Two new Pristimantis (Anura: Strabomantidae) of the P. lacrimosus species group from the eastern Guiana Shield lowlands
Abstract
Many anuran species remain to be formally named and described in Amazonia, notably in the Guiana Shield and particularly in megadiverse groups such as Pristimantis. These descriptions are urgently needed to include species in local conservation and protection policies. In French Guiana, two species of Pristimantis of the P. lacrimosus group have been reported as Pristimantis sp. 3 and Pristimantis sp. 5. Using integrative taxonomy, we compared these two species with all other species of the lacrimosus group and confirmed their status as distinct species. Herein, we describe Pristimantis fouqueti sp. nov. and Pristimantis flavus sp. nov., both endemic to the eastern Guiana Shield lowlands.
References
Altschul S.F., Madden T.L., Schäffer A.A., Zhang J., Zhang Z., Miller W. & Lipman D.J. 1997. Gapped BLAST and PSI-BLAST: A new generation of protein database search programs. Nucleic Acids Research 25: 3389–3402. https://doi.org/10.1093/nar/25.17.3389
Arteaga-Navarro A.F., Bustamante M.R. & Guayasamin J.M. 2013. The Amphibians and Reptiles of Mindo. Life in the Cloudforest. Universidad Tecnológica Indoamérica, Quito.
Batallas-Revelo D. & Brito-M J. 2014. Nueva especie de rana del género Pristimantis del grupo lacrimosus (Amphibia: Craugastoridae) del Parque Nacional Sangay, Ecuador. Papéis Avulsos de Zoologia 54: 51–62. https://doi.org/10.1590/0031-1049.2014.54.05
Batallas-Revelo D. & Brito-M J. 2023. The advertisement calls of Pristimantis galdi Jiménez de la Espada, 1870 and Pristimantis katoptroides (Flores, 1988) (Anura, Strabomantidae). Herpetozoa 36: 59–64. https://doi.org/10.3897/herpetozoa.36.e97705
Blanc M. 2016. Inventaire herpétologique du Pic Coudreau du Sud. Les Cahiers scientifiques du Parc amazonien de Guyane 2 (1): 327–339.
Carrión-Olmedo J.C. & Ron S.R. 2021. A new cryptic species of the Pristimantis lacrimosus group (Anura, Strabomantidae) from the eastern slopes of the Ecuadorian Andes. Evolutionary Systematics 5: 151–175. https://doi.org/10.3897/evolsyst.5.62661
Castillo-Urbina E., Vences M., Aguilar-Puntriano C., Glaw F. & Köhler J. 2023. Contributing to the taxonomic inventory of green-colored rain frogs: A new species of the Pristimantis lacrimosus group (Anura: Strabomantidae) from the southern Cordillera Azul, central Peru. Vertebrate Zoology 73: 1047–1061. https://doi.org/10.3897/vz.73.e109309
Chávez G. & Catenazzi A. 2016. A new species of frog of the genus Pristimantis from Tingo María National Park, Huánuco Department, central Peru (Anura, Craugastoridae). ZooKeys 610: 113–130. https://doi.org/10.3897/zookeys.610.8507
Despax R. 1911. Mission géodésique de l’Équateur. Collections recueilles par M. le Dr. Rivet. Batraciens anoures. Bulletin du Museum national d’Histoire naturelle 17: 90–94. Available from https://www.biodiversitylibrary.org/page/34198121 [accessed 26 Nov. 2024].
Dewynter M., Courtois E.A., Massary J.-C. de, Uriot Q., Uriot S., Premel V., Villette B., Rufray V., Perrier M., Bouchet L., Le Pape T., Foxonet H., Remérand E., Baudain D., Baglan A., Marty C. & Bonnefond A. 2020. La base de données Faune-Guyane (Amphibiens, Squamates, Tortues & Caïmans): Synthèse 2019. Herp me! 2: 1– 87.
Dewynter M., Courtois E.A., Massary J.-C. de, Uriot Q., Uriot S., Premel V., Villette B., Rufray V., Perrier M., Bouchet L., Le Pape T., Foxonet H., Remérand E., Baudain D., Baglan A., Marty C. & Bonnefond A. 2021. La base de données Faune-Guyane (Amphibiens, Squamates, Tortues & Caïmans): Synthèse 2020. Herp me! 4: 1– 87.
Duellman W.E. 1978. Three new species of Eleutherodactylus from Amazonian Perú (Amphibia: Anura: Leptodactylidae). Herpetologica 34: 264–270.
Duellman W.E. 1990. A new species of Eleutherodactylus from the Andes of northern Peru (Anura: Leptodactylidae). Journal of Herpetology 24: 348–350. https://doi.org/10.2307/1565048
Duellman W.E. & Lehr E. 2009. Terrestrial-Breeding Frogs (Strabomantidae) in Peru. Natur und Tier Verlag.
Duellman W.E. & Pramuk J.B. 1999. Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers. Natural History Museum, The University of Kansas 13: 1–78. https://doi.org/10.5962/bhl.title.16169
Fouquet A., Martinez Q., Courtois E.A., Dewynter M., Pineau K., Gaucher P., Blanc M., Marty C. & Kok P.J.R. 2013. A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately evelated massifs of French Guiana. Zootaxa 3750: 569–586. https://doi.org/10.11646/zootaxa.3750.5.8
Fouquet A., Vidal N. & Dewynter M. 2019a. The amphibians of the Mitaraka massif, French Guiana. Zoosystema 41: 359–374. https://doi.org/10.5252/zoosystema2019v41a19
Fouquet A., Vacher J.-P., Courtois E.A., Deschamps C., Ouboter P.E., Jairam R., Gaucher P., Dubois A. & Kok P.J.R. 2019b. A new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group with the redescription of A. baeobatrachus (Boistel and de Massary, 1999), and an amended definition of A. leopardus Ouboter and Jairam, 2012. Zootaxa 4576: 439–460. https://doi.org/10.11646/zootaxa.4576.3.2
Fouquet A., Cornuault J., Rodrigues M.T., Werneck F.P., Hrbek T., Acosta-Galvis A.R., Massemin D., Kok P.J.R. & Ernst R. 2022a. Diversity, biogeography and reproductive evolution in the genus Pipa (Amphibia: Anura: Pipidae). Molecular Phylogenetics and Evolution 170: 107442. https://doi.org/10.1016/j.ympev.2022.107442
Fouquet A., Réjaud A., Rodrigues M.T., Ron S.R., Chaparro J.C., Osorno M., Werneck F.P., Hrbek T., Lima A.P., Camacho-Badani T., Jaramillo-Martinez A.F. & Chave J. 2022b. Diversification of the Pristimantis conspicillatus group (Anura: Craugastoridae) within distinct neotropical areas throughout the Neogene. Systematics and Biodiversity 20: 1–16. https://doi.org/10.1080/14772000.2022.2130464
Fouquet A., Ferrão M. & Jairam R. 2023. Two new species of Allobates of the trilineatus clade (Anura: Aromobatidae) from the Eastern Guiana Shield. Zootaxa 5297: 533–561. https://doi.org/10.11646/zootaxa.5297.4.4
Fouquet A., Courtois É.A. & Dewynter M. 2024. Atlas of the Amphibians of French Guiana. Éditions Biotope, Mèze, Muséum national d’Histoire naturelle, Paris (collection inventaires & biodiversité).
Frost D.R. 2025. Amphibian Species of the World: an Online Reference. Version 6.2 Available from https://amphibiansoftheworld.amnh.org/ [accessed 15 Jul. 2025].
Guayasamin J.M., Ron S.R., Cisneros-Heredia D.F., Lamar W.W. & McCracken S.F. 2006. A new species of frog of the Eleutherodactylus lacrimosus assemblage (Leptodactylidae) from the western Amazon Basin, with comments on the utility of canopy surveys in lowland rainforest. Herpetologica 62: 191–202. https://doi.org/10.1655/05-40.1
Guayasamin J.M., Hutter C.R., Tapia E.E., Culebras J., Peñafiel N., Pyron R.A., Morochz C., Funk W.C. & Arteaga-Navarro A.F. 2017. Diversification of the rainfrog Pristimantis ornatissimus in the lowlands and Andean foothills of Ecuador. PLoS ONE 12 (3): e0172615. https://doi.org/10.1371/journal.pone.0172615
Heinicke M.P., Duellman W.E. & Hedges S.B. 2007. Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences 104: 10092–10097. https://doi.org/10.1073/pnas.0611051104
Heyer W.R. & Hardy L.M. 1991. A new species of frog of the Eleutherodactylus lacrimosus assembly from Amazonia, South America (Amphibia: Anura: Leptodactylidae). Proceedings of the Biological Society of Washington 104: 436–447. Available from https://www.biodiversitylibrary.org/page/34809134 [accessed 26 Nov. 2024].
Hoang D.T., Chernomor O., Von Haeseler A., Minh B.Q. & Vinh L.S. 2018. UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–522. https://doi.org/10.1093/molbev/msx281
Hutter C.R., Liu V., Kell T., Lyons J.A. & Guayasamin J.M. 2016. The natural history, distribution, and conservation of Lonely Rainfrogs, Pristimantis eremitus. Herpetologica 72: 13–22. https://doi.org/10.1655/Herpetologica-D-14-00020
K. Lisa Yang Center for Conservation Bioacoustics at the Cornell Lab of Ornithology 2024. Raven Pro: Interactive Sound Analysis Software. Version 1.6.5 [Computer software]. Available from https://www.ravensoundsoftware.com/ [accessed 11 Dec. 2024].
Kalyaanamoorthy S., Minh B.Q., Wong T.K., Von Haeseler A. & Jermiin L.S. 2017. ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14 (6): 587–589. https://doi.org/10.1038/nmeth.4285
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. https://doi.org/10.1093/molbev/mst010
Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16: 111–120. https://doi.org/10.1007/BF01731581
Köhler J., Morales V.R., Lötters S., Reichle S. & Aparicio J. 1998. A new green species of frog, genus Eleutherodactylus, from Bolivia and Peru (Amphibia, Anura, Leptodactylidae). Studies on Neotropical Fauna and Environment 33: 93–99. https://doi.org/10.1076/snfe.33.2.93.2158
Kok P.J. 2013. Two new charismatic Pristimantis species (Anura: Craugastoridae) from the tepuis of the “Lost World”(Pantepui region, South America). European Journal of Taxonomy 60: 1–24. https://doi.org/10.5852/ejt.2013.60
Kok P.J., Means D.B. & Bossuyt F. 2011. A new highland species of Pristimantis Jiménez de la Espada, 1871 (Anura: Strabomantidae) from the Pantepui region, northern South America. Zootaxa 2934: 1–19. https://doi.org/10.11646/zootaxa.2934.1.1
Lehr E., Lundberg M., Aguilar C. & von May R. 2006. New species of Eleutherodactylus (Anura: Leptodactylidae) from the eastern Andes of central Peru with comments on central Peruvian Eleutherodactylus. Herpetological Monographs 20: 105–128. https://doi.org/10.1655/0733-1347(2007)20[105:NSOEAL]2.0.CO;2
Lehr E., Torres-Gastello C.P. & Suárez-Segovia J. 2007. A new species of arboreal Eleutherodactylus (Anura: Leptodactylidae) from the Amazonian lowlands of central Peru. Herpetologica 63: 94–99. https://doi.org/10.1655/0018-0831(2007)63[94:ANSOAE]2.0.CO;2
Lescure J. & Marty C. 2000. Atlas des Amphibiens de Guyane. Muséum d’Histoire naturelle, Paris.
Lescure J., Dewynter M., Frétey T., Ineich I., Ohler A., Vidal N. & De Massary J.-C. 2022. Liste taxinomique de l’herpétofaune dans l’outre-mer français: VII. Les Amphibiens de la Guyane française. Bulletin de la Société herpétologique de France 181: 17
Lynch J.D. 1970. Redescriptions of three little-known Eleutherodactylus from northwestern Ecuador (Amphibia: Leptodactylidae). Transactions of the Kansas Academy of Science 73: 169–180. https://doi.org/10.2307/3627301
Lynch J.D. 1976. New species of frogs (Leptodactylidae: Eleutherodactylus) from the Pacific versant of Ecuador. Occasional Papers of the Museum of Natural History, University of Kansas 55: 1–33. https://doi.org/10.5962/bhl.part.29037
Lynch J.D. 1979. Leptodactylid frogs of the genus Eleutherodactylus from the Andes of southern Ecuador. Miscellaneous Publication (University of Kansas. Museum of Natural History) 66: 1–62. https://doi.org/10.5962/bhl.title.16268
Lynch J.D. 1980a. Eleutherodactylus eremitus, a new trans-Andean species of the lacrimosus assembly from Ecuador (Amphibia: Leptodactylidae). Breviora 462: 1–7. Available from https://www.biodiversitylibrary.org/page/3189673 [accessed 26 Nov. 2024].
Lynch J.D. 1980b. Systematic status and distribution of some poorly known frogs of the genus Eleutherodactylus from the Chocoan lowlands of South America. Herpetologica 36: 175–189.
Lynch J.D. 1994. A new species of frog (genus Eleutherodactylus: Leptodactylidae) from a cloud forest in Departamento de Santander, Colombia. Revista de la Academia colombiana de Ciencias exactas, físicas y naturales 19: 205–208.
Lynch J.D. & Duellman W.E. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous publication (University of Kansas. Museum of Natural History) 69: 1–86. https://doi.org/10.5962/bhl.title.16222
Lynch J.D. & Duellman W.E. 1997. Frogs of the genus Eleutherodactylus in western Ecuador. Systematics, ecology, and biogeography. Special Publication (University of Kansas. Natural History Museum) 23: 1–236. https://doi.org/10.5962/bhl.title.7951
Maddison D.R. & Maddison W.P. 2000. MacClade 4, Analysis of Phylogeny and Character Evolution. M.S. Associates, Sunderland.
McCracken S.F. & Forstner M.R.J. 2006. Reproductive ecology and behavior of Eleutherodactylus aureolineatus (Anura, Brachycephalidae) in the canopy of the Upper Amazon Basin, Ecuador. Phyllomedusa Journal of Herpetology 5: 135–143. https://doi.org/10.11606/issn.2316-9079.v5i2p135-143
McCracken S.F. & Forstner M.R.J. 2014. Herpetofaunal community of a high canopy tank bromeliad (Aechmea zebrina) in the Yasuní Biosphere Reserve of Amazonian Ecuador, with comments on the use of “arboreal” in the herpetological literature. Amphibian & Reptile Conservation 8 (1): 65–75.
McCracken S.F., Forstner M.R.J. & Dixon J.R. 2007. A new species of the Eleutherodactylus lacrimosus assemblage (Anura, Brachycephalidae) from the lowland rainforest canopy of Yasuni National Park, Amazonian Ecuador. Phyllomedusa Journal of Herpetology 6: 23–36. https://doi.org/10.11606/issn.2316-9079.v6i1p23-35
Means D.B. & Savage J.M. 2007. Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America. Zootaxa 1658: 39–55. https://doi.org/10.5281/zenodo.179899
Means D.B., Heinicke M.P., Hedges S.B., Macculloch R.D. & Lathrop A. 2023. Exceptional diversity of Pristimantis Landfrogs (Anura: Terraranae) on the Wokomung Massif, Guyana, with descriptions of three new species. Journal of Vertebrate Biology 72: 23026.1. https://doi.org/10.25225/jvb.23026
Mendoza A.M., Ospina O.E., Cárdenas-Henao H. & García-R J.C. 2015. A likelihood inference of historical biogeography in the world’s most diverse terrestrial vertebrate genus: Diversification of direct-developing frogs (Craugastoridae: Pristimantis) across the Neotropics. Molecular Phylogenetics and Evolution 85: 50–58. https://doi.org/10.1016/j.ympev.2015.02.001
Mônico A.T., Ferrão M., Chaparro J.C., Fouquet A. & Lima A.P. 2022. A new species of rain frog (Anura: Strabomantidae: Pristimantis) from the Guiana Shield and amended diagnosis of P. ockendeni (Boulenger, 1912). Vertebrate Zoology 72: 1035–1065. https://doi.org/10.3897/vz.72.e90435
Mônico A.T., Koch E.D., Ferrão M., Fernandes I.Y., Marques G.M.G., Chaparro J.C., Rodrigues M.T., Lima A.P. & Fouquet A. 2024. The small and inconspicuous majority: Revealing the megadiversity and historical biogeography of the Pristimantis unistrigatus species group (Anura, Strabomantidae). Molecular Phylogenetics and Evolution 201: e108203. https://doi.org/10.1016/j.ympev.2024.108203
Moraes L.J., Werneck F.P., Réjaud A., Rodrigues M.T., Prates I., Glaw F., Kok P.J., Ron S.R., Chaparro J.C., Osorno-Muñoz M., Dal Vechio F., Recoder R.S., Marques-Souza S., Rojas R.R., Demay L., Herbek T. & Fouquet A. 2022. Diversification of tiny toads (Bufonidae: Amazophrynella) sheds light on ancient landscape dynamism in Amazonia. Biological Journal of the Linnean Society 136 (1): 75–91. https://doi.org/10.1080/10.1093/biolinnean/blac006
Morales V.R. 2007. Una especie nueva de Eleutherodactylus (Amphibia: Anura: Brachycephalidae) de la Amazonía central del Perú. Biotempo 7: 5–11.
Moravec J., Lehr E., Pérez-Peña P.E., López-Rojas J.J., Gagliardi-Urrutia L.A.G. & Arista-Tuanama I. 2010. A new green, arboreal species of Pristimantis (Anura: Strabomantidae) from Amazonian Peru. Vertebrate Zoology 60: 225–232. https://doi.org/10.3897/vz.60.e31018
Nguyen L.-T., Schmidt H.A., Von Haeseler A. & Minh B.Q. 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
Ortega-Andrade H.M., Rojas-Soto O.R., Valencia J.H., Espinosa de los Monteros A., Morrone J.J., Ron S.R. & Cannatella D.C. 2015. Insights from integrative systematics reveal cryptic diversity in Pristimantis frogs (Anura: Craugastoridae) from the Upper Amazon Basin. PLoS ONE 10 (11): e0143392. https://doi.org/10.1371/journal.pone.0143392
Ospina-Sarria J.J. & Duellman W.E. 2019. Two new Species of Pristimantis (Amphibia: Anura: Strabomantidae) from southwestern Colombia. Herpetologica 75: 85–95. https://doi.org/10.1655/D-18-00019
Padial J.M., Castroviejo-Fisher S., Köhler J., Domic E. & De la Riva I. 2007. Systematics of the Eleutherodactylus fraudator species group (Anura: Brachycephalidae). Herpetological Monographs 21: 213–240. https://doi.org/10.1655/06-007.1
Padial J.M., Castroviejo-Fisher S., Köhler J., Vilà C., Chaparro J.C. & De la Riva I. 2009. Deciphering the products of evolution at the species level: The need for an integrative taxonomy. Zoologica Scripta 38 (4): 431–447. https://doi.org/10.1111/j.1463-6409.2008.00381.x
Padial J.M., Miralles A., De la Riva I. & Vences M. 2010. The integrative future of taxonomy. Frontiers in Zoology 7: 1–14. https://doi.org/10.1186/1742-9994-7-16
Páez N.B. & Ron S.R. 2019. Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species. ZooKeys 868: 1–112. https://doi.org/10.3897/zookeys.868.26766
Rivera-Correa M. & Daza J.M. 2020. Out of the blue: A new rain frog species of the genus Pristimantis (Anura: Craugastoridae) from the northern Cordillera Central in Colombia. Zootaxa 4838: 83–101. https://doi.org/10.11646/zootaxa.4838.1.4
Ron S.R., Carrión J., Caminer M.A., Sagredo Y., Navarrete M.J., Ortega J.A., Varela-Jaramillo A., Maldonado-Castro G.A. & Terán C. 2020. Three new species of frogs of the genus Pristimantis (Anura, Strabomantidae) with a redefinition of the P. lacrimosus species group. ZooKeys 993: 121–155. https://doi.org/10.3897/zookeys.993.53559
Savage J.M. 1965. A new bromeliad frog of the genus Eleutherodactylus from Costa Rica. Bulletin of the Southern California Academy of Sciences 64: 106–110. Available from https://www.biodiversitylibrary.org/page/34186172 [accessed 26 Nov. 2024].
Savage J.M. 2002. The Amphibians and Reptiles of Costa Rica. University of Chicago Press, Chicago.
Shepack A., von May R., Ttito A. & Catenazzi A. 2016. A new species of Pristimantis (Amphibia, Anura, Craugastoridae) from the foothills of the Andes in Manu National Park, southeastern Peru. ZooKeys 594: 143-164. https://doi.org/10.3897/zookeys.594.8295
Shreve B. 1935. On a new teiid and Amphibia from Panama, Ecuador, and Paraguay. Occasional Papers of the Boston Society of Natural History 8: 209–218.
Tamura K., Stecher G. & Kumar S. 2021. MEGA11: Molecular Evolutionary Genetics Analysis version 11. Molecular Biology and Evolution 38: 3022–3027. https://doi.org/10.1093/molbev/msab120
Taucce P.P., Costa-Campos C.E., Carvalho T.R. & Michalski F. 2022. Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps. European Journal of Taxonomy 836: 96–130. https://doi.org/10.5852/ejt.2022.836.1919
Trifinopoulos J., Nguyen L.-T., von Haeseler A. & Minh B.Q. 2016. W-IQ-TREE: A fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44: W232–W235. https://doi.org/10.1093/nar/gkw256
Vacher J.P., Kok P.J., Rodrigues M.T., Lima J.D., Lorenzini A., Martinez Q., Fallet M., Courtois E.A., Blanc M., Gaucher P., Dewynter M., Jairam R., Ouboter P., Thébaud C. & Fouquet A. 2017. Cryptic diversity in Amazonian frogs: Integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield. Molecular Phylogenetics and Evolution 112: 158–173. https://doi.org/10.1016/j.ympev.2017.04.017
Vacher J., Chave J., Ficetola F.G., Sommeria-Klein G., Tao S., Thébaud C., Blanc M., Camacho A., Cassimiro J., Colston T.J., Dewynter M., Ernst R., Gaucher P., Gomes J.G., Jairam R., Kok P.J.R., Lima J.D., Martinez Q., Marty C., ... & Fouquet A. 2020. Large-scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism. Journal of Biogeography 47: 1781–1791. https://doi.org/10.1111/jbi.13847
Waddell E.H., Crotti M., Lougheed S.C., Cannatella D.C. & Elmer K.R. 2018. Hierarchies of evolutionary radiation in the world’s most species rich vertebrate group, the Neotropical Pristimantis leaf litter frogs. Systematics and Biodiversity 16: 807–819. https://doi.org/10.1080/14772000.2018.1503202
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