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Type: Article
Published: 2022-12-15
Page range: 113–311
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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

Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, 2100, Copenhagen Ø, Denmark
CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485-661 Vairão, Portugal, Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, 4099-002 Porto, Portugal, BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485-661 Vairão, Portugal
Museum of Natural Sciences and Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, USA
Hauptstr. 13, 82234 Weßling, Germany
Museo Regionale di Scienze Naturali, Via G. Giolitti, 36, 10123 Torino, Italy
Natural and Environmental Sciences, Regional University Centre of the SAVA Region (CURSA), Antalaha, Madagascar, Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar
Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, D-60325, Frankfurt, Germany
Institute of Zoology, Ecology and Conservation, Biocentre Grindel, University of Hamburg, Martin-Luther-King-Platz 3, 29146 Hamburg, Germany
Institut de Systématique, Evolution, Biodiversité, UMR 7205 CNRS, MNHN, Sorbonne Université, EPHE, Université des Antilles, Muséum National d'Histoire Naturelle, CP 51, 57 rue Cuvier, 75231 PARIS Cedex 05 France
Institute for Biochemistry and Biology, University of Potsdam, Karl-Liebknecht-Str. 24–25, 14476 Potsdam, Germany
Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar, School for International Training, VN 41A Bis Ankazolava Ambohitsoa, Antananarivo, 101 Madagascar
Zoologisches Institut, Technische Universität Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany, Uppsala University, Department of Ecology and Genetics, Animal Ecology, Norbyvägen 18 D, 752 36 Uppsala, Sweden
Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar, Association Vahatra, Lot V A 38 LBA Ter Ambohidempona Tsiadana, BP 3972, Antananarivo, 101 Madagascar
Institute of Zoology, Ecology and Conservation, Biocentre Grindel, University of Hamburg, Martin-Luther-King-Platz 3, 29146 Hamburg, Germany
Museum für Naturkunde—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany
Institute of Zoology, Zoological Society of London, London NW1 4RY, UK, Centre for Ecology, Evolution and Environmental Changes (cE3c), Faculdade de Ciências, Universidade de Lisboa, 1749-016 Lisboa, Portugal
Department of Life Sciences, The Natural History Museum, Cromwell Road, London, UK
Department of Biogeography and Global Change, Museo Nacional de Ciencias Naturales—CSIC. Calle José Gutierrez Abascal 2, 28006, Madrid, Spain
Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany
Institute for Biochemistry and Biology, University of Potsdam, Karl-Liebknecht-Str. 24–25, 14476 Potsdam, Germany
Zoologische Staatssammlung München (ZSM-SNSB), Münchhausenstr. 21, 81247 München, Germany
Technische Universität Braunschweig, Zoological Institute, Mendelssohnstr. 4, 38106 Braunschweig
Amphibia Anura Mantellidae Madagascar FrogCap target enrichment museomics museum genomics phylogenomics ASAP integrative taxonomy


Malagasy frogs of the subgenus Brygoomantis in the mantellid frog genus Mantidactylus currently comprise 14 described species of mostly brown, riparian frogs. Data from DNA barcoding suggested that the diversity of this subgenus is dramatically underestimated by current taxonomy. We here provide a comprehensive revision of this subgenus. We use hybrid-enrichment based DNA barcode fishing to obtain mitochondrial DNA fragments from the name-bearing type material of 16 of the 20 available names for members of this subgenus, and integrate these into a genetic dataset consisting of 1305 individuals sampled across Madagascar. By thus assigning the nomina to genetic lineages, we can confidently establish synonyms, revalidate old names, and describe the remaining diversity. We take an integrative approach to our descriptions, drawing together genetics, morphometrics and morphology, and bioacoustics for assignment. We also provide a robust phylogenomic hypothesis for the subgenus, based on 12,951 nuclear-encoded markers (almost 10 million base pairs) for 58 representative samples, sequenced using a hybrid-enrichment bait set for amphibians. Those data suggest a division of the subgenus into eight major clades and shows that morphological species complexes are often paraphyletic or polyphyletic. Lectotypes are designated for Rana betsileana Boulenger, 1882; Rana biporus Boulenger, 1889; Rana curta Boulenger, 1882; Mantidactylus ambohimitombi Boulenger, 1918; Mantidactylus tripunctatus Angel, 1930; and Rana inaudax Peracca, 1893. For several other nomina, previous authors had considered a certain syntype as holotype; this has been seen as lectotype designation by implication, which, however, is ambiguous according to the provisions of the International Code of Zoological Nomenclature. Hence, we validate a previous lectotype designation by implication for Limnodytes ulcerosus Boettger, 1880 by explicitly designating the same individual as lectotype. In one other such case, that of Mantidactylus brauni Ahl, 1929, we deviate from previous authors and designate a different specimen as lectotype. We revalidate Rana inaudax Peracca, 1893 as Mantidactylus inaudax (Peracca, 1893) bona species, and Mantidactylus tripunctatus Angel, 1930 bona species. The identities of three further species (M. ambohimitombi, M. biporus, M. tricinctus) are largely redefined based on new genetic data. By designating the lectotype of Rana aluta (MZUT An725.1) as the neotype of Mantidactylus laevis Angel, 1929 we also stabilize the latter nomen (as junior synonym of M. alutus) whose original type material is lost. Based on DNA sequences of its lectotype, we consider Mantidactylus brauni Ahl, 1929 as junior synonym of M. ulcerosus (rather than M. biporus). We formally name 20 new species and four new subspecies: M. ambohimitombi marefo ssp. nov., M. ambohimitombi miloko ssp. nov., M. mahery sp. nov., M. steinfartzi sp. nov., M. incognitus sp. nov., M. jonasi sp. nov., M. katae sp. nov., M. kortei sp. nov., M. riparius sp. nov., M. fergusoni sp. nov., M. georgei sp. nov., M. jahnarum sp. nov., M. marintsoai sp. nov., M. grubenmanni sp. nov., M. gudrunae sp. nov., M. augustini sp. nov., M. bletzae sp. nov., M. brevirostris sp. nov., M. eulenbergeri sp. nov., M. glosi sp. nov., M. stelliger sp. nov., M. manerana sp. nov., M. manerana fotaka ssp. nov., and M. manerana antsanga ssp. nov. Based on our taxonomic revision, we discuss (i) the importance of definitive assignment of historical names via archival DNA analysis; (ii) the relevance of the subspecies category to name geographic variation within species; (iii) the value of molecular characters in formal species diagnoses in taxa with substantial individual variation of morphology; (iv) the use of phylogenomic approaches for taxonomy, by confirming that some morphologically similar taxa are not each other’s closest relatives, and in several cases belong to entirely different major subclades within Brygoomantis, thus facilitating lineage diagnosis; and (v) the need to interpret genetic distances in a probabilistic framework rather than using fixed thresholds, where higher distances confer a higher likelihood of genetic incompatibilities across the genome and thus completion of speciation.


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