Abstract
Nephelomys albigularis is a sigmodontine rodent of the tribe Oryzomyini distributed in the Andean forests from central Ecuador to central Peru. Although several studies recognize this species as monotypic, significant morphological variation has been reported in Peruvian populations that were not properly assessed by direct comparisons with the type series from central Ecuador. We present a preliminary review of N. albigularis with an integrative approach and emphasis on Peruvian populations. We analyzed specimens using morphological and morphometric methods, complemented with phylogenetic analyses and species delimitation using sequence data from the cytochrome-b gene. Our results reveal that N. albigularis (sensu lato) comprises two taxa: N. albigularis s.s., from the montane forests in central and southern Ecuador and northwestern Peru, and Nephelomys sp. nov. from montane forest east of the Marañón River. These taxa are morphologically distinct and are separated by a genetic distance of 5.90 ± 1.01%. Nephelomys sp. nov. differs from N. albigularis s.s. by longer rump hairs, narrow hypothenar pads, faintly bicolor tail; absent interorbital ridges, low zygomatic plates, smoothly squared posterior margin of the hard palate or with a small median postpalatal process, smoothly edged ventral margin of the external auditory meatus, and slightly angular mandibular sigmoid notches. In this work, we present a diagnosis and description of the new species of Nephelomys and discuss the role of the Marañón River as a potential driver for speciation in the genus Nephelomys.
References
Aguilera, M., Pérez-Zapata, A. & Martino, A. (1995) Cytogenetics and karyosystematics of Oryzomys albigularis (Rodentia, Cricetidae) from Venezuela. Cytogenetics and Cell Genetics, 69, 44–49. https://doi.org/10.1159/000133935
Bandelt, H.J., Forster, P. & Röhl, A. (1999) Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16, 37–48. https://doi.org/10.1093/oxfordjournals.molbev.a026036
Brito, J., Tinoco, N. & Zornoza, F. (2015) Nuevo registro distribucional del ratón endémico Nephelomys nimbosus (Rodentia: Cricetidae) en el suroriente de Ecuador. Therya, 6, 667–674. https://doi.org/10.12933/therya-15-298
Brito, J., Koch, C., Percequillo, A.R., Tinoco, N., Weksler, M., Pinto, C.M. & Pardiñas, U.F.J. (2020) A new genus of oryzomyine rodents (Cricetidae, Sigmodontinae) with three new species from montane cloud forests, western Andean cordillera of Colombia and Ecuador. PeerJ, 8, e10247. https://doi.org/10.7717/peerj.10247
Brito, J., Vaca-Puente, S., Koch, C. & Tinoco, N. (2021) Discovery of the first Amazonian Thomasomys (Rodentia, Cricetidae, Sigmodontinae): a new species from the remote Cordilleras del Cóndor and Kutukú in Ecuador. Journal of Mammalogy, 2021, gyaa183. https://doi.org/10.1093/jmammal/gyaa183
Britto, B. (2017) Actualización de las ecorregiones terrestres de Perú propuestas en el Libro Rojo de plantas endémicas del Perú. Gayana Botanica, 74, 15–29. https://doi.org/10.4067/S0717-66432017005000318
Cabrera, A. (1961) Catálogo de los mamíferos de América del sur. Revista del Museo Argentino de Ciencias Naturales “Bernadino Rivadavia” e Instituto Nacional de Investigación de las Ciencias Naturales, Ciencias Zoológicas, 4, 309–732.
Carleton, M.D. & Musser, G.G. (1989) Systematic studies of oryzomyine rodents (Muridae, Sigmodontinae): A synopsis of Microryzomys. Bulletin of the American Museum of Natural History, 191, 1–83.
Chávez Argandoña, A.M. (2018) Influencia del hábitat y la estacionalidad sobre la dieta del oso andino (Tremarctos ornatus) en el distrito de Corosha, departamento de Amazonas. Tesis para obtener el título profesional de Licenciada en Biología. Universidad Peruana Cayetano Heredia. Lima. Available from: https://www.researchgate.net/publication/323497475_INFLUENCIA_DEL_HABITAT_Y_LA_ESTACIONALIDAD_SOBRE_LA_DIETA_DEL_OSO_ANDINO_Tremarctos_ornatus_EN_EL_DISTRITO_DE_COROSHA_DEPARTAMENTO_DE_AMAZONAS
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772. https://doi.org/10.1038/nmeth.2109
Duellman, W.E. (1979) The herpetofauna of the Andes: Patterns of distribution, origin, differentiation, and present communities. In: Duellman, W.E. (Ed.), The South American herpetofauna: Its origin, evolution, and dispersal. Monographs of the Museum of Natural History, University of Kansas, Lawrence, Kansas, pp. 371–459. https://doi.org/10.5962/bhl.title.3207
Duellman, W.E. & Wild, E.R. (1993) Anuran amphibians from the Cordillera de Huancabamba, northern Peru: Systematics, ecology, and biogeography. Occasional Papers of the Museum of Natural History of the University of Kansas, 157, 1–53.
Dunnum, J.L., McLean B.S. & Dowler R.C. (2018) Systematic collections committee of the American Society of Mammalogists, mammal collections of the Western Hemisphere: a survey and directory of collections, Journal of Mammalogy, 99, 1307–1322. https://doi.org/10.1093/jmammal/gyy151
Excoffier, L., Smouse, P.E. & Quattro, J.M. (1992) Analysis of Molecular Variance Inferred from Metric Distances among DNA Haplotypes: Application to Human Mitochondrial DNA Restriction Data. Genetics, 131, 479–491. https://doi.org/10.1093/genetics/131.2.479
Excoffier, L. & Lischer, H.E.L. (2010) Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and Windows. Molecular Ecology Resources, 10, 564–567. https://doi.org/10.1111/j.1755-0998.2010.02847.x
Fischer, G. (1817) Adversaria zoologica. Fasciculus primus. Quaedam ad Mammalium systema et genera illustranda. Mémoires de la Société impériale des naturalists de Moscou, 5, 357–428.
Fujisawa, T. & Barraclough, T.G. (2013) Delimiting species using single-locus data and the Generalized Mixed Yule Coalescent approach: A revised method and evaluation on simulated data sets. Systematic Biology, 62, 707–724. https://doi.org/10.1093/sysbio/syt033
Gardner, A.L. & Patton, J.L. (1976) Karyotypic variation in oryzomyine rodents (Cricetinae) with comments on chromosomal evolution in the Neotropical cricetine complex. Occasional Papers of the Museum of Zoology, Louisiana State University, 49, 1–48.
Gardner, A.L. & Romo, R. M. (1993) A new Thomasomys (Mammalia: Rodentia) from the Peruvian Andes. Proceedings of the Biological Society of Washington, 106, 762–774.
Gyldenstolpe, N. (1932) A manual of Neotropical sigmodont rodents. Kungliga Svenska Vetenskapsakademiens Handlingar, Stockholm, 11 (3), 1–164.
Hershkovitz, P. (1944) A systematic review of the Neotropical water rats of the genus Nectomys (Cricetinae). Miscellaneous Publications, Museum of Zoology, University of Michigan, 58, 1–88.
Hurtado, N. & D’Elìa, G. (2018) Taxonomy of the long-tailed mouse Oligoryzomys destructor (Sigmodontinae: Oryzomyini) with the designation of neotypes for Hesperomys destructor Tschudi, 1844 and Hesperomys melanostoma Tschudii, 1844. Journal Zoological Systematics Evolutionary Research, 57 (1), 127–144. https://doi.org/10.1111/jzs.12232
Hurtado, N. & Pacheco, V. (2017) Revision of Neacomys spinosus (Thomas, 1882) (Rodentia: Cricetidae) with emphasis on Peruvian populations and the description of a new species. Zootaxa, 4242 (3), 401–440. https://doi.org/10.11646/zootaxa.4242.3.1
INRENA (Instituto Nacional de Recursos Naturales). (2003) Plan Maestro 2003–2007. Parque Nacional del Río Abiseo. Imprenta Cano S.R.L., Lima, 228 pp.
International Commission on Zoological Nomenclature. (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.
Jiménez, C.F., Pacheco, V. & Vivas, D. (2013) An introduction to the systematics of Akodon orophilus Osgood, 1913 (Rodentia: Cricetidae) with the description of a new species. Zootaxa, 3669 (3), 223–242. https://doi.org/10.11646/zootaxa.3669.3.2
Koch, C., Venegas, P.J., Garcia-Bravo, A. & Böhme, W. (2011) A new bush anole (Iguanidae: Polychrotinae: Polychrus) from the upper Marañón basin, Peru, with a redescription of Polychrus peruvianus (Noble, 1924) and additional information on P. gutturosus Berthold, 1845. Zookeys, 141, 79–107. https://doi.org/10.3897/zookeys.141.1678
Koopman, K.F. (1978) Zoogeography of Peruvian bats with special emphasis on the role of the Andes. American Museum Novitates, 2651, 1–6.
Kumar, S., Stecher, G. & Tamura, K. (2015) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0. Molecular Biology and Evolution, 33, 1870–1874. https://doi.org/10.1093/molbev/msw054
Lance, R.F, Kennedy, M.L. & Lebero, P.L. (2000) Classification bias in discriminant function analyses used to evaluate putatively different taxa. Journal of Mammalogy, 81, 245–249. https://doi.org/10.1644/1545-1542(2000)081%3C0245:CBIDFA%3E2.0.CO;2
Lau, M.K. (2013) DTK: Dunnett-Tukey-Kramer pairwise multiple comparison test adjusted for unequal variances and unequal sample sizes. (R package version 3.5.) Available from: http://CRAN.R-project.org/package=DTK (accessed 1 February 2020)
Lehr, E., Gregory, C. & Catenazzi, A. (2013) A new species of Pristimantis (Amphibia: Anura: Strabomantidae) from the Río Abiseo National Park, Peru. Zootaxa, 3731, 201–211. https://doi.org/10.11646/zootaxa.3731.2.1
Leo L., M. & Gardner, A.L. (1993) A new species of a giant Thomasomys (Mammalia: Muridae: Sigmodontinae) from the Andes of northcentral Peru. Proceedings of the Biological Society of Washington, 106, 417–428.
Leo, M. (1995) The importance of tropical montane cloud forest for preserving vertebrate endemism in Peru: The Río Abiseo National Park as a case study. In: Hamilton, L.S., Juvik, J.O. & Scatena, F.N. (Eds.), Tropical Montane Cloud Forests. Vol. 110. Springer, New York, New York, pp. 198–211. https://doi.org/10.1007/978-1-4612-2500-3_13
Leo, M. & Romo, M. (1992) Distribución altitudinal de roedores sigmodontinos (Cricetidae) en el Parque Nacional Rio Abiseo, San Martin, Perú. Memorias del Museo de Historia Natural UNMSM, 21, 105–118.
Letunic, I. & Bork, P. (2016) Interactive Tree of Life (iTOL) v3: an online tool for the display and annotation of phylogenetic and other trees. Nucleic Acids Research, 44, W242–W245. https://doi.org/10.1093/nar/gkw290
Librado, P. & Rozas, J. (2009) DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452. https://doi.org/10.1093/bioinformatics/btp187
Márquez, E.J., Aguilera, M. & Corti, M. (2000) Morphometric and chromosomal variation in populations of Oryzomys albigularis (Muridae: Sigmodontinae) from Venezuela: multivariate aspects. Zeitschrift fur Saugetierkunde, 65, 84–99.
Mitchell, B.A., Stolton, S., Bezaury-Creel, J., Bingham, H.C., Cumming, T.L., Dudley, N., Fitzsimons, J.A., Malleret-King, D., Redford, K.H. & Solano, P. (2018) Guidelines for privately protected areas. Best practice protected area guidelines series No. 29. IUCN, Gland, xii + 100 pp. https://doi.org/10.2305/IUCN.CH.2018.PAG.29.en
Monteferri, B. (Ed.) (2019) Áreas de conservación privada en el Perú: avances y propuestas a 20 años de su creación. Sociedad Peruana de Derecho Ambiental, Lima, 114 pp.
Musser, G.M., Carleton, M.D., Brothers, E.M. & Gardner, A.L. (1998). Systematic studies of oryzomyine rodents (Muridae, Sigmodontinae): Diagnoses and distributions of species formerly assigned to Oryzomys “capito”. Bulletin of the American Museum of Natural History, 236, 1–376.
Musser, G.G. & Carleton M.D. (1993) Family Muridae. In D.E. Wilson and D. M. Reeder, Eds., Mammal species of the world: A Taxonomic and Geographic Reference, Smithsonian University Press, Washington, 1206 pp.
Musser, G.G. & Carleton M.D. (2005) Superfamily Muroidea. In Wilson, D. E. and Reeder, D. M., Eds., Mammal Species of the World: A Taxonomic and Geographic Reference, The Johns Hopkins University Press, Baltimore, Maryland, 2142 pp.
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
Pacheco, V. (2002) Mamíferos del Perú. In: Ceballos, G. & Simonetti J. (Eds.), Diversidad y Conservación de los Mamíferos Neotropicales. CONABIO-UNAM, Mexico D.F., pp. 503–550.
Pacheco, V. (2015) Genus Thomasomys Coues, 1884. In: Patton, J.L., Pardiñas, U.F.J. & D’Elia, G. (Eds.), Mammals of South America. Vol. 2. University of Chicago Press, Chicago, pp. 617–683.
Pacheco, V., Cadenillas, R., Salas, E., Tello, C. & Zeballos, H. (2009) Diversidad y endemismo de los mamíferos del Perú. Revista Peruana de Biología, 16, 5–32. https://doi.org/10.15381/rpb.v16i1.111
Pantigoso Pecero M. (2018) Ricardo Palma, el “Bibliotecario mendigo”: amor al libro a través de sus cartas. Aula Palma, 16, 285–308. https://doi.org/10.31381/test2.v0i16.1353
Parada, A., Pardiñas, U.F.J., Salazar-Bravo, J., D’Elía, G. & Palma, R.E. (2013) Dating an impressive Neotropical radiation: Molecular time estimates for the Sigmodontinae (Rodentia) provide insights into its historical biogeography. Molecular Phylogenetics and Evolution, 66, 960–968. https://doi.org/10.1016/j.ympev.2012.12.001
Parada, A., D’Elía, G. & Palma, R.E. (2015) The influence of ecological and geographical context in the radiation of Neotropical sigmodontine rodents. BMC Evolutionary Biology, 15, 172. https://doi.org/10.1186/s12862-015-0440-z
Parker, T.A., Schulenberg, T.S., Graves, G.R. & Braun, M.J. (1985) The avifauna of the Huancabamba region, northern Peru. Ornithological Monographs, 36, 169–197. https://doi.org/10.2307/40168282
Patton, J.L. (1986) Patrones de distribucion y especiacion de fauna de mamiferos de los bosques nublados andinos del Peru. Anales del Museo de Historia Natural Valparaíso, 17, 87–94.
Patton, J.L., Myers, P. & Smith, M.F. (1990) Vicariant versus gradient model of diversification: The small mammal fauna of eastern Andean slopes of Peru. In: Peters, G. & Hutterer, R. (Eds.), Vertebrates in the Tropics. Museum Alexander Koenig, Bonn, pp. 355–371.
Patterson, B.D. (1999) Contingency and determinism in mammalian biogeography: The role of history. Journal of Mammalogy, 80, 345–360. https://doi.org/10.2307/1383284
Percequillo, A.R. (2003) Sistemática de Oryzomys Baird, 1858: definição dos grupos de espécies e revisão do grupo albigularis (Rodentia, Sigmodontinae). Tese de Doutorado, Universidade de São Paulo. São Paulo. [unknown pagination]
Percequillo, A.R. (2015) Genus Nephelomys Weksler, Percequillo, and Voss, 2006. In Patton, J.L., Pardiñas, U.F.J. & D’Elia, G. (Eds.), Mammals of South America, vol. 2. University of Chicago Press, Chicago, Illinois, pp. 377–399.
Pons, J., Barraclough, T.G., Gomez-Zurita, J., Cardoso, A., Duran, D.P., Hazell, S., Kamoun, S., Sumlin, W.D. & Vogler, A.P. (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology, 55, 595–609. https://doi.org/10.1080/10635150600852011
Porzecanski, A.L. & Cracraft, J. (2005) Cladistic Analysis of Distributions and Endemism (CADE): Using raw distributions of birds to unravel the biogeography of the South American arid lands. Journal of Biogeography, 32, 261–275. https://doi.org/10.1111/j.1365-2699.2004.01138.x
Pulliandre, N., Lambert, A., Brouillet, S. & Achaz,G. (2011) ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology, 21 (8), 1864–1877. https://doi.org/10.1111/j.1365-294X.2011.05239.x
R Development Core Team (2017) R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. Available from: http://www.R-project.org/ (accessed 2 August 2021)
Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904. https://doi.org/10.1093/sysbio/syy032
Reig, O.A. (1980) A new fossil genus of South American cricetid rodents allied to Wiedomys with an assessment of the Sigmodontinae. Journal of Zoology, 192, 257–281. https://doi.org/10.1111/j.1469-7998.1980.tb04233.x
Rengifo, E.M. & Pacheco, V. (2015) Taxonomic revision of the Andean leaf-eared mouse, Phyllotis andium Thomas 1912 (Rodentia: Cricetidae), with the description of a new species. Zootaxa, 4018 (3), 349–380. https://doi.org/10.11646/zootaxa.4018.3.2
Rodriguez, L.O. & Catenazzi, A. (2017) Four new species of terrestrial-breeding frogs of the genus Phrynopus (Anura: Terrarana: Craugastoridae) from Río Abiseo National Park, Peru. Zootaxa, 4273 (3), 381–406. https://doi.org/10.11646/zootaxa.4273.3.4
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. https://doi.org/10.1093/sysbio/sys029
Ruelas, D., Pardiñas, U.F.J. & Brito, J. (2017) White-throated rice rat Nephelomys albigularis. In Wilson, D.E., Lacher Jr., T.E. & Mittermeier, R.A. (Eds.), Handbook of the mammals of the world. Rodents II. Vol. 7. Lynx Edicions, Barcelona, pp. 416.
Schenk, J.J. & Steppan, S. J. (2018) The role of geography in adaptive radiation. The American Naturalist, 192, 415–431. https://doi.org/10.1086/699221
Shanee, S., Shanee, N. & Allgas-Marchena, N. (2013) Primate surveys in the Marañón-Huallaga landscape, northern Peru with notes on conservation. Primate Conservation, 27, 3–11. https://doi.org/10.1896/052.027.0114
Smith, M.F. & Patton, J.L. (1991) Variation in mitochondrial cytochrome b sequence in natural populations of South American akodontine rodents (Muridae: Sigmodontinae). Molecular Biology and Evolution, 8, 85–103.
Smith, M.F. & Patton, J.L. (1993) The diversification of South American murid rodents: evidence from mitochondrial DNA sequence data for the akodontine tribe. Biological Journal of the Linnean Society, 50, 149–177. https://doi.org/10.1111/j.1095-8312.1993.tb00924.x
Smith, M.F. & Patton, J.L. (1999) Phylogenetic relationship and the radiation of sigmodontine rodents in South America: evidence from cytochrome b. Journal of Mammalian Evolution, 6, 89–128. https://doi.org/10.1023/A:1020668004578
Smithe, F.B. (1975) Naturalist’s color guide. American Museum of Natural History, New York.
Tate, G.H.H. (1939) The mammals of the Guiana region. Bulletin of the American Museum of Natural History, 76, 151–229.
Thomas, O. (1882) On a collection of rodents from north Peru. Proceedings of the Zoological Society of London, 50, 98–111. https://doi.org/10.1111/j.1096-3642.1882.tb02730.x
Thomas, O. (1926) The Godman-Thomas expedition to Peru III. On mammals collected by Mr RW Hendee in the Chachapoyas region of north Peru. Annals and Magazine of Natural History, including Zoology, Botany and Geology, London, 18 (9), 156–167. https://doi.org/10.1080/00222932608633489
Tinoco, N.O. (2015) Caracterización molecular, morfológica y morfométrica del complejo Nephelomys albigularis Tomes, 1860 (Rodentia: Cricetidae), y su distribución en el Ecuador. Disertación previa a la obtención del título de Licenciado Ciencias Biológicas, Pontificia Universidad Católica del Ecuador. Quito, Ecuador. Available from: http://repositorio.puce.edu.ec/handle/22000/9667 (accessed 4 January 2019)
Tirira, D.G. & Boada, C.E. (2009) Diversidad de mamíferos en bosques de Ceja Andina alta del nororiente de la provincia de Carchi, Ecuador. Boletín Técnico, 8, Serie Zoológica, 4–5, 1–24.
Tomes, R.F. (1860) Notes on a third collection of Mammalia made by Mr. Fraser in the Republic of Ecuador. Proceedings of the Zoological Society of London, 1860, 260–268.
Trouessart, E.L. (1897) Catalogus mammalium tam viventium quam fossilium. Fasciculus III. Rodentia II (Myomorpha, Histricomorpha, Lagomorpha). R. Friedlander & Sohn, 1, Berolini, vi + 664 pp. https://doi.org/10.5962/bhl.title.130824
Uturunco, A.C. & Pacheco, V. (2016) Redescripción de Eremoryzomys polius (Rodentia: Cricetidae) y descripción de una nueva especie del género Eremoryzomys. Mastozoología Neotropical, 23, 483–503.
Venables, W.N. & Ripley, B.D. (2002) Modern applied statistics with S. 4th Edition. Springer, New York, xi + 495 pp. https://doi.org/10.1007/978-0-387-21706-2
Venegas, P.J., Townsend, J.H., Koch, C. & Böhme, W. (2008) Two new sympatric species of leaf-toed geckos (Gekkonidae: Phyllodactylus) from the Balsas region of the upper Marañón Valley, Peru. Journal of Herpetology, 42, 386–396. https://doi.org/10.1670/07-1341.1
Vorontsov, N.N. (1959) The system of hamster (Cricetinae) in the sphere of the world fauna and their phylogenetic relations. Bjulleten’ Moskovskogo Obščestva Ispytatelej Prirody Otdel biologičeskij, 64, 134–37.
Voss, R.S. (1988) Systematics and ecology of ichthyomyine rodents (Muroidea): Patterns of morphological evolution in a small adaptive radiation. Bulletin of the American Museum of Natural History, 188, 259–493.
Voss, R.S. (1991) An introduction to the Neotropical muroid rodent genus Zygodontomys. Bulletin of the American Museum of Natural History, 210, 1–113.
Voss, R.S. & Carleton, M.D. (1993) A new genus for Hesperomus molitor Winge and Holochilus magnus Hershkovitz (Mammalia, Muridae) with an analysis of its phylogenetic relationships. American Museum Novitates, 3085, 1–39.
Vuilleumier, F. (1969) Pleistocene speciation in birds living in the high Andes. Nature, 223, 1179–1180. https://doi.org/10.1038/2231179a0
Wagner, J.A. (1843) s.n. In: Die säugethiere in abbildungen nach der natur mit beschreibungen Dr. Johann Christian Daniel von Schreber, Supplemental 3, 398.
Weberbauer, A. (1945) El mundo vegetal de los Andes peruanos, estudio fitogeográfico. Estación Experimental Agrícola de la Molina, Dirección de Agricultura, Ministerio de Agricultura, Lima, 776 pp.
Weksler, M. (2006) Phylogenetic relationships of oryzomyine rodents (Muridae: Sigmodontinae): separate and combined analyses of morphological and molecular data. Bulletin of the American Museum of Natural History, 296, 1–149. https://doi.org/10.1206/0003-0090(2006)296[0001:PROORM]2.0.CO;2
Weksler, M., Percequillo, A.R. & Voss, R.S. (2006) Ten new genera of oryzomyine rodents (Cricetidae: Sigmodontinae). American Museum Novitates, 3537, 1–30. https://doi.org/10.1206/0003-0082(2006)3537[1:TNGOOR]2.0.CO;2
Winger, B.M. & Bates, J.M. (2015) The tempo of trait divergence in geographic isolation: Avian speciation across the Marañon Valley of Peru. Evolution, 69, 772–787. https://doi.org/10.1111/evo.12607
Zhang, J., Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29, 2869–2876. https://doi.org/10.1093/bioinformatics/btt499