Abstract
We describe a new species of arboreal microhylid frog from northern Madagascar. The new species is assigned to the genus Cophyla based on molecular phylogeny and morphological similarities to other species of this genus. Molecular phylogenetic analysis resolved Cophyla fortuna sp. nov. as closely related to C. noromalalae. However, the two species were reciprocally monophyletic both in a tree reconstructed from a combination of mitochondrial genes, and in a tree based on the nuclear RAG-1 gene. The new species, previously identified as candidate species Cophyla sp. Ca4, occurs in lowland bamboo forests around the Marojejy Massif in northeastern Madagascar. It differs from the allopatric C. noromalalae, so far only known from the Montagne d’Ambre Massif in northern Madagascar, by slightly smaller body size and shorter duration and higher spectral frequency of advertisement calls. Several additional genetically divergent mitochondrial lineages of Cophyla, related to the C. fortuna/noromalalae complex and to C. phyllodactyla, occur in intervening areas between Montagne d’Ambre and Marojejy, and their status requires further study.
References
Andreone, F., Vences, M., Vieites, D.R., 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
Bletz, M.C., Scherz, M.D., Rakotoarison, A., Lehtinen, R.M., Glaw, F. & Vences, M. (2018) Stumbling upon a new frog species of Guibemantis (Anura: Mantellidae) on top of the Marojejy Massif in northern Madagascar. Copeia, 106, 255–263.
https://doi.org/10.1643/CH-17-655
Blommers-Schlösser, R.M.A. (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, R.M.A. & Blanc, C.P. (1991) Amphibiens (première partie). Faune de Madagascar, 75, 1–379.
Boettger, O. (1880) Diagnoses Reptilium et Batrachiorum novorum a Carolo Ebenau in insula Nossi-Bé Madagascariensi lectorum. Zoologischer Anzeiger, 3, 279–283.
Boumans, L., Vieites, D.R., Glaw, F. & Vences, M. (2007) Geographical patterns of deep mitochondrial differentiation in widespread Malagasy reptiles. Molecular Phylogenetics and Evolution, 45, 822–839.
https://doi.org/10.1016/j.ympev.2007.05.028
Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Single-locus and multilocus DNA fingerprinting. In: Hoelzel, A.R. (Ed.), Molecular genetic analysis of populations: a practical approach. IRL Press, Oxford, pp. 225–270.
Frost, D.R. (2018) Amphibian Species of the World: an Online Reference. Version 6.0. Available from: http://research.amnh.org/herpetology/amphibia/index.html (accessed 20 April 2019)
Glaw, F., Köhler, J. & Vences, M. (2012) A tiny new species of Platypelis from the Marojejy National Park in northeastern Madagascar (Amphibia: Microhylidae). European Journal of Taxonomy, 9, 1–9.
https://doi.org/10.5852/ejt.2012.9
Glaw, F. & Vences, M. (1992) A fieldguide to the amphibians and reptiles of Madagascar. – Vences & Glaw Verlag, Köln, 331 pp.
Glaw, F. & Vences, M. (1994) A fieldguide to the amphibians and reptiles of Madagascar, 2nd edition. – Vences & Glaw Verlag, Köln, 480 pp.
Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. 3rd Edition. Vences & Glaw Verlags GbR, Köln, 496 pp. [Germany]
Köhler, J., Jansen, M., Rodríguez, A., Kok, P.J.R., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F.B., Rödel, M.-O. & Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa, 4251 (1), 1–124.
https://doi.org/10.11646/zootaxa.4251.1.1
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874.
https://doi.org/10.1093/molbev/msw054
Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34, 772–773.
https://doi.org/10.1093/molbev/msw260
Peloso, P.L.V., Frost, D.R., Richards, S.J., Rodrigues, M.T., Donnellan, S., Matsui, M., Raxworthy, C.J., Biju, S.D., Lemmon, E.M., Lemmon, A.R. & Wheeler, W.C. (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, P.L.V., Raxworthy, C.J., Wheeler, W.C. & Frost, D.R. (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
R Development Core Team (2015) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: http://www.R-project.org (accessed 20 April 2019)
Rakotoarison, A., Glaw, F., Vieites, D.R., Raminosoa, N.R. & 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), 1–25.
https://doi.org/10.11646/zootaxa3563.1.1
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 (1), 61–89.
https://doi.org/10.11646/zootaxa.3937.1.3
Rakotoarison, A., Glaw, F., Vieites, D.R., Raminosoa, N.R. & 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), 1–25.
https://doi.org/10.11646/zootaxa3563.1.1
Rakotoarison, A, Scherz, M.D., Glaw, F., Köhler, J., Andreone, F., Franzen, M., Glos, J., Hawlitschek, O., Jono, T., Mori, A., Ndriantsoa, S.H., Raminosa, N.R., Riemann, J.C., Rödel, M.-O., Rosa, G.M., Vieites, D.R., 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.
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180
Scherz, M.D., Hawlitschek, O., Razafindraibe, J.H., Megson, S., Ratsoavina, F.M., Rakotoarison, A., Bletz, M.C., Glaw, F. & Vences, M. (2018) A distinctive new frog species (Anura, Mantellidae) supports the biogeographic linkage of two montane rainforest massifs in northern Madagascar. Zoosystematics and Evolution, 94, 247–261.
https://doi.org/10.3897/zse.94.21037
Scherz, M.D., 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 (6), 547–560. https://doi.org/10.11646/zootaxa.3860.6.3
Scherz, M.D., 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, M.D., 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, M.D., Razafindraibe, J.H., Rakotoarison, A., Dixit, N.M., Bletz, M.C., Glaw, F. & Vences, M. (2017a) Yet another small brown frog from high elevation on the Marojejy Massif, northeastern Madagascar (Anura: Mantellidae). Zootaxa, 4347 (3), 572–482.
https://doi.org/10.11646/zootaxa.4347.3.9
Scherz, M.D., 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.
Sueur, J., Aubin, T. & Simonis, C. (2008) Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics, 18, 213–226.
https://doi.org/10.1080/09524622.2008.9753600
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.
https://doi.org/10.1016/j.ympev.2018.03.036
Vences, M., Andreone, F.& Glaw, F.(2005) A new microhylid frog of the genus Cophyla from a transitional forest in northwestern Madagascar. African Zoology, 40, 143–149.
Vences, M., Kosuch, J., Glaw, F., Böhme, W. & Veith, M. (2003) 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
Vieites, D.R., Wollenberg, K.C., 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 United States of America, 106, 8267–8272.
https://doi.org/10.1073/pnas.0810821106
Wohltmann, A., du Preez, L., Rödel, M.-O., Köhler, J. & Vences, M. (2007) Endoparasitic mites of the genus Endotrombicula Ewing, 1931 (Acari: Prostigmata: Parasitengona: Trombiculidae) from African and Madagascan anurans, with description of a new species. Folia Parasitologica, 54, 225–235.
https://doi.org/10.14411/fp.2007.031
Wollenberg, K.C., Vieites, D.R., van der Meijden, A., Glaw, F., Cannatella, D.C. & Vences, M. (2008) Patterns of endemism and species richness in Malagasy cophyline frogs support a key role of mountainous areas for speciation. Evolution, 62, 1890–1907.
https://doi.org/10.1111/j.1558-5646.2008.00420.x