A new subspecies of Conepatus semistriatus (Boddaert, 1784) (Mammalia, Carnivora, Mephitidae) from Venezuela, and the first known case of insular dwarfism in living skunks

Una subespecie nueva de Conepatus semistriatus (Boddaert, 1784) (Mammalia, Carnivora, Mephitidae) de Venezuela, y el primer caso conocido de enanismo insular en mofetas vivientes

• Jesús Molinari Universidad de Los Andes, Mérida, Venezuela. https://orcid.org/0000-0002-9393-5483
• María R. Abarca-Medina Instituto Venezolano de Investigaciones Científicas, Caracas, Venezuela. https://orcid.org/0000-0002-5079-5134
• Belkis A. Rivas-Rodríguez Universidad de Los Andes, Mérida, Venezuela. https://orcid.org/0000-0002-3801-7989

DOI: https://doi.org/10.5281/zenodo.13755829

Keywords: Caribbean islands, endemism, geographic variation, island rule, Margarita Island, morphometry, Striped Hognosed Skunk

Abstract

The Striped Hog-nosed Skunk, Conepatus semistriatus (Boddaert, 1784), is distributed from southern Mexico to northern Peru. It is one of the least studied carnivorans of the New World. Not even basic morphometric information exists for the species. This study assesses the taxonomic status of the Striped Hog-nosed Skunk occurring on Margarita Island, off the coast of eastern Venezuela. Multivariate analyses of 25 cranial measurements revealed the Margaritan skunk to be much dwarfed with respect to its nearby mainland congener, thus complying with the ‘island rule’, which predicts that under insular conditions large continental animals become smaller, and small continental animals become larger. The craniometric differences between the Margaritan and mainland skunks are of similar or greater magnitude to those found in interspecific comparisons within genera of several carnivoran families. Geographic variation in cranial size was not evident within the Margaritan sample, or within the mainland sample, analyzed separately. The Margaritan form is sufficiently distinct to be deemed a new subspecies, thus it is here formally described as C. semistriatus elieceri ssp. nov. This skunk becomes the seventh mammal known to be endemic to Margarita Island. Its conservation status is undetermined.

References

Adler, G. H. & R. Levins. 1994. The island syndrome in rodent populations. Quarterly Review of Biology 69: 473–490. https://doi.org/10.1086/418744

ASM [American Society of Mammalogists]. 2024. Mammal diversity database, version 1.12.1. https://doi.org/10.5281/zenodo.10595931

Baeckens, S. & R. Damme. 2020. The island syndrome. Current Biology 30: 338–339. https://doi.org/10.1016/j. cub.2020.03.029

Belant, J. L., J. Schipper & J. Conroy. 2009. The conservation status of small carnivores in the Americas. Small Carnivore Conservation 41: 3–8.

Bertrand, A. S. & A. Morisot. 2012. Neotropical spotted cat species discrimination using morphometrics. Natureza & Conservação 10: 40–44. https://doi.org/10.4322/natcon.2012.007

Bisbal, F. J. 1983. Dos nuevos mamíferos para la Isla de Margarita, Venezuela. Acta Científica Venezolana 34: 366–367.

Boher-Bentti, S. & G. A. Cordero-Rodríguez. 2000. Distribution of brown capuchin monkeys (Cebus apella) in Venezuela: a piece of the puzzle. Neotropical Primates 8: 152–153.

Boher-Bentti, S., M. Salazar-Candelle & C. Ferreira-Marques. 2023. Mamíferos de Venezuela: lista actualizada 2023 y comentarios taxonómicos. Anartia 36: 7–35. https://doi. org/10.5281/zenodo.10433912

Bornholdt, R., K. Helgen, K. P. Koepfli, L. Oliveira, M. Lucherini & E. Eizirik. 2013. Taxonomic revision of the genus Galictis (Carnivora: Mustelidae): Species delimitation, morphological diagnosis, and refined mapping of geographical distribution. Zoological Journal of the Linnean Society 167: 449–472. https://doi.org/10.1111/j.1096-3642.2012.00859.x

Cabrera, A. 1958. Catálogo de los mamíferos de América del Sur. Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Ciencias Zoológicas 4: 1–308.

Castillo, D. F. & N. C. Caruso. 2024. Potential distribution and conservation of the hognosed skunk (genus Conepatus, Mammalia: Mephitidae). Journal for Nature Conservation 77: 126519. https://doi.org/10.1016/j.jnc.2023.126519

Coonan, T. J., C. A. Schwemm & D. K. Garcelon. 2010. Decline and recovery of the island fox a case study for population recovery. Cambridge, United Kingdom: Cambridge University Press, xi + 212 p.

Cuarón, A. D., D. Valenzuela-Galván, D. García-Vasco, M. E. Copa, S. Bautista, H. Mena, D. Martínez-Godínez, C. González-Baca, L. A. Bojórquez-Tapia, L. Barraza, P. C. De Grammont, F. Galindo-Maldonado, M. A. Martínez-Morales, E. Vázquez-Domínguez, E, Andresen, J. Benítez-Malvido, D. Pérez-Salicrup, K. W. McFadden & M. E. Gompper. 2009. Conservation of the endemic dwarf carnivores of Cozumel Island, Mexico. Small Carnivore Conservation 41: 15–21.

Dragoo, J. W., R. L. Honeycutt & D. J. Schmidly. 2003. Taxonomic status of white-backed hognosed skunks, genus Conepatus (Carnivora: Mephitidae). Journal of Mammalogy 84: 159–176. https://doi.org/10.1644/1545- 1542(2003)084%3C0159:TSOWBH%3E2.0.CO;2

Dragoo, J. W. & S. R. Sheffield. 2009. Conepatus leuconotus (Carnivora: Mephitidae). Mammalian Species 827: 1–8. https://doi.org/10.1644/827.1

Emmons, L. H. 2005. A revision of the genera of arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae), with descriptions of two new genera. pp. 247–310. In: Lacey, E. A. & P. Myers (eds). Mammalian diversification: from chromosomes to phylogeography. Berkeley, USA: University of California Publications in Zoology. https://doi.org/10.1525/califor- nia/9780520098534.003.0009

Ferguson, A. W., R. E. Strauss & R. C. Dowler. 2022. Beyond black and white: Addressing colour variation in the context of local environmental conditions for the aposematic North American Hognosed skunk. pp. 107–130. In: Do Linh San, E., J. J. Sato, J. L. Belant & M. J. Somers (eds). Small carnivores: Evolution, ecology, behaviour, and conservation. New York, USA: Wiley.

Floyd, C. H., D. H. Van Vuren, K. R. Crooks, K. L. Jones, D. K. Garcelon, N. M. Belfiore, J. W. Dragoo & B. May. 2011. Genetic differentiation of island spotted skunks, Spilogale gracilis amphiala. Journal of Mammalogy 92: 148–158. https:// doi.org/10.1644/09-MAMM-A-204.1

Foster, J. B. 1964. Evolution of mammals on islands. Nature 202: 234–235. https://doi.org/10.1038/202234a0

García‐Perea, R. 1996. Patterns of postnatal development in skulls of lynxes, genus Lynx (Mammalia: Carnivora). Journal of Morphology 229: 241–254. https://doi.org/10.1002/ (S I C I)1097-4687(199609)229:3%3C241:: A I D - JMOR1%3E3.0.CO;2-1

Gompper, M. E., A. E. Petrites & R. L. Lyman. 2006. Cozumel Island fox (Urocyon sp.) dwarfism and possible divergence history based on subfossil bones. Journal of Zoology 270: 72–77. https://doi.org/10.1111/j.1469-7998.2006.00119.x

Groves, C. P. 2001. Primate Taxonomy. Washington, USA: Smithsonian Institution Press, 350 pp.

Gutiérrez, E. E. & J. Molinari. 2008. Morphometrics and taxonomy of bats of he genus Pteronotus (subgenus Phyllodia) in Venezuela. Journal of Mammalogy 89: 292–305. https:// doi.org/10.1644/06-MAMM-A-452R.1

Hall, E. R. 1981. The mammals of North America, 2nd ed. New York, USA: Wiley, xv + vi + 1181 pp.

Hammer, Ø. 2024. PAST, Paleontological Statistics, Version 4.16, Reference manual. Oslo, Norway: University of Oslo.

Hernández-Sánchez, A., A. Santos-Moreno & G. Pérez-Irineo. 2022. The Mephitidae in the Americas: A review of the current state of knowledge and future research priorities. Mammalian Biology 102: 307–320. https://doi.org/10.1007/ s42991-022-00249-z

Hershkovitz, P. 1959. Nomenclature and taxonomy of the Neo- tropical mammals described by Olfers, 1818. Journal of Mammalogy 40: 337–353. https://doi.org/10.2307/1376558

Jolicoeur, P. & J. R. Mosimann. 1960. Size and shape variation in the painted turtle: A principal component analysis. Growth 24: 339–354.

Lichter, M., D. Zviely, M. Klein & D. Sivan. 2010. Sea-level changes in the Mediterranean: Past, present, and future –a review. pp. 3–17. In: Seckbach, J., R. Einav & A. Israel (eds). Seaweeds and their role in globally changing environments (cellular origin, life in extreme habitats and astrobiology). Dordrecht, Netheerlands: Springer. https://doi.org/10.1007/978- 90-481-8569-6_1

Linares, O. J. 1998. Mamíferos de Venezuela. Caracas, Venezuela: Sociedad Conservacionista Audubon de Venezuela, 691 pp. Lyras, G. A., A. A. Van Der Geer & L. Rook. 2010. Body size of insular carnivores: evidence from the fossil record. Journal of Biogeography 37: 1007–1021. https://doi.org/10.1111/j.1365-2699.2010.02312.x

McDonough, M. M., A. W. Ferguson, R. C. Dowler, M. E. Gompper & J. E. Maldonado. 2022. Phylogenomic systematics of the spotted skunks (Carnivora, Mephitidae, Spilogale): Additional species diversity and Pleistocene climate change as a major driver of diversification. Molecular Phylogenetics and Evolution 167: 107266. https://doi.org/10.1016/j. ympev.2021.107266

McFadden, K. W. 2004. The ecology, evolution, and natural history of the dwarf carnivores of Cozumel Island, Mexico. New York, USA: Columbia University, 148 pp. [PhD Thesis]

McGarigal, K., S. Cushman & S. Stafford. 2000. Multivariate statistics for wildlife and ecology research. New York, USA: Springer, xiii + 283 pp.

Mead, R. A. 1967. Age determination in the spotted skunk. Journal of Mammalogy 48: 606–616. https://doi. org/10.2307/1377584

Meiri, S., T. Dayan & D. Simberloff. 2004. Body size of insular carnivores: Little support for the island rule. American Naturalist 163: 469-479. https://doi.org/10.1086/382229

Meza-Joya, F. L., E. Ramos, F. Cediel, V. Martínez-Arias, J. Colmenares & D. Cardona. 2018. Predicted distributions of two poorly known small carnivores in Colombia: The greater grison and striped hognosed skunk. Mastozoología Neotropical 25: 89–105. https://doi.org/10.31687/ saremMN.18.25.1.0.09

Miller, A. 2015. Skunk. London, United Kingdom: Reaktion Books, 199 pp.

Molinari, J. 2007. Variación geográfica en los venados de cola blanca (Cervidae, Odocoileus) de Venezuela, con énfasis en O. margaritae, la especie enana de la Isla de Margarita. Memoria de la Fundación La Salle de Ciencias Naturales 167: 29–72.

Molinari, J. 2023a. A global assessment of the ‘island rule’ in bats based on functionally distinct measures of body size. Journal of Biogeography 50: 1179–1190. https://doi.org/10.1111/ jbi.14624

Molinari, J. 2023b. A barebones scheme to choose between the species, subspecies, and ‘evolutionarily significant unit’ categories in taxonomy and conservation. Journal for Nature Conservation 72: 126335. https://doi.org/10.1016/j. jnc.2023.126335

Molinari, J., E. E. Gutiérrez & B. K. Lim. 2023. Systematics and biogeography of Anoura cultrata (Mammalia, Chiroptera, Phyllostomidae): A morphometric, niche modeling, and genetic perspective, with a taxonomic reappraisal of the genus. Zootaxa 5297: 151–188. https://doi.org/10.11646/ zootaxa.5297.2.1

Moscarella, R. A. 2001. Filogeografía y genética de la conservación del venado caramerudo de Venezuela. Caracas, Venezuela: Universidad Simón Bolívar, 87 pp. [MSc Thesis]

Pacheco, V., S. Diaz, L. Graham-Angeles, M. Flores-Quispe, G. Calizaya-Mamani, D. Ruelas & P. Sánchez-Vendizú. 2021. Lista actualizada de la diversidad de los mamíferos del Perú y una propuesta para su actualización. Revista Peruana de Biología 28: e21019. http://dx.doi.org/10.15381/rpb. v28i4.21019

Patten, M. A., & P. Unitt. 2002. Diagnosability versus mean differences of Sage Sparrow subspecies. Auk 119: 26–35. https://doi.org/10.1093/auk/119.1.26

Pilgrim, G. E. 1933. A fossil skunk from Samos. American Museum Novitates 663: 1–15.

Rodrigues, M. L. F. 2013. História evolutiva de Conepatus (Carnivora: Mephitidae): padrões biogeográficos de diversificação, investigação filogenética e revisão taxonómica do gênero. Porto Alegre, Brazil: Pontifícia Universidade Católica do Río Grande do Sul, 162 pp. [PhD Thesis]

Rossi, R. V., R. S. Voss & D. P. Lunde. 2010. A revision of the didelphid marsupial Genus Marmosa Part 1. The species in Tate’s ‘mexicana’ and ‘mitis’ sections and other closely related forms. Bulletin of the American Museum of Natural History 334: 1–83. https://doi.org/10.1206/334.1

Ruíz-García, M., D. Chacón, T. Plese, I. Schuler & J. M. Shostell. 2017. Mitogenomics phylogenetic relationships of the current sloth’s genera and species (Bradypodidae and Megalonychidae). Mitochondrial DNA Part A 29: 281–299. http://dx.doi.org/10.1080/24701394.2016.1275602

Sánchez-Villagra, M. R., Geiger, M., & R. A. Schneider. 2016. The taming of the neural crest: a developmental perspective on the origins of morphological covariation in domestic cated mammals. Royal Society Open Science 3: 160107. http:// dx.doi.org/10.1098/rsos.160107

Schiaffini, M. I., M. Gabrielli, F. J. Prevosti, Y. P. Cardoso, D. Castillo, R. Bo, E. Casanave & M. Lizarralde. 2013. Taxonomic status of southern South American Conepatus (Carnivora: Mephitidae). Zoological Journal of the Linnean Society 167: 327–344. https://doi.org/10.1111/zoj.12006

Siddall, M., J. Chappell & E. K. Potter. 2006. Eustatic sea level during past interglacials. pp. 75–92. In: Sirocko, F., M. Claussen, T. Litt & M. F. Sanchez-Goni (eds). The climate of past interglacials. Amsterdam, The Netherlands: Elsevier. https://doi.org/10.1016/S1571-0866(07)80032-7

Simpson, G. G. 1951. The species concept. Evolution 5: 285– 298. https://doi.org/10.1111/j.1558-5646.1951.tb02788.x

Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Miscellaneous Publication, Museum of Natural History, University of Kansas 56: 1–132.

Smith, J. D. & H. H. Genoways. 1974. Bats of Margarita island, Venezuela, with zoogeographic comments. Bulletin of the Southern California of Sciences 73: 64–79.

Solari, S., Y. Muñoz-Saba, J. V. Rodríguez-Mahecha, T. R. Defler, H. E. Ramírez-Chaves & F. Trujillo. 2013. Riqueza, endemismo y conservación de los mamíferos de Colombia. Mastozoología Neotropical 20: 301–365.

Srinivas, Y. & Y. Jhala. 2021. Morphometric variation in wolves and golden jackal in India (Mammalia, Carnivora). Biodiversity Data Journal 9: e67677. https://doi.org/10.3897/ BDJ.9.e67677

Taylor, M. E. & J. Matheson. 1999. A craniometric comparison of the African and Asian mongooses in the genus Herpestes (Carnivora: Herpestidae). Mammalia 63: 449–464. https:// doi.org/10.1515/mamm.1999.63.4.449

Teta, P., G. D’Elía, P. Jayat, G. S. Libardi, J. A. Oliveira, R. Moratelli, A. R. Percequillo, J. R. Prado, P. E. Ortiz, N. Hurtado, M. E. Schiaffini, E. F. Abreu, Jr., E. A. Chiquito, L. Giménez & J. Torres. 2020. On the distinction and availability of the new taxa proposed by Agnolin et al., (2019). Mastozoología Neotropical 27: 155–171. https://doi. org/10.31687/saremMN.20.27.1.0.20

Tirira, D. G., J. Brito, S. F. Burneo, C. M. Pinto & J. A. Salas. 2023. Mamíferos del Ecuador: lista oficial actualizada de especies. Quito, Ecuador: Asociación Ecuatoriana de Mastozoología, 83 pp.

van der Geer, A. A. E. 2019. Effect of isolation on coat colour polymorphism of Polynesian rats in Island Southeast Asia and the Pacific. PeerJ 7:e6894. http://doi.org/10.7717/ peerj.6894

Van Gelder, R. G. 1959. A taxonomic revision of the spotted skunks (genus Spilogale). Bulletin of the American Museum of Natural History 117: 229–392.

Van Gelder, R. G. 1968. The genus Conepatus (Mammalia, Mustelidae): variation within a population. American Museum Novitates 2322: 1–37.

Van Valen, L. 1973. Body size and numbers of plants and animals. Evolution 27: 27–35. https://doi. org/10.1111/j.1558-5646.1973.tb059 14.x

Van Valkenburgh, B. & R. K. Wayne. 1994. Shape divergence associated with size convergence in sympatric East African jackals. Ecology 75: 1567–1581. https://doi. org/10.2307/1939618

Villa-Meza, A., R. Avila-Flores, A. D. Cuarón & D. Valenzuela- Galván. 2011. Procyon pygmaeus (Carnivora: Procyonidae). Mammalian Species 877: 87–93. https://doi.org/10.1644/877.1 Vivo, M. & A. P. Carmignotto. 2015. Family Sciuridae G. Fischer, 1817. pp. 1–48. In: Patton, J. L., U. F. Pardiñas & G. D’Elía (eds). Mammals of South America, volume 2. Rodents. Chicago, USA: University of Chicago Press.

Waelbroeck, C., L. Labeyrie, E. Michel, J. C. Duplessy, J. F. Mc- Manus, K. Lambeck, E. Balbon & M. Labracherie. 2002. Sea-level and deep water temperature changes derived from benthonic foraminifera isotopic records. Quaternary Science Reviews 21: 295–305. https://doi.org/10.1016/S0277-
3791(01)00101-9

Wang, X. & Z. Qiu. 2004. Late Miocene Promephitis (Carnivora, Mephitidae) from China. Journal of Vertebrate Paleontology 24: 721–731. https://doi.org/10.1671/0272- 4634(2004)024[0721:LMPCMF]2.0.CO;2

Wang, X., D. P. Whistler & G. T. Takeuchi. 2005. A new basal skunk Martinogale (Carnivora, Mephitinae) from late Miocene Dove Spring Formation, California, and origin of new world mephitines. Journal of Vertebrate Paleontology 25: 936–949. https://doi.org/10.1671/0272- 4634(2005)025[0936:ANBSMC]2.0.CO;2

Wilson, D. E. 1991. Mammals of the Tres Marías Islands. Bulletin of the American Museum of Natural History 206: 214– 250.

Wozencraft, W. C. 2005. Order Carnivora. pp. 532–562. In: Wilson, D. E. & D. M. Reeder (eds) Mammal species of the world: A taxonomic and geographic reference. Washington, USA: Smithsonian Institution Press, 2142 pp.