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Type: Monograph
Published: 2022-09-01
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Integrative revision of the Lygodactylus madagascariensis group reveals an unexpected diversity of little brown geckos in Madagascar’s rainforest

Zoologisches Institut, Technische Universität Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany
Zoologisches Institut, Technische Universität Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany
Zoologisches Institut, Technische Universität Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany
Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire Naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, Paris, France
CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485-661 Vairão, Portugal 4Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, 4099-002 Porto, Portugal 5BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485-661 Vairão, Portugal
Faculty of Biology / Biologiedidaktik, University Bielefeld, Universitätsstraße 25, 33615 Bielefeld, Germany
Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar 8School for International Training, VN 41A Bis Ankazolava Ambohitsoa, Antananarivo, 101 Madagascar
Mention Zoologie et Biodiversité Animale, Université d’Antananarivo, BP 906, Antananarivo, 101 Madagascar
Zoologische Staatssammlung München (ZSM-SNSB), Münchhausenstr. 21, 81247 München, Germany
Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, 2100, Copenhagen Ø, Denmark
Amphibia Squamata Gekkonidae Domerguella dwarf geckos cryptic species integrative taxonomy


The Lygodactylus madagascariensis species group, constituting the subgenus Domerguella, currently contains five valid species of inconspicuous dwarf geckos from Madagascar’s humid forests, but at least 18 deep genetic lineages have been revealed by recent molecular studies. Given the high morphological similarity of these lineages, taxonomic resolution of this astonishing diversity requires efforts to correctly delimit species, as well as assigning the available nomina to the species-level lineages identified. We here combine DNA sequences of one mitochondrial and two nuclear-encoded gene fragments with morphometric measurements and scale counts, and report evidence for a species status of most of the previously identified lineages. In particular, we rely on sympatric and often even syntopic occurrence of several of these lineages without evidence for genetic admixture, and consistent with subtle morphological differences. Furthermore, the very high divergences of 7.4–23.8% pairwise distances in the relatively conserved mitochondrial 16S rRNA gene, combined with a lack of haplotype sharing in the nuclear-encoded genes and differences in scale counts convinced us that most of the other, allopatrically distributed lineages also represent distinct species. We elevate L. madagascariensis petteri to species level and formally name eight new species: L. salvi sp. nov., a species from the Sambirano region in northern Madagascar, previously called L. sp. 8; L. tantsaha sp. nov. (L. sp. 10), a species occurring sympatrically with L. madagascariensis and L. petteri on Montagne d’Ambre in far northern Madagascar; L. roellae sp. nov. (L. sp. 17), a species characterized by a striped coloration in all known specimens, from northern Madagascar; L. winki sp. nov. (L. sp. 18), an unstriped species from northern Madagascar but belonging to a subclade mostly distributed in the eastern rainforests of the island; L. ulli sp. nov. (L. sp. 21), a species from the same subclade as L. winki but known only from the Marojejy Massif in the North East; L. fritzi sp. nov. (L. sp. 11), a further species of this subclade from coastal lowlands in the Northern Central East; L. hodikazo sp. nov. (L. sp. 23) known from a single specimen collected at the Tsingy de Bemaraha and therefore the only Domerguella species known from the West region of Madagascar; and L. hapei sp. nov. (L. sp. 26), an enigmatic species from the Sambirano region characterized by a striped pattern on the throat that is otherwise unknown in the subgenus. Three additional deep mitochondrial lineages of Domerguella were identified in our analysis, but could not be further analyzed due to the lack or scarcity of voucher specimens. More field work and collection of voucher specimens is needed to understand their status. Furthermore, the taxonomy of the Domerguella subclade occurring in eastern Madagascar, with three described species (L. guibei, L. miops, L. fritzi), two synonyms (L. septemtuberculatus, Microscalabotes spinulifer) and at least two further deep genetic lineages co-occurring in a relatively small area, requires further revisionary work, possibly aided by target-enrichment sequencing of the respective name-bearing types.



  1. Ahrens, D., Fujisawa, T., Krammer, H.J., Eberle, J., Fabrizi, S. & Vogler, A.P. (2016) Rarity and incomplete sampling in DNA-based species delimitation. Systematic Biology, 65, 478–494.
    Angel, F. (1942) Les lézards de Madagascar. Mémoires de l’Académie Malgache, 36, 1–139.
    Avise, J.C. & Ball, R.M. (1990) Principles of genealogical concordance in species concepts and biological taxonomy. In: Futuyma, D. & Antonovics, J. (Eds.), Oxford surveys in evolutionary biology. Oxford Univ. Press, Oxford, UK, pp. 45–67.
    Avise, J. & Wollenberg, K. (1997) Phylogenetics and the origin of species. Proceedings of the National Academy of Sciences of the United States of America, 94, 7748–7755.
    Blommers-Schlösser, R.M.A. & Blanc, C.P. (1991) Amphibiens (première partie). Faune de Madagascar, 75 (1), 1–379.
    Boettger, O. (1881) Diagnoses reptilium et batrachiorum novorum ab ill. Antonio Stumpff in insula Nossi-Bé Madagascariensis lectorum. Zoologischer Anzeiger, 4, 358–362.
    Boettger, O. (1913) Reptilien und Amphibien von Madagascar, den Inseln und dem Festland Ostafrikas. Voeltzkow, A., Reise in Ostafrika, 3 (4), 1–269.
    Bollen, A. & Donati, G. (2005) Phenology of the littoral forest of Sainte Luce, southeastern Madagascar. Biotropica, 37, 32–43
    Bora, P., Randrianantoandro, J.C., Randrianavelona, R., Hantalalaina, E.F., Andriantsimanarilafy, R.R., Rakotondravony, D., Ramilijaona, O.R., Vences, M., Jenkins, R.K.B., Glaw, F. & Köhler, J. (2010) Amphibians and reptiles of the Tsingy de Bemaraha Plateau, Western Madagascar: Checklist, biogeography and conservation. Herpetological Conservation and Biology, 5, 111–125.
    Bott, R. (1965) Die Süsswasserkrabben von Madagaskar (Crustacea, Decapoda). Bulletin du Muséum national d’Histoire naturelle, series 2, 37, 335–350.
    Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Printed by order of the Trustees London, 450 pp.
    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.
    Brown, J.L., Sillero, N., Glaw, F., Bora, P., Vieites, D.R. & Vences, M. (2016) Spatial biodiversity patterns of Madagascar’s amphibians and reptiles. PLoS One, 11, e0144076.
    Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Single-locus and multilocus DNA fingerprint. In: Hoelzel, A.R. (Ed.), Molecular Genetic Analysis of Populations: A Practical Approach. IRL Press, Oxford, pp. 225–270.
    Chan, K.O., Hutter, C.R., Wood, P.L., Grismer, L.L., Das, I. & Brown, R.M. (2020) Gene flow creates a mirage of cryptic species in a Southeast Asian spotted stream frog complex. Molecular Ecology, 29, 3970–3987.
    Colwell, R.K. & Lees, D.C. (2000) The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology and Evolution, 15, 70–76.
    Crottini A., Gehring, P.-.S., Glaw, F., Harris, D.J., Lima, A. & Vences, M. (2011) Deciphering the cryptic species diversity of dull-coloured day gekos Phelsuma (Squamata: Gekkonidae) from Madagascar, with description of a new species. Zootaxa, 2982, 40–48.
    Crottini, A., Harris, D.J., Miralles, A., Glaw, F., Jenkins, R.K.B., Randrianantoandro, J.C., Bauer, A.M. & Vences, M. (2015) Morphology and molecules reveal two new species of the poorly studied gecko genus Paragehyra (Squamata: Gekkonidae) from Madagascar. Organisms Diversity & Evolution, 15, 175–198.
    de Gouvenain, R.C. & Silander, J.A. (2003) Littoral forest. In: Goodman, S.M. & Benstead, J.P. (Eds.), The Natural History of Madagascar.The University of Chicago Press, London, pp. 103–111
    de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms. Species and Speciation. Oxford University Press, Oxford, pp. 49–89.
    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.
    de Queiroz, K. & Good, D.A. (1997) Phenetic clustering in biology: A critique. Quarterly Review of Biology, 72, 3–30.
    DeSalle, R., Egan, M.G. & Siddall, M. (2005) The unholy trinity: taxonomy, species delimitation and DNA barcoding. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 360, 1905–1916.
    Dufresnes, C., Brelsford, A., Jeffries, D.L., Mazepa, G., Suchan, T., Canestrelli, D., Nicieza, A., Fumagalli, L., Dubey, S., Martínez-Solano, I., Litvinchuk, S.N., Vences, M., Perrin, N. & Crochet P.-A. (2021) Mass of genes rather than master genes underlie the genomic architecture of amphibian speciation. Proceedings of the National Academy of Sciences of the U.S.A., 118, e2103963118.
    Fisher, B.L. & Girman, D. (2000) Biogeography of ants in eastern Madagascar. In: Lourenço, W.R. & Goodman, S.M. (Eds.), Diversity and Endemism in Madagascar. Mémoires de la Société de Biogéographie, Paris, pp. 331–344.
    Fölling, M., Knogge, C. & Böhme, W. (2001) Geckos are milking honeydew-producing planthoppers in Madagascar. Journal of Natural History, 35, 279–284.
    Fujisawa, T., Aswad, A. & Barraclough, T.G. (2016) A rapid and scalable method for multilocus species delimitation using Bayesian model comparison and rooted triplets. Systematic Biology, 65, 759–771.
    Ganzhorn, J.U., Goodman, S.M., Ramanamanjato, J.B., Rakotondavony, D., Rakotosamimanana, B. & Vallan, D. (2000) Vertebrate species in fragmented littoral forest of Madagascar. In: Lourenco, W.R. & Goodman, S.M. (Eds.), Diversity and endemism in Madagascar. Mémoires de la Société de Biogéographie Edition. Museum Histoire Naturelle, Paris, pp. 155–164.
    Gehring, P.S., Ratsoavina, F.M. & Vences, M. (2010) Filling the gaps – amphibian and reptile records from lowland rainforests in eastern Madagascar. Salamandra, 46, 214–234.
    Gehring, P.-S., Ratsoavina, F.M., Vences, M. & Glaw, F. (2011) Calumma vohibola, a new chameleon species (Squamata: Chamaeleonidae) from the littoral forests of eastern Madagascar. African Journal of Herpetology, 60, 130–154.
    Gehring, P.-S., Tolley, K.A., Eckhardt, F.S., Townsend, T.M., Ziegler, T., Ratsoavina, F., Glaw, F. & Vences, M. (2012) Hiding deep in the trees: discovery of divergent mitochondrial lineages in Malagasy chameleons of the Calumma nasutum group. Ecology and Evolution, 2, 1468–1479.
    Gippner, S., Travers S.L., Scherz M.D., Colston T.J., Lyra M.L., Mohan A.V., Multzsch M., Nielsen S.V., Rancilhac L., Glaw F., Bauer A.M. & Vences M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311.
    Glaw, F., Gehring, P.-S., Köhler, J., Franzen, M. & Vences, M. (2010) A new dwarf species of day gecko, genus Phelsuma, from the Ankarana pinnacle karst in northern Madagascar. Salamandra, 46, 83–92.
    Glaw, F., Köhler, J., Bora, P., Rabibisoa, N.H.C., Ramilijaona, O. & Vences, M. (2007) Discovery of the genus Plethodontohyla (Anura: Microhylidae) in dry western Madagascar: description of a new species and biogeographic implications. Zootaxa, 1577, 61–68.
    Glaw, F., Köhler, J. & Vences, M. (2018) Three new species of nocturnal geckos of the Paroedura oviceps clade from xeric environments of Madagascar (Squamata: Gekkonidae). Zootaxa, 4433, 305–324.
    Glaw, F., Rösler, H., Ineich, I., Gehring, P.-S., Köhler, J. & Vences, M. (2014) A new species of nocturnal gecko (Paroedura) from karstic limestone in northern Madagascar. Zoosystematics and Evolution, 90, 249–259.
    Glaw, F. & Vences, M. (1992) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 331 pp. [First Edition.]
    Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 480 pp. [Second Edition.]
    Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. [Third Edition.]
    Goodman, S.M. & Ramanamanjato, J.P. (2007) A perspective on the Paleo–ecology and biogeography of extreme southeastern Madagascar, with special reference to animals. In: Ganzhorn, J.U., Goodman, S.M. & Vincelette, M. (Eds.), Biodiversity, Ecology and Conservation of the Littoral Ecosystems of South–eastern Madagascar, Tolagnaro (Fort Dauphin). SI/MAB Series #11. Smithsonian Institution, Washington DC, USA, pp. 25– 48.
    Gray, J.E. (1864) Notes on some new lizards from south-eastern Africa, with the descriptions of several new species. Proceedings of the Zoological Society of London, 1864, 58–62.
    Grismer, L., Perry, L.W., Quah, E.S.H. & Thura, M.K. (2020) Origin, diversity, and conservation of karst-associated Bent-toed Geckos (Genus Cyrtodactylus) in Myanmar (Burma). Israel Journal of Ecology and Evolution, 66, 202–208.
    Günther, A. (1891) Eleventh Contribution to the knowledge of the Fauna of Madagascar. Annals and Magazine of Natural History, 8, 287–288.
    Hawlitschek, O., Scherz, M. D., Ruthensteiner, B., Crottini, A. & Glaw, F. (2018) Computational molecular species delimitation and taxonomic revision of the gecko genus Ebenavia Boettger, 1878. Naturwissenschaften, 105, 49.
    Hutter, C.R., Andriampenomanana, Z.F., Razafindraibe, J., Rakotoarison, A. & Scherz, M.D. (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.
    Ineich, I., Glaw, F. & Vences, M. (2016) A new species of Blaesodactylus (Squamata: Gekkonidae) from Tsingy limestone outcrops in Namoroka National Park, north-western Madagascar. Zootaxa, 4109, 523–541.
    IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. IUCN, Gland, Switzerland and Cambridge, UK.
    Jono, T., Bauer, A.M., Brennan, I. & Mori, A. (2015) New species of Blaesodactylus (Squamata: Gekkonidae) from Tsingy karstic outcrops in Ankarana National Park, northern Madagascar. Zootaxa, 3980, 406–416.
    Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.
    Kluge, A.G. (1991) Checklist of Gekkonoid Lizards. Smithsonian Herpetological Information Service 85, 36 pp.
    Köhler, J., Vences, M., Scherz, M.D. & Glaw, F. (2019) A new species of nocturnal gecko, genus Paroedura, from the karstic Tsingy de Bemaraha formation in western Madagascar. Salamandra, 55, 73–81.
    Kollár, J., Poulíčková, A. & Dvořák, P. (2022) On the relativity of species, or the probabilistic solution to the species problem. Molecular Ecology, 31, 411–418.
    Krüger, J. (2001) Die madagassischen Gekkoniden. Teil II: Die Geckos der Gattung Lygodactylus GRAY 1864 (Reptilia: Sauria: Gekkonidae). Gekkota, 3, 3–28.
    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.
    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.
    Librado, P. & Rozas, J. (2009) DnaSP. Version 5. A software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452.
    Malone, J.H. & Fontenot, B.E. (2008) Patterns of reproductive isolation in toads. PLoS One, 3, e3900.
    Mertens, R. (1967) Die herpetologische Sektion des Naturmuseums und Forschungsinstitutes Senckenberg in Frankfurt a. M. nebst einem Verzeichnis ihrer Typen. Senckenbergiana Biologica, 48, Sonderheft A, 1–106.
    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010. New Orleans, LA. pp. 1‒8.
    Michaelsen, W. (1891) Beschreibung der von Herrn Dr. Fr. Stuhlmann auf Sansibar und dem gegenüberliegenden Festlande gesammelten Terricolen. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 9, 1–72.
    Miralles, A., Bruy, T., Crottini, A., Rakotoarison, A., Ratsoavina, F.M., Scherz, M.D., Schmidt, R., Köhler, J., Glaw, F. & Vences, M. (2021) Completing a taxonomic puzzle: integrative review of geckos of the Paroedura bastardi species complex (Squamata, Gekkonidae). Vertebrate Zoology, 71, 27–48.
    Miralles, A. & Vences, M. (2013) New metrics for comparison of taxonomies reveal striking discrepancies among species delimitation methods in Madascincus lizards. PLoS One, 8, e68242.
    Moat, J. & Smith, P. (2007) Atlas of the Vegetation of. Madagascar. Royal Botanic Gardens, Kew.
    Nagy, Z.T., Sonet, G., Glaw, F. & Vences, M. (2012) First large-scale DNA barcoding assessment of reptiles in the biodiversity hotspot of Madagascar, based on newly designed COI primers. PLoS One, 7, e34506.
    Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, e16.
    Pasteur, G. (1965a) Notes préliminaries sur les lygodactyles (Gekkonidés). IV Diagnoses de quelques formes africaines et malgaches. Bulletin du Muséum national d’Histoire naturelle, 36, 311–314.
    Pasteur, G. (1965b) Recherches sur l’évolution des lygodactyles, lézards afro-malgaches actuels. Travaux de l’Institut scientifique chérifien, Séries, Zoology, 29, 1–132.
    Pasteur, G. & Blanc, C.P. (1967) Les lézards du sous-genre malgache de lygodactyles Domerguella (Gekkonidés). Bulletin de la Société Zoologique de France, 92, 583–597.
    Pasteur, G. & Blanc, C.P. (1973) Nouvelles études sur les lygodactyles (Sauriens Gekkonidés). I. Données récentes sur Domerguella et sur ses rapports avec la Phytogéographie malgache. Bulletin de la Société Zoologique de France, 98, 165–174.
    Pasteur, G. & Lumaret, R. (1976) Evolution morphologique brusque dans une population contemporaine d’un lézard de Madagascar. Comptes rendus de l’Académie des sciences de Paris, 282, 1045–1047.
    Penny, S.G., Crottini, A., Andreone, F., Bellati, A., Rakotozafy, L.M.S., Holderied, M.W., Schwitzer, C. & Rosa, G.M. (2017) Combining old and new evidence to increase the known biodiversity value of the Sahamalaza Peninsula, Northwest Madagascar. Contributions to Zoology, 86, 273–296.
    Pellegrin, J. (1933) Les poissons des eaux douces de Madagascar et des iles voisines (Comores, Seychelles, Mascareignes). Mémoires de L’Académie Malgache, Tananarive, 14, 1–222.
    Pocock, R.L. (1895) XXVIII.— Descriptions of new genera and species of Trap-door Spiders belonging to the Group Trionychi. Annals and Magazine of Natural History, Series, 6, 16, 187–197.
    Prötzel, D., Ruthensteiner, B., Scherz, M.D. & Glaw, F. (2015) Systematic revision of the Malagasy chameleons Calumma boettgeri and C. linotum (Squamata: Chamaeleonidae). Zootaxa, 4048, 211–231.
    Prötzel, D., Scherz, M.D., Ratsoavina, F.M., Vences, M. & Glaw, F. (2020) Untangling the trees: Revision of the Calumma nasutum complex (Squamata: Chamaeleonidae). Vertebrate Zoology, 70, 23–59.
    Prötzel, D., Vences, M., Hawlitschek, O., Scherz, M.D., Ratsoavina, F.M. & Glaw, F. (2018) Endangered beauties: micro-CT cranial osteology, molecular genetics and external morphology reveal three new species of chameleons in the Calumma boettgeri complex (Squamata: Chamaeleonidae). Zoological Journal of the Linnean Society, 184, 471–498.
    Prötzel, D., Vences, M., Scherz, M.D., Vieites, D.R. & Glaw, F. (2017) Splitting and lumping: An integrative taxonomic assessment of Malagasy chameleons in the Calumma guibei complex results in the new species C. gehringi sp. nov. Vertebrate Zoology, 67, 231–249.
    Puente, M., Glaw, F., Vieites, D.R. & Vences, M. (2009) Review of the systematics, morphology and distribution of Malagasy dwarf geckos, genera Lygodactylus and Microscalabotes (Squamata: Gekkonidae). Zootaxa, 2103, 1–76.
    Puente, M., Thomas, M. & Vences, M. (2005) Phylogeny and biogeography of Malagasy dwarf geckos, Lygodactylus Gray, 1864: Preliminary data from mitochondrial DNA sequences (Squamata: Gekkonidae). In: Huber, B.A. & Lampe, K.H. (Eds.), African Biodiversity: Molecules, Organisms, Ecosystems. Proc. 5th Intern. Symp. Trop. Biol., Museum Koenig, Bonn. Springer, pp. 229–235.
    Puillandre, N., Brouillet, S. & Achaz, G. (2021) ASAP: assemble species by automatic partitioning. Molecular Ecology Resources, 21, 609–622.
    Puillandre, N., Lambert, A., Brouillet, S. & Achaz, G. (2012) ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology, 21, 1864–1877.
    Ramananjara, H.C. (2009) Etude des sources archivistiques et documentaires sur les terres dans la Basse Vallée de l’Ivoloina (Toamasina) de 1897 à 1939 : Essai documentaire/ Mémoire de maîtrise. Antananarivo: Université d’Antananarivo / F. L. S. H / Département d’Histoire / Formation en Archivistique- Documentation-Muséologie, 255 pp.
    Randrianarimanana, H.L.L. (2009) Etude comparative de l’alimentation et du comportement des deux espèces sympatriques d’indriides: Propithecus diadema et Indri indri dans la Reserve Naturelle Integrale N01 de Betampona (Toamasina). DEA dissertation, Université d’Antananarivo, Departement de Paleontologie et d’Anthropologie Biologique. Antananarivo, 88 pp.
    Randriatavy, L.D. (2003) Etude de regeneration forestiere et de quelques especies vegetales utilisées a la fois par Varecia variegata et par l’homme dans la Reserve Naturelle Integrale No 1 de Betampona. DEA dissertation, Université d’Antananarivo, Département de Biologie et Ecologie Végétales. Antananarivo, 114 pp.
    Ratsoavina, F.M., Gehring, P.S., Scherz, M.D., Vieites, D.R., Glaw, F. & Vences, M. (2017) Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar. Zootaxa, 4347, 446–464.
    Ratsoavina, F.M., Glaw, F., Raselimanana, A.P., Rakotoarison, A., Vieites, D.R., Hawlitschek, O., Vences, M. & Scherz, M.D. (2020) Towards completion of the species inventory of small-sized leaf-tailed geckos: two new species of Uroplatus from northern Madagascar. Zootaxa, 4895, 251–271.
    Ratsoavina, F.M., Louis, E.E.Jr., Crottini, A., Randrianiaina, R.D., Glaw, F. & Vences, M. (2011) A new leaf tailed gecko species from northern Madagascar with a preliminary assessment of molecular and morphological variability in the Uroplatus ebenaui group. Zootaxa, 3022, 39–57.
    Ratsoavina, F.M., Raminosoa, N.R., Louis, E.E.Jr., Raselimanana, A.P., Glaw, F. & Vences, M. (2013) An overview of Madagascar’s leaf tailed geckos (genus Uroplatus): species boundaries, candidate species and review of geographical distribution based on molecular data. Salamandra, 49, 115–148.
    Ratsoavina, F.M., Ranjanaharisoa, F.A., Glaw, F., Raselimanana, A.P., Miralles, A. & Vences, M. (2015) A new leaf-tailed gecko of the Uroplatus ebenaui group (Squamata: Gekkonidae) from Madagascar’s central eastern rainforests. Zootaxa, 4006, 143–160.
    Ratsoavina, F.M., Raselimanana, A.P., Scherz, M.D., Rakotoarison, A., Razafindraibe, J.H., Glaw, F. & Vences, M. (2019a) Finaritra! A new leaf-tailed gecko (Uroplatus) species from Marojejy National Park in north-eastern Madagascar. Zootaxa, 4545, 563–577.
    Ratsoavina, F.M., Scherz, M.D., Tolley, K.A., Raselimanana, A.P., Glaw, F. & Vences, M. (2019b) A new species of Uroplatus (Gekkonidae) from Ankarana National Park, Madagascar, of remarkably high genetic divergence. Zootaxa, 4683, 84–96.
    Ratsoavina, F.M., Vences, M. & Louis Jr., E.E. (2012) Phylogeny and phylogeography of the Malagasy leaf-tailed geckos in the Uroplatus ebenaui group. African Journal of Herpetology, 61, 143–158.
    Rösler, H. (2000a) Studien an den Begattungsorganen der Geckos (Reptilia: Gekkomorpha) – 2. Die Hemipenismorphologie von Lygodactylus Gray, 1864 (Gekkonidae: Gekkoninae). Gekkota, 2, 208–219.
    Rösler, H. (2000b) Kommentierte Liste der rezent, subrezent und fossil bekannten Geckotaxa (Reptilia: Gekkonomorpha). Gekkota, 2, 28–153.
    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.
    Rosa, G.M., Andreone, F., Crottini, A., Hauswald, J.S., Noël, J., Rabibisoa, N.H., Randriambahiniarime, M.O., Rebelo, R. & Raxworthy, C.J. (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.
    Salzburger, W., Ewing, G.B. & Von Haeseler, A. (2011) The performance of phylogenetic algorithms in estimating haplotype genealogies with migration. Molecular Ecology, 20, 1952–1963.
    Scherz, M.D., Daza, J.D., Köhler, J., Vences, M. & Glaw, F. (2017) Off the scale: a new species of fish-scale gecko (Squamata: Gekkonidae: Geckolepis) with exceptionally large scales. PeerJ, 5, e2955. 10.7717/peerj.e2955
    Scherz, M.D., Hutter, C.R., Rakotoarison, A., Riemann, J.C., Rödel, M.O., Ndriantsoa, S.H., Glos, J., Hyde Roberts, S., Crottini, A., Vences, M. & Glaw, F. (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. PLoS One, 14, e0213314.
    Schultz, M. (2013) The orchid types of the Herbarium Hamburgense (HBG). Verhandlungen des Naturwissenschaftlichen Vereins Hamburg NF, 47, 5–251.
    Sharpe, R.B. & Ogilvie-Grant, W.R. (1898) Catalogue of the Birds in the British Museum. Volume XXVI. Taylor and Francis, London, 314 pp.
    Sparks, J.S. & Nelson, D.W. (2004) Review of the Malagasy sicydiine gobies (Teleostei: Gobiidae), with description of a new species and comments on the taxonomic status of Gobius lagocephalus Pallas, 1770. American Museum Novitates, 3440, 1–20.;2
    Speybroeck, J., Beukema, W., Dufresnes, C., Fritz, U., Jablonski, D., Lymberakis, P., Martinez-Solano, I., Razzettis, E., Vamberger, M., Vences, M., Vörös, J. & Crochet, P.-A. (2020) Species list of the European herpetofauna – 2020 update by the Taxonomic Committee of the Societas Europaea Herpetologica. Amphibia-Reptilia, 41, 139–189.
    Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
    Stephens, M., Smith, N.J. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics, 68, 978–989.
    Uetz, P., Freed, P, Aguilar, R. & Hošek, J. (eds.) (2021) The Reptile Database. Available from: (accessed 01 December 2021)
    Vences, M., Galán, P., Miramontes, K. & Vieites, D.R. (2004) Weak expression of reproductive seasonality in a dwarf gecko (Lygodactylus) from arid south-western Madagascar. Journal of Arid Environments, 56, 329–338.
    Zhang, J., Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29, 2869–2876.
    Zieliński, P., Nadachowska-Brzyska, K., Wielstra, B., Szkotak, R., Covaciu-Marcov, S.D., Cogălniceanu, D. & Babik, W. (2013) No evidence for nuclear introgression despite complete mtDNA replacement in the Carpathian newt (Lissotriton montandoni). Molecular Ecology, 22, 1884–1903.