the type series does not contain any males. Nevertheless, we have confirmed this observation with a newly collected male specimen (see Variation above). Natural history. At Andranomadio, specimens were found during the day in high-altitude rainforest with rather open canopy (2503 m a.s.l.), burrowed several centimetres deep in the ground. Another individual (ZSM 608/2014) was collected above the treeline (2732 m a.s.l.), in an area of grassland with some scattered heath, hidden under a stone during the day. Distribution and conservation status. Rhombophryne guentherpetersi is currently listed in the IUCN Red List as Endangered under criterion B1ab(iii) (IUCN SSC Amphibian Specialist Group 2016a), meaning an extent of occurrence of R. guentherpetersi is known only from the Tsaratanana Massif. The exact collecting locality of the type specimens is not clear (Guib�� 1974), but three specimens were collected on a recent expedition to the Tsaratanana Massif (ZSM 606���607/2014 at 14.0801��S, 48.9854��E at 2503 m a.s.l; ZSM 608/2014 at 14.0665��S, 48.9832��E, 2732 m a.s.l.; Fig. 8). These two localities are well inside the protected area of Tsaratanana Strict Nature Reserve, to which only researchers and conservation workers are permitted access. Although it is possible that the species may occur in the forest corridor between Marojejy and Tsaratanana (COMATSA; see Rabearivony et al. 2015), which is pending official protected status, we consider this unlikely: we suspect that R. guentherpetersi is found only at high elevations, possibly near to and above the tree line, which in Tsaratanana is around 2550 m a.s.l. No part of the COMATSA corridor exceeds 2300 m a.s.l. The likely extent of occurrence of this species is therefore limited to an area of 36.4 km 2. Deforestation and habitat degradation pressure at such high altitude is minimal relative to the lower reaches of the forest, especially due to the strict conservation status of this forest. However, it is not free from threats: Climate change may be a significant factor for species living near the tops of mountains, as their niche space diminishes with rising temperatures. Fires in the high mountain grassland could swiftly eradicate the suitable open habitat and leave only forest habitat available. To balance (1) the small range of this species, (2) the highly protected status it enjoys, and (3) the threats that persist despite this protection, we propose to maintain a status of Endangered B1ab(iii) (IUCN 2012), but encourage further surveys within COMATSA and Tsaratanana Strict Nature Reserve in order to better understand the distribution and ecology of this species., Published as part of Scherz, Mark D., Hawlitschek, Oliver, Andreone, Franco, Rakotoarison, Andolalao, Vences, Miguel & Glaw, Frank, 2017, 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, pp. 301-340 in Zootaxa 4273 (3) on pages 307-312, DOI: 10.11646/zootaxa.4273.3.1, http://zenodo.org/record/803049, {"references":["Guibe, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Museum national d'Histoire Naturelle, 3, 1170 - 1192.","Frank, N. & Ramus, E. (1995) Complete Guide to Scientific and Common Names of Amphibians and Reptiles of the World. N. G. Publishing Inc., Pottsville, Pennsylvania, 377 pp.","Blommers-Schlosser, R. M. A. & Blanc, C. P. (1991) Amphibiens (premiere partie). Faune de Madagascar, 75, 1 - 397.","Stuart, S. N., Hoffmann, M., Chanson, J. S., Cox, N. A., Berridge, R. J., Ramani, P. & Young, B. E. (2008) Threatened Amphibians of the World. Lynx Edicions, Barcelona; IUCN, Gland & Conservation International, Arlington, Virginia, 758 pp.","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Koln, 496 pp.","Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Caudata in the collection of the British Museum. 2 nd Edition. Taylor and Francis, London, xvi + 503 pp.","Vieites, D. R., Wollenberg, K. C., Andreone, F., Kohler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the United States of America, 106, 8267 - 8272.","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","Rabearivony, J., Rasamoelina, M., Raveloson, J., Rakotomanana, H., Raselimanana, A. P., Raminosoa, N. R. & Zaonarivelo, J. R. (2015) Roles of a forest corridor between Marojejy, Anjanaharibe-Sud and Tsaratanana protected areas, northern Madagascar, in maintaining endemic and threatened Malagasy taxa. Madagascar Conservation & Development, 10, 85 - 92."]}