Published July 22, 2026 | Version v1

A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species

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Abstract

The diamond frogs of the genus Rhombophryne are a poorly understood clade of morphologically diverse frogs native to the humid forests of Madagascar. We here present a monographic revision of the taxonomy and natural history of these frogs. We first provide morphological diagnoses and contents of the established species groups in Rhombophryne. We then integrate molecular phylogenetic data from mitochondrial and nuclear genetic markers, with morphometrics, bioacoustics, and micro-CT-based osteology, to describe seven new species from across Madagascar. This brings the number of species in the genus to 27. We have obtained DNA sequences from name-bearing specimens of 23 of these species, plus topotypical specimens of two further species; we used museomics methods to sequence name-bearing types of three species collected before 1976, as well as several other specimens of interest, allowing us to clarify taxonomic assignment of those individuals. In our treatments of all existing and new species, we provide new insights into their anatomy, diet, ecology, and distribution. We provide the first published key to the genus, which we hope will make their identification more straightforward. Finally, we suggest a revised conservation assessment in light of the data that we present. We estimate 24 of the 27 species to be threatened, with two species (R. kilonjy sp. nov. and R. matavy) probably worthy of being classified as 'Critically Endangered'. Several candidate species remain taxonomically unaddressed, due to the poor representation of specimens in museum collections. We therefore emphasise the need for increased survey efforts to fill knowledge gaps in both the taxonomy and the natural history of these frogs.

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References

  • Andreone F (2003) Distribution record. Plethodontohyla coudreaui. Herpetological Review 34: 259.
  • Andreone F, Glaw F, Mattioli F, Jesu R, Schimmenti G, Randrianirina JE, Vences M (2009) The peculiar herpetofauna of some Tsaratanana rainforests and its affinities with Manongarivo and other massifs and forests of northern Madagascar. Italian Journal of Zoology 76: 92–110. https://doi.org/10.1080/11250000802088603
  • Andreone F, Glaw F, Nussbaum RA, Raxworthy CJ, Vences M, Randrianirina JE (2003) The amphibians and reptiles of Nosy Be (NW Madagascar) and nearby islands: A case study of diversity and conservation of an insular fauna. Journal of Natural History 37: 2119–2149. https://doi.org/10.1080/00222930210130357
  • Andreone F, Randrianirina JE (2008) An unexpected Rhombophryne record at Tsingy de Bemaraha confirms the presence of cophyline frogs in western Madagascar. Zootaxa 1812: 46–48.
  • Andreone F, Randrianirina JE, Jenkins PD, Aprea G (2000) Species diverstiy of Amphibia, Reptilia and Lipotyphla (Mammalia) at Ambolokopatrika, a rainforest between the Anjanaharibe-Sud and Marojejy massifs, NE Madagascar. Biodiversity and Conservation 9: 1587–1622.
  • Andreone F, Vences M, Randrianirina JE (2001) Patterns of amphibian and reptile diversity at Berara Forest (Sahamalaza Peninsula), NW Madagascar. Italian Journal of Zoology 68: 235–241. https://doi.org/10.1080/11250000109356414
  • Andreone F, Vences M, Vieites DR, Glaw F, Meyer A (2005) Recurrent ecological adaptations revealed through a molecular analysis of the secretive cophyline frogs of Madagascar. Molecular Phylogenetics and Evolution 34: 315–322. https://doi.org/10.1016/j.ympev.2004.10.013
  • Andrianarimisa A, Andrianjakarivelo V, Rakotomalala Z, Anjeriniaina M (2009) Vertébrés terrestres des fragments forestiers de la Montagne d'Ambatotsirongorongo, site dans le Système des Aires Protégées de Madagascar de la Région Anosy, Tolagnaro. Malagasy Nature 2: 30–51.
  • Angel F (1938) Description d'un Amphibien nouveau, de Madagascar, appartenant au genre Plethodontohyla. Bulletin du Muséum National d'Histoire Naturelle (2e Série) 10: 260–261.
  • Antonelli A, Smith RJ, Perrigo AL, Crottini A, Hackel J, Testo W, Farooq H, Torres Jiménez MF, Andela N, Andermann T, Andriamanohera AM, Andriambololonera S, Bachman SP, Bacon CD, Baker WJ, Belluardo F, Birkinshaw C, Borrell JS, Cable S, Canales NA, Carrillo JD, Clegg R, Clubbe C, Cooke RSC, Damasco G, Dhanda S, Edler D, Faurby S, de Lima Ferreira P, Fisher BL, Forest F, Gardiner LM, Goodman SM, Grace OM, Guedes TB, Henniges MC, Hill R, Lehmann CER, Lowry PP, Marline L, Matos-Maraví P, Moat J, Neves B, Nogueira MGC, Onstein RE, Papadopulos AST, Perez-Escobar OA, Phelps LN, Phillipson PB, Pironon S, Przelomska NAS, Rabarimanarivo M, Rabehevitra D, Raharimampionona J, Rajaonah MT, Rajaonary F, Rajaovelona LR, Rakotoarinivo M, Rakotoarisoa AA, Rakotoarisoa SE, Rakotomalala HN, Rakotonasolo F, Ralaiveloarisoa BA, Ramirez-Herranz M, Randriamamonjy JEN, Randriamboavonjy T, Randrianasolo V, Rasolohery A, Ratsifandrihamanana AN, Ravololomanana N, Razafiniary V, Razanajatovo H, Razanatsoa E, Rivers M, Sayol F, Silvestro D, Vorontsova MS, Walker K, Walker BE, Wilkin P, Williams J, Ziegler T, Zizka A, Ralimanana H (2022) Madagascar's extraordinary biodiversity: Evolution, distribution, and use. Science 378: eabf0869. https://doi.org/10.1126/science.abf0869
  • Aprea G, Odierna G, Andreone F, Glaw F, Vences M (2007) Karyological evolution and systematics of Malagasy microhylid frogs. Zoologischer Anzeiger 246: 23–41. https://doi.org/10.1016/j.jcz.2006.10.001
  • Barbour T, Loveridge A (1929) Typical reptiles and amphibians in the Museum of Comparative Zoölogy. Bulletin of the Museum of Comparative Zoology 69: 205–360.
  • Bauer AM, Günther R, Robeck HE (1996) An annotated type catalogue of the hemisotid, microhylid, myobatrachid, pelobatid and pipid frogs in the Zoological Museum, Berlin. Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe 72: 259–275. https://doi.org/10.1002/mmnz.19960720205
  • Bellati A, Scherz MD, Megson S, Hyde Roberts S, Andreone F, Rosa GM, Noël J, Randrianirina JE, Fasola M, Glaw F, Crottini A (2018) Resurrection and re-description of Plethodontohyla laevis (Boettger, 1913) and transfer of Rhombophryne alluaudi (Mocquard, 1901) to the genus Plethodontohyla (Microhylidae: Cophylinae). Zoosystematics and Evolution 94: 109–135. https://doi.org/10.3897/zse.94.14698
  • Belluardo F, Scherz MD, Santos B, Andreone F, Antonelli A, Glaw F, Muñoz-Pajares J, Randrianirina JE, Raselimanana AP, Vences M, Crottini A (2022) Molecular taxonomic identification and species-level phylogeny of the narrow-mouthed frogs of the genus Rhombophryne (Anura: Microhylidae: Cophylinae) from Madagascar. Systematics and Biodiversity 20: 1–13. https://doi.org/10.1080/14772000.2022.2039320
  • Blommers-Schlösser RMA (1975) Observations on the larval development of some Malagasy frogs, with notes on their ecology and biology (Anura: Dyscophinae, Scaphiophryninae, and Cophylinae). Beaufortia 24: 7–26.
  • Blommers-Schlösser RMA (1976) Chromosomal analysis of twelve species of Microhylidae (Anura) from Madagascar. Genetica 46: 199–210.
  • Blommers-Schlösser RMA, Blanc CP (1991) Amphibiens (première partie). Faune de Madagascar 75: 1–397.
  • Boettger O (1880a) Diagnoses Batrachiorum novorum insulae Madagascar. Zoologischer Anzeiger 3: 567–568.
  • Boettger O (1880b) Diagnoses reptilium et batrachiorum novorum a Carolo Ebenau in insula Nossi-Bé madagascariensi lectorum. Zoologischer Anzeiger 3: 279–283.
  • Boettger O (1881) Diagnoses reptilium et batrachiorum novorum ab ill. Antonio Stumpff in insula Nossi-Bé Madagascariensi lectorum. Zoologischer Anzeiger 4: 358–362.
  • Boulenger GA (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London, 503 pp. https://doi.org/10.5962/bhl.title.20903
  • Brown JL, Sillero N, Glaw F, Bora P, Vieites DR, Vences M (2016) Spatial biodiversity patterns of Madagascar's amphibians and reptiles. PLoS One 11: e0144076. https://doi.org/10.1371/journal.pone.0144076
  • Bruford MW, Hanotte O, Brookefield JFY, Burke T (1992) Single-locus and multilocus DNA fingerprint. In: Hoelzel AR (Ed.) Molecular Genetic Analysis of Populations: A Practical Approach. IRL Press, Oxford, 225–270.
  • Carné A, Vieites DR (2024) A race against extinction: The challenge to overcome the Linnean amphibian shortfall in tropical biodiversity hotspots. Diversity and Distributions 30: e13912. https://doi.org/10.1111/ddi.13912
  • Carné A, Vieites DR, Ferrer X, Moreno M, ponz-Segrelles G, Rahagalala TNAA, Rakotomanga SE, Laca AR, Ratsoavina FM, van den Burg MP (2025) Microendemism in Madagascar: Small ranges or sampling gaps? The case of the frog Wakea madinika. Salamandra 61: 160–170.
  • Clement M, Posada DC, Crandall KA (2000) TCS: A computer program to estimate gene genealogies. Molecular Ecology 9: 1657–1659.
  • D'Cruze N, Köhler J, Franzen M, Glaw F (2008) A conservation assessment of the amphibians and reptiles of the Forêt d'Ambre Special Reserve, north Madagascar. Madagascar Conservation & Development 3: 44–54.
  • D'Cruze N, Köhler J, Vences M, Glaw F (2010) A new fat fossorial frog (Microhylidae: Cophylinae: Rhombophryne) from the rainforest of the Forêt d'Ambre Special Reserve, northern Madagascar. Herpetologica 66: 182–191. https://doi.org/10.1655/09-008r1.1
  • Dabney J, Knapp M, Glock I, Gansauge M, Weihmann A, Nickel B, Valdioserad C, García N, Pääbo S, Arsuag J, Meyer M (2013) Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments. Proceedings of the National Academy of Sciences of the USA 110: 15758–15763.
  • de Queiroz K (1998) The General Lineage Concept of species, species criteria, and the process of speciation. In: Howard DJ, Berlocher SH (Eds) Endless Forms: Species and Speciation. Oxford University Press, Oxford, 57–75.
  • de Queiroz K (2007) Species concepts and species delimitation. Systematic Biology 56: 879–886. https://doi.org/10.1080/10635150701701083
  • Edmonds D, Rakotoarisoa JC, Dolch R, Pramuk J, Gagliardo R, Andreone F, Rabibisoa N, Rabemananjara F, Rabesihanaka S, Robsomanitrandrasana E (2012) Building capacity to implement conservation breeding programs for frogs in Madagascar: Results from year one of Mitsinjo's amphibian husbandry research and captive breeding facility. Amphibian & Reptile Conservation 5: 57–69
  • Eudeline R, Wang C, Scherz MD (2015) Predation attempt of Rhombophryne laevipes by Compsophis albiventris. Herpetology Notes 8: 393–394.
  • Fedosov A, Achaz G, Gontchar A, Puillandre N (2022) mold, a novel software to compile accurate and reliable DNA diagnoses for taxonomic descriptions. Molecular Ecology Resources 22: 2038–2053. https://doi.org/10.1111/1755-0998.13590
  • Fouquet A, Gilles A, Vences M, Marty C, Blanc M, Gemmell NJ (2007) Underestimation of species richness in neotropical frogs revealed by mtDNA analyses. PLoS One 2: e1109. https://doi.org/10.1371/journal.pone.0001109
  • Frappier-Brinton T, Lehman SM (2022) The burning island: Spatiotemporal patterns of fire occurrence in Madagascar. PLoS One 17: e0263313. https://doi.org/10.1371/journal.pone.0263313
  • Frost DR (2026) Amphibian Species of the World: An Online Reference. Version 6.2 Electronic Database accessible at https://amphibiansoftheworld.amnh.org/. American Museum of Natural History, New York, NY [accessed 26 June 2026].
  • Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CFB, de Sá RO, Channing A, Wilkinson M, Donnellan SC, Raxworthy CJ, Campbell JA, Blotto BL, Moler P, Drewes RC, Nussbaum RA, Lynch JD, Green DM, Wheeler WC (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370. https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
  • Fulton TL, Shapiro B (2019) Setting up an ancient DNA laboratory. In: Shapiro B, Barlow A, Heintzman PD, Hofreiter M, Paijmans JLA, Soares AER (Eds) Ancient DNA: Methods and Protocols. Springer New York, New York, NY, 1–13. https://doi.org/10.1007/978-1-4939-9176-1_1
  • Gansauge M-T, Gerber T, Glocke I, Korlević P, Lippik L, Nagel S, Riehl LM, Schmidt A, Meyer M (2017) Single-stranded DNA library preparation from highly degraded DNA using T4 DNA ligase. Nucleic Acids Research 45: e79. https://doi.org/10.1093/nar/gkx033
  • Garg S, Biju SD (2019) New microhylid frog genus from Peninsular India with Southeast Asian affinity suggests multiple Cenozoic biotic exchanges between India and Eurasia. Scientific Reports 9: 1906. https://doi.org/10.1038/s41598-018-38133-x
  • Garg S, Senevirathne G, Wijayathilaka N, Phuge S, Deuti K, Manamendra-Arachchi K, Meegaskumbura M, Biju SD (2018) An integrative taxonomic review of the South Asian microhylid genus Uperodon. Zootaxa 4384: 1–88. https://doi.org/10.11646/zootaxa.4384.1.1
  • Garg S, Sivaperuman C, Gogulakrishnan G, Chandramouli SR, Biju SD (2022) Hiding in plain sight: Rain water puddles in Nicobar Islands of India reveal abundance of a new frog species of the genus Microhyla Tschudi, 1838 (Anura: Microhylidae). Zoological Studies 61: 1–23. https://doi.org/10.6620/ZS.2022.61-02
  • Garg S, Suyesh R, Das A, Jiang J, Wijayathilaka N, Amarasinghe AAT, Alhadi F, Vineeth KK, Aravind NA, Senevirathne G, Meegaskumbura M, Biju SD (2019) Systematic revision of Microhyla (Microhylidae) frogs of South Asia: A molecular, morphological, and acoustic assessment. Vertebrate Zoology 69: 1–71. https://doi.org/10.26049/VZ69-1-2019-01
  • Glaw F, Crottini A, Rakotoarison A, Scherz MD, Vences M (2022) Diversity and exploration of the Malagasy amphibian fauna. In: Goodman SM (Ed.) The New Natural History of Madagascar. Princeton University Press, Princeton, NJ, 1305–1322.
  • Glaw F, Köhler J, de la Riva I, Vieites DR, Vences M (2010a) Integrative taxonomy of Malagasy treefrogs: Combination of molecular genetics, bioacoustics and comparative morphology reveals twelve additional species of Boophis. Zootaxa 2383: 1–82.
  • Glaw F, Köhler J, Vences M (2010b) A new fossorial frog, genus Rhombophryne, from Nosy Mangabe Special Reserve, Madagascar. Zoosystematics and Evolution 86: 235–243. https://doi.org/10.1002/zoos.201000006
  • Glaw F, Scherz MD, Rakotoarison A, Crottini A, Raselimanana AP, Andreone F, Köhler J, Vences M (2020) Genetic variability and partial integrative revision of Platypelis frogs (Microhylidae) with red flash marks from eastern Madagascar. Vertebrate Zoology 70: 141–156. https://doi.org/10.26049/VZ70-2-2020-04
  • Glaw F, Vences M (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, 335 pp.
  • Glaw F, Vences M (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, 478 pp.
  • Glaw F, Vences M (2002) Rhombophryne testudo. Correction of locality record. Madagascar: Réunion. Herpetological Review 33: 64.
  • Glaw F, Vences M (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. Vences & Glaw Verlags GbR, Cologne, 496 pp.
  • Goodman SM, Raherilalao MJ, Wohlhauser S (Eds) (2018) Les Aires Protégées Terrestres de Madagascar: Leur Histoire, Description et Biote/The Terrestrial Protected Areas of Madagascar: Their History, Description, and Biota. Vol. II: Northern and Eastern Madagascar. Association Vahatra, Antananarivo, 806 pp.
  • Gouveia CM, Silva M, Russo A (2024) The severity of dry and hot climate extremes and their related impacts on vegetation in Madagascar. iScience 27. https://doi.org/10.1016/j.isci.2023.108658
  • Guibé J (1946) Sur la validité de Mantipus laevipes (Mocq.) (Batraciens-Microhylidae). Bulletin du Muséum National d'Histoire Naturelle (2e Série) 18: 50–51.
  • Guibé J (1973) Batraciens nouveaux de Madagascar. Bulletin du Muséum National d'Histoire Naturelle (3e Série) 171: 1169–1192.
  • Guibé J (1975) Batraciens nouveaux de Madagascar. Bulletin du Muséum National d'Histoire Naturelle (3e Série) 323: 1081–1089.
  • Guibé J (1978) Les batraciens de Madagascar. Bonner zoologische Monographien 11: 1–140.
  • Günther ACLG (1858) On the systematic arrangement of the tailless batrachians and the structure of Rhinophrynus dorsalis. Proceedings of the Zoological Society of London 1858: 339–352.
  • Hahn EE, Alexander M, Stiller J, Grewe PM, Holleley CE (2024) Hot alkaline lysis gDNA extraction from formalin-fixed archival tissues. PLoS One 19: e0296491. https://doi.org/10.1371/journal.pone.0296491
  • Hall TA (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
  • Hernández-Palma TL, Rueda-Solano LA, Valkonen JK, Rojas B (2023) Predator response to the coloured eyespots and defensive posture of Colombian four-eyed frogs. Journal of Evolutionary Biology 36: 1040–1049. https://doi.org/10.1111/jeb.14193
  • Howes GB, Ridewood W (1888) On the carpus and tarsus of the Anura. Proceedings of the Zoological Society of London 1888: 141–182. https://doi.org/10.1111/j.1469-7998.1888.tb06691.x
  • Hutter CR, Andriampenomanana ZF, Razafindraibe J, Rakotoarison A, Scherz MD (2018) New dietary data from Compsophis and Alluaudina species (Squamata: Lamprophiidae: Pseudoxyrhophiinae), and implications for their dietary complexity and evolution. Journal of Natural History 52: 2497–2510. https://doi.org/10.1080/00222933.2018.1543732
  • Hutter CR, Cobb KA, Portik DM, Travers SL, Wood Jr PL, Brown RM (2021) FrogCap: A modular sequence capture probe-set for phylogenomics and population genetics for all frogs, assessed across multiple phylogenetic scales. Molecular Ecology Resources 22: 1100–1119. https://doi.org/10.1111/1755-0998.13517
  • Hyde Roberts S, Daly C (2014) A rapid herpetofaunal assessment of Nosy Komba Island, northwestern Madagascar, with new locality records for seventeen species. Salamandra 50: 18–26.
  • IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. IUCN, Gland and Cambridge, 38 pp.
  • IUCN SSC Amphibian Specialist Group (2016a) Boophis ankarafensis. IUCN Red List of Threatened Species 2016: e.T68568328A68568331. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T68568328A68568331.en
  • IUCN SSC Amphibian Specialist Group (2016b) Gephyromantis ambohitra. IUCN Red List of Threatened Species 2016: e.T57458A84168942. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T57458A84168942.en
  • IUCN SSC Amphibian Specialist Group (2016c) Rhombophryne coronata. IUCN Red List of Threatened Species 2016: e.T57969A84180747. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T57969A84180747.en
  • IUCN SSC Amphibian Specialist Group (2016d) Rhombophryne coudreaui. IUCN Red List of Threatened Species 2016: e.T57970A84180885. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T57970A84180885.en
  • Joseph GS, Seymour CL, Rakotoarivelo AR (2024) Fire incongruities can explain widespread landscape degradation in Madagascar's forests and grasslands. Plants, People, Planet 6: 656–669. https://doi.org/10.1002/ppp3.10471
  • Kalyaanamoorthy S, Minh BQ, Wong TK, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587. https://doi.org/10.1038/nmeth.4285
  • Keefe R, Blackburn DC (2020) Comparative morphology of the humerus inforward-burrowing frogs. Biological Journal of the Linnean Society 20: 1–13. https://doi.org/10.1093/biolinnean/blaa092
  • Klages J, Glaw F, Köhler J, Müller J, Hipsley CA, Vences M (2013) Molecular, morphological and osteological differentiation of a new species of microhylid frog of the genus Stumpffia from northwestern Madagascar. Zootaxa 3717: 280–300. https://doi.org/10.11646/zootaxa.3717.2.8
  • Köhler J, Glaw F, Vences M (1997) Notes on the reproduction of Rhombophryne (Anura: Microhylidae) at Nosy Be, northern Madagascar. Revue français d'Aquariologie- Herpetologie 24: 53–54.
  • Köhler J, Jansen M, Rodríguez A, Kok PJR, Toledo LF, Emmrich M, Glaw F, Haddad CFB, Rödel M-O, Vences M (2017) The use of bioacoustics in anuran taxonomy: Theory, terminology, methods and recommendations for best practice. Zootaxa 4251: 1–124. https://doi.org/10.11646/zootaxa.4251.1.1
  • Lambert SM, Hutter CR, Scherz MD (2017) Diamond in the rough: A new species of fossorial diamond frog (Rhombophryne) from Ranomafana National Park, southeastern Madagascar. Zoosystematics and Evolution 93: 143–155. https://doi.org/10.3897/zse.93.10188
  • Laurence JD (1987) Rev. Richard Baron's "Compendium des Plantes Malgaches". Taxon 36: 39–46. https://doi.org/10.2307/1221349
  • Laurent RF (1946) Mises au point dans la taxonomie des ranides. Revue de Zoologie et de Botanique Africaines 39: 336–338.
  • Lê S, Josse J, Husson F (2008) FactoMineR: An R package for multivariate analysis. Journal of Statistical Software 25: 1–18. https://doi.org/10.18637/jss.v025.i01
  • Mertens R (1967) Die herpetologische Sektion des Natur-Museums und Forschungs-Institutes Senckenberg in Frankfurt a. M. nebst einem Verseichnis ihrer Typen. Senckenbergiana Biologica 48: 1–106.
  • Mocquard MF (1895) Sur une collection de reptiles recueillis a Madagascar par MM. Alluaud et Belly. Bulletin de la Société Philomathique de Paris (Huitième Série) 7: 112–136.
  • Mocquard MF (1901) Note préliminaire sur une collection de Reptiles et de Batraciens recuellis par M. Aluaud dans le sud de Madagascar. Bulletin du Muséum National d'Histoire naturelle 7: 251–256.
  • Mocquard MF (1909) Synopsis des familles, genres et espèces des reptiles écailleux et des batraciens de Madagascar. Nouvelles Archives du Muséum (5e Série) 1: 1–110.
  • Müller F (1892) Siebenter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel 10: 195–215.
  • Nguyen L-T, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. https://doi.org/10.1093/molbev/msu300
  • Noble GK (1931) The Biology of the Amphibia. McGraw-Hill, New York, NY & London, 577 pp.
  • Padial JM, Miralles A, De La Riva I, Vences M (2010) The integrative future of taxonomy. Frontiers in Zoology 7: 16. https://doi.org/10.1186/1742-9994-7-16
  • Paijmans JLA, Baleka S, Henneberger K, Taron UH, Trinks A, Westbury MV, Barlow A (2017) Sequencing single-stranded libraries on the Illumina NextSeq 500 platform. arXiv. https://doi.org/10.48550/arXiv.1711.11004
  • Palumbi SR, Martin A, Romano S, McMillan WO, Stice L, Grabowski G (1991) The Simple Fool's Guide to PCR, Version 2.0. Privately published, University of Hawaii, Honolulu, HI, 45 pp.
  • Parker HW (1934) Monograph of the Frogs of the Family Microhylidae. British Museum, London, 208 pp.
  • Peloso PLV, Frost DR, Richards SJ, Rodrigues MT, Donnellan S, Matsui M, Raxworthy CJ, Biju SD, Lemmon EM, Lemmon AR, Wheeler WC (2016) The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in narrow-mouthed frogs (Anura, Microhylidae). Cladistics 32: 113–140. https://doi.org/10.1111/cla.12118
  • Peloso PLV, Raxworthy CJ, Wheeler WC, Frost DR (2017) Nomenclatural stability does not justify recognition of paraphyletic taxa: A response to Scherz et al. (2016). Molecular Phylogenetics and Evolution 111: 56–64. https://doi.org/10.1016/j.ympev.2017.03.016
  • Penny SG, Andreone F, Crottini A, Holderied MW, Rakotozafy LS, Schwitzer C, Rosa GM (2014) A new species of the Boophis rappiodes group (Anura, Mantellidae) from the Sahamalaza Peninsula, northwest Madagascar, with acoustic monitoring of its nocturnal calling activity. Zookeys: 111–132. https://doi.org/10.3897/zookeys.435.7383
  • Penny SG, Andreone F, Crottini A, Holderied MW, Rosa GM, Schwitzer C (2016) The Amphibians of the Sahamalaza Peninsula, Northwest Madagascar — Actions for their Conservation. Bristol Zoological Society, Bristol, 34 pp.
  • Penny SG, Crottini A, Andreone F, Bellati A, Rakotozafy LMS, Holderied MW, Schwitzer C, Rosa GM (2017) Combining old and new evidence to increase the known biodiversity value of the Sahamalaza Peninsula, Northwest Madagascar. Contributions to Zoology 86: 273–296.
  • Perl RGB, Nagy ZT, Sonet G, Glaw F, Wollenberg KC, Vences M (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia 35: 197–206. https://doi.org/10.1163/15685381-00002942
  • Peters WCH (1883) Über Mantipus und Phrynocara, zwei neue Batrachiergattungen aus dem Hinterlasse des Reisenden J. M. Hildebrandt von Madagascar. Sitzungsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin 1883: 165–168.
  • Petzold A, Glaw F, Mullin KE, Rakotoarison A, Raselimanana AP, Crottini A, Orozco-terWengel P, Köhler J, Prötzel D, Vences M, Hofreiter M, Scherz MD (2025) A preliminary assessment of the diversity of the frog genus Anilany from Madagascar (Microhylidae, Cophylinae) with description of a new species from western Madagascar. Salamandra 61: 70–94.
  • Phelps LN, Andela N, Gravey M, Davis DS, Kull CA, Douglass K, Lehmann CER (2022) Madagascar's fire regimes challenge global assumptions about landscape degradation. Global Change Biology 28: 6944–6960. https://doi.org/10.1111/gcb.16206
  • Phillips CA (2015) Herpetological type specimens in the University of Illinois Museum of Natural History: An update to the primary types. https://www.inhs.illinois.edu/files/8914/3110/1369/Prim._Types2015.pdf
  • Posit team (2024) RStudio: Integrated Development Environment for R. Posit Software, PBC, Boston, MA.
  • Puillandre N, Brouillet S, Achaz G (2021) ASAP: Assemble species by automatic partitioning. Molecular Ecology Resources 21: 609–620. https://doi.org/10.1111/1755-0998.13281
  • R Core Team (2024) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.
  • Rabearivony J, Raselimanana AP, Andriamazava MA, Thorstrom R, de Roland L-AR (2010) A new locality for the endangered microhylid frog Scaphiophryne boribory from northern Madagascar and a rapid survey of other amphibians of the Bemanevika region. Herpetology Notes 3: 105–109.
  • Raherilalao MJ, Raselimanana AP, Soarimalala V, Goodman S (2016) Evaluation biologique de la flore et de la faune du Parc National de Masoala, Madagascar. Association Vahatra, Antananarivo, 116 pp.
  • Rakotoarimalala F, Raselimanana AP (2023) Aperçu global de la tendance de la structure de la communauté herpétofaunique le long du gradient altitudinal du versant Est du Parc National de Marojejy au cours de ces 25 dernières années. Malagasy Nature 17: 165–186.
  • Rakotoarisoa AJ (2012) Analyse comparative des communautes herpetofauniques de la foret d'Ambatovy-Analamay, en vue de concevoir une approche de conservation convenable. Université d'Antananarivo, Antananarivo, 50 pp.
  • Rakotoarisoa JC, Rakotoarison A, Rasoanantenaina S, Robsomanitrandrasana E, Edmonds SSS, Soamiarimampionona J, Tsimialomanana E, Wolf S, Edmonds D (2025) Captive breeding reveals insights into the ecology and reproductive biology of 11 little-known Malagasy frog species. Zoo Biology 44: 24–35. https://doi.org/10.1002/zoo.21876
  • Rakotoarison A, Crottini A, Müller J, Rödel M-O, Glaw F, Vences M (2015) Revision and phylogeny of narrow-mouthed treefrogs (Cophyla) from northern Madagascar: Integration of molecular, osteological, and bioacoustic data reveals three new species. Zootaxa 3937: 61–89. https://doi.org/10.11646/zootaxa.3937.1.3
  • Rakotoarison A, Glaw F, Vieites DR, Raminosoa NR, Vences M (2012) Taxonomy and natural history of arboreal microhylid frogs (Platypelis) from the Tsaratanana Massif in northern Madagascar, with description of a new species. Zootaxa 3563: 1–25.
  • Rakotoarison A, Scherz MD, Bletz MC, Razafindraibe JH, Glaw F, Vences M (2019) Diversity, elevational variation, and phylogenetic origin of stump-toed frogs (Microhylidae: Cophylinae: Stumpffia) on the Marojejy massif, northern Madagascar. Salamandra 55: 115–123.
  • Rakotoarison A, Scherz MD, Glaw F, Köhler J, Andreone F, Franzen M, Glos J, Hawlitschek O, Jono T, Mori A, Ndriantsoa SH, Raminosoa Rasoamampionona N, Riemann JC, Rödel M-O, Rosa GM, Vieites DR, Crottini A, Vences M (2017) Describing the smaller majority: Integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar. Vertebrate Zoology 67: 271–398. https://doi.org/10.3897/vz.67.e31595
  • Rakotomalala D (2002) Diversité des reptiles et amphibiens de la Réserve Spéciale de Manongarivo, Madagascar. In: Gautier L, Goodman SM (Eds) Inventaire Floristique et Faunistique de la Réserve Spéciale de Manongarivo (NW Madagascar). Conservatoire et Jardin Botaniques de Genève, Geneva, 340–358.
  • Ralimanana H, Perrigo AL, Smith RJ, Borrell JS, Faurby S, Rajaonah MT, Randriamboavonjy T, Vorontsova MS, Cooke RSC, Phelps LN, Sayol F, Andela N, Andermann T, Andriamanohera AM, Andriambololonera S, Bachman SP, Bacon CD, Baker WJ, Belluardo F, Birkinshaw C, Cable S, Canales NA, Carrillo JD, Clegg R, Clubbe C, Crottini A, Damasco G, Dhanda S, Edler D, Farooq H, de Lima Ferreira P, Fisher BL, Forest F, Gardiner LM, Goodman SM, Grace OM, Guedes TB, Hackel J, Henniges MC, Hill R, Lehmann CER, Lowry PP, Marline L, Matos-Maraví P, Moat J, Neves B, Nogueira MGC, Onstein RE, Papadopulos AST, Perez-Escobar OA, Phillipson PB, Pironon S, Przelomska NAS, Rabarimanarivo M, Rabehevitra D, Raharimampionona J, Rajaonary F, Rajaovelona LR, Rakotoarinivo M, Rakotoarisoa AA, Rakotoarisoa SE, Rakotomalala HN, Rakotonasolo F, Ralaiveloarisoa BA, Ramirez-Herranz M, Randriamamonjy JEN, Randrianasolo V, Rasolohery A, Ratsifandrihamanana AN, Ravololomanana N, Razafiniary V, Razanajatovo H, Razanatsoa E, Rivers M, Silvestro D, Testo W, Torres Jiménez MF, Walker K, Walker BE, Wilkin P, Williams J, Ziegler T, Zizka A, Antonelli A Madagascar's extraordinary biodiversity: Threats and opportunities. Science 378: eadf1466. https://doi.org/10.1126/science.adf1466
  • Ramanamanjato J-B, McIntyre PB, Nussbaum RA (2002) Reptile, amphibian, and lemur diversity of the Malahelo Forest, a biogeographical transition zone in southeastern Madagascar. Biodiversity and Conservation 11: 1791–1807.
  • Ramanamanjato J-B, Soanary CH (2011) The herpetofauna of Tsitongambarika Forest. In: BirdLie International (Ed.) Tsitongambarika Forest, Madagascar—Biological and socio-economic surveys, with conservation recommendations. BirdLife International, Cambridge, 34–41.
  • Raselimanana AP (1999) L'herpétofaune. In: Goodman SM, Rasolonandrasana BPN (Eds) Inventaire Biologique de la Réserve Spéciale du Pic d'Ivohibe et du Couloir Forestier qui la Relie au Parc National d'Andringitra. Centre d'Information et de Documentation scientifique et technique, Antananarivo, 81–97.
  • Raselimanana AP (2010) The amphibians and reptiles of the Ambatovy-Analamay region. Malagasy Nature 3: 99–123.
  • Raselimanana AP, Raxworthy CJ, Nussbaum RA (2000) Herpetofaunal species diversity and elevational distribution within the Parc National de Marojejy, Mad[a]gascar. Fieldiana Zoology (New Series) 97: 157–174.
  • Ratsimbazafy J, Ralison JM, Rabarison H, Randrianarison RM, Mensimana RM, Rasolondraibe LT, Rasoarivelo DS, Randrianasolo RM, Rakotosamimanana TH, Andrianandrasanana H (2008) Préservation de la biodiversité de Maromizaha. Antananarivo, 109 pp.
  • Raxworthy CJ, Nussbaum RA (1994) A rainforest survey of amphibians, reptiles and small mammals at Montagne d'Ambre, Madagascar. Biological Conservation 69: 65–73.
  • Raxworthy CJ, Nussbaum RA (1996) Amphibians and reptiles of the Réserve Naturelle Intégrale d'Andringitra, Madagascar: A study of elevational distribution and local endemicity. Fieldiana Zoology 85: 158–170.
  • Raxworthy CJ, Pearson RG, Rabibisoa N, Rakotondrazafy AM, Ramanamanjato J-B, Raselimanana AP, Wu S, Nussbaum RA, Stone DA (2008) Extinction vulnerability of tropical montane endemism from warming and upslope displacement: A preliminary appraisal for the highest massif in Madagascar. Global Change Biology 14: 1703–1720. https://doi.org/10.1111/j.1365-2486.2008.01596.x
  • Rohland N, Siedel H, Hofreiter M (2004) Nondestructive DNA extraction method for mitochondrial DNA analyses of museum specimens. BioTechniques 36: 814–821.
  • Rosa GM, Andreone F, Crottini A, Hauswaldt JS, Noël J, Rabibisoa NH, Randriambahiniarime MO, Rebelo R, Raxworthy CJ (2012) The amphibians of the relict Betampona low-elevation rainforest, eastern Madagascar: An application of the integrative taxonomy approach to biodiversity assessments. Biodiversity and Conservation 21: 1531–1559. https://doi.org/10.1007/s10531-012-0262-x
  • Scherz MD (2020) Diamond frogs forever: A new species of Rhombophryne Boettger, 1880 (Microhylidae, Cophylinae) from Montagne d'Ambre National Park, northern Madagascar. Zoosystematics and Evolution 96: 313–323. https://doi.org/10.3897/zse.96.51372
  • Scherz MD, Crottini A, Hutter CR, Hildenbrand A, Andreone F, Fulgence TR, Köhler G, Ndriantsoa SH, Ohler A, Preick M, Rakotoarison A, Rancilhac L, Raselimanana AP, Riemann JC, Rödel M-O, Rosa GM, Streicher JW, Vieites DR, Köhler J, Hofreiter M, Glaw F, Vences M (2022a) An inordinate fondness for inconspicuous brown frogs: integration of phylogenomics, archival DNA analysis, morphology, and bioacoustics yields 24 new taxa in the subgenus Brygoomantis (genus Mantidactylus) from Madagascar. Megataxa 7: 113–311. https://doi.org/10.11646/megataxa.7.2.1
  • Scherz MD, Crottini A, Rakotoarison A (2022b) Microhylidae: Cophylinae, Microhylid frogs. In: Goodman SM (Ed.) The New Natural History of Madagascar. Princeton University Press, Princeton, NJ, 1382–1390.
  • Scherz MD, Glaw F, Hutter CR, Bletz MC, Rakotoarison A, Köhler J, Vences M (2019a) Species complexes and the importance of Data Deficient classification in Red List assessments: The case of Hylobatrachus frogs. PLoS One 14: e0219437. https://doi.org/10.1371/journal.pone.0219437
  • Scherz MD, Glaw F, Vences M, Andreone F, Crottini A (2016a) Two new species of terrestrial microhylid frogs (Microhylidae: Cophylinae: Rhombophryne) from northeastern Madagascar. Salamandra 52: 91–106.
  • Scherz MD, Hawlitschek O, Andreone F, Rakotoarison A, Vences M, Glaw F (2017a) 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 4273: 301–340. https://doi.org/10.11646/zootaxa.4273.3.1
  • Scherz MD, Hutter CR, Rakotoarison A, Riemann JC, Rödel M-O, Ndriantsoa SH, Glos J, Hyde Roberts S, Crottini A, Vences M, Glaw F (2019b) Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera. PLoS One 14: e0213314. https://doi.org/10.1371/journal.pone.0213314
  • Scherz MD, Rakotoarison A, Hawlitschek O, Vences M, Glaw F (2015a) Leaping towards a saltatorial lifestyle? An unusually long-legged new species of Rhombophryne (Anura, Microhylidae) from the Sorata massif in northern Madagascar. Zoosystematics and Evolution 91: 105–114. https://doi.org/10.3897/zse.91.4979
  • Scherz MD, Ruthensteiner B, Vences M, Glaw F (2014) A new microhylid frog, genus Rhombophryne, from northeastern Madagascar, and a re-description of R. serratopalpebrosa using micro-computed tomography. Zootaxa 3860: 547–560. https://doi.org/10.11646/zootaxa.3860.6.3
  • Scherz MD, Ruthensteiner B, Vieites DR, Vences M, Glaw F (2015b) Two new microhylid frogs of the genus Rhombophryne with superciliary spines from the Tsaratanana Massif in northern Madagascar. Herpetologica 71: 310–321. https://doi.org/10.1655/HERPETOLOGICA-D-14-00048
  • Scherz MD, Vences M, Rakotoarison A, Andreone F, Köhler J, Glaw F, Crottini A (2016b) Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae). Molecular Phylogenetics and Evolution 100: 372–381. https://doi.org/10.1016/j.ympev.2016.04.019
  • Scherz MD, Vences M, Rakotoarison A, Andreone F, Köhler J, Glaw F, Crottini A (2017b) 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 53: 479–483.
  • Stephens M, Smith NJ, Donnelly P (2001) A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics 68: 978–989.
  • Straube N, Lyra ML, Paijmans JLA, Preick M, Basler N, Penner J, Rödel M-O, Westbury MV, Haddad CF, Barlow A, Hofreiter M (2021) Successful application of ancient DNA extraction and library construction protocols to museum wet collection specimens. Molecular Ecology Resources 21: 2299–2315. https://doi.org/10.1111/1755-0998.13433
  • Trueb L (1968) Cranial osteology of the hylid frog, Smilisca baudini. University of Kansas Publications, Museum of Natural History 18: 11–35.
  • Trueb L (1973) Bones, frogs, and evolution. In: Vial JL (Ed.) Evolutionary Biology of the Anurans: Contemporary Research on Major Problems. University of Missouri Press, Columbia, MO, 65–132.
  • Trueb L (1993) Patterns of diversity in the lissamphibian skull. In: Hanken J, Hall BK (Eds) The Skull, Volume 2: Patterns of Structural and Systematic Diversity. University of Chicago Press, Chicago, IL, 255–343.
  • Tu N, Yang M, Liang D, Zhang P (2018) A large-scale phylogeny of Microhylidae inferred from a combined dataset of 121 genes and 427 taxa. Molecular Phylogenetics and Evolution 126: 85–91.
  • Vallan D, Andreone F, Raherisoa VH, Dolch R (2004) Does selective wood exploitation affect amphibian diversity? The case of An'Ala, a tropical rainforest in eastern Madagascar. Oryx 38: 410–417. https://doi.org/10.1017/S003060530400078X
  • van der Meijden A, Vences M, Hoegg S, Boistel R, Channing A, Meyer A (2007) Nuclear gene phylogeny of narrow-mouthed toads (Family: Microhylidae) and a discussion of competing hypotheses concerning their biogeographical origins. Molecular Phylogenetics and Evolution 44: 1017–1030. https://doi.org/10.1016/j.ympev.2007.02.008
  • van der Meijden A, Vences M, Hoegg S, Meyer A (2005) A previously unrecognized radiation of ranid frogs in southern Africa revealed by nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 37: 674–685.
  • Veith M, Lötters S, Andreone F, Rödel M-O (2004) Measuring and monitoring amphibian diversity in tropical forests. II. Estimating species richness from standardized transect censing. Ecotropica 10: 85–99.
  • Venables WN, Ripley BD (2002) Modern Applied Statistics with S. Springer, New York, NY, 498 pp.
  • Vences M, Glaw F (2003) New microhylid frog (Plethodontohyla) with a supraocular crest from Madagascar. Copeia 2003: 789–793. https://doi.org/10.1643/ha02-285.1
  • Vences M, Glaw F, Marquez R (Eds) (2006) The Calls of the Frogs of Madagascar. 3 Audio CDs and booklet. Vol. Fonoteca Zoológica, Madrid, 44 pp.
  • Vences M, Köhler J, Vieites DR, Glaw F (2011) Molecular and bioacoustic differentiation of deep conspecific lineages of the Malagasy treefrogs Boophis tampoka and B. luteus. Herpetology Notes 4: 239–246.
  • Vences M, Kosuch J, Glaw F, Böhme W, Veith M (2003a) Molecular phylogeny of hyperoliid treefrogs: Biogeographic origin of Malagasy and Seychellean taxa and re-analysis of familial paraphyly. Journal of Zoological Systematics and Evolutionary Research 41: 205–215. https://doi.org/10.1046/j.1439-0469.2003.00205.x
  • Vences M, Miralles A, Brouillet S, Ducasse J, Fedosov A, Kharchev V, Kostadinov I, Kumari S, Patmanidis S, Scherz MD, Puillandre N, Renner SS (2021) iTaxoTools 0.1: Kickstarting a specimen-based software toolkit for taxonomists. Megataxa 6: 77–92. https://doi.org/10.11646/megataxa.6.2.1
  • Vences M, Multzsch M, Gippner S, Miralles A, Crottini A, Gehring P-S, Rakotoarison A, Ratsoavina FM, Glaw F, Scherz MD (2022) Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar's rainforest. Zootaxa 5179: 1–61. https://doi.org/10.11646/zootaxa.5179.1.1
  • Vences M, Patmanidis S, Fodosov A, Miralles A, Puillandre N (2024a) iTaxoTools 1.0: Improved DNA Barcode Exploration with TaxI2. In: DeSalle R (Ed.) DNA Barcoding Methods in Molecular Biology, vol 2744. Humana, New York, NY, https://doi.org/10.1007/978-1-0716-3581-0_18
  • Vences M, Patmanidis S, Schmidt J-C, Matschiner M, Miralles A, Renner SS (2024b) Hapsolutely: A user-friendly tool integrating haplotype phasing, network construction, and haploweb calculation. Bioinformatics Advances 4: vbae083. https://doi.org/10.1093/bioadv/vbae083
  • Vences M, Raxworthy CJ, Nussbaum RA, Glaw F (2003b) New microhylid frog (Plethodontohyla) from Madagascar, with semiarboreal habits and possible parental care. Journal of Herpetology 37: 629–636. https://doi.org/10.1670/258-01A
  • Vieites DR, Wollenberg KC, Andreone F, Köhler 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 USA 106: 8267–8272. https://doi.org/10.1073/pnas.0810821106
  • Wickham H (2016) ggplot2: Elegant Graphics for Data Analysis. Springer, New York, NY, 260 pp.
  • Wickham H, Averick M, Bryan J, Chang W, D'Agostino McGowan L, François R, Grolemund G, Hayes A, Henry L, Hester J, Kuhn M, Pedersen TL, Miller E, Milton Bache S, Müller K, Ooms J, Robinson D, Seidel DP, Spinu V, Takahashi K, Vaughan D, Wilke C, Woo K, Yutani H (2019) Welcome to the Tidyverse. The Journal of Open Source Software 4: 1686. https://doi.org/10.21105/joss.01686
  • Wollenberg KC, Vieites DR, Glaw F, Vences M (2011) Speciation in little: The role of range and body size in the diversification of Malagasy mantellid frogs. BMC Evolutionary Biology 11: 217. https://doi.org/10.1186/1471-2148-11-217
  • Wollenberg KC, Vieites DR, van der Meijden A, Glaw F, Cannatella DC, Vences M (2008) Patterns of endemism and species richness in Malagasy cophyline frogs support a key role of mountainous areas for seciation. Evolution 62: 1890–1907. https://doi.org/10.1111/j.1558-5646.2008.00420.x