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Dermatemys mawii (The Hicatee, Tortuga Blanca, or Central American River Turtle): A Working Bibliography
Venetia Briggs-Gonzalez, Sergio C. Gonzalez, Dustin Smith, Kyle Allen, Thomas R. Rainwater, and Frank J. Mazzotti

Caribbean Naturalist, Special Issue No. 2 (2018):1–22

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Caribbean Naturalist 1 V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 Dermatemys mawii (The Hicatee, Tortuga Blanca, or Central American River Turtle): A Working Bibliography Venetia Briggs-Gonzalez1,*, Sergio C. Gonzalez1, Dustin Smith2, Kyle Allen1, Thomas R. Rainwater3, and Frank J. Mazzotti1 Abstract - Dermatemys mawii (Central American River Turtle), locally known in Belize as the “Hicatee” and in Guatemala and Mexico as Tortuga Blanca, is a large, highly aquatic freshwater turtle that has been extirpated from much of its historical range of southern Mexico, northern Guatemala, and lowland Belize. Throughout its restricted range, Dermatemys has been intensely harvested for its meat and eggs and sold in local markets. Despite being formally protected throughout their range, most D. mawaii populations have disappeared in Guatemala and Mexico and continue to decline in Belize because of intensive overhunting, which now includes commercial harvesting. As the last remaining species of the family Dermatemydidae, the Hicatee is one of the top 25 most exploited turtles worldwide and is classified as critically endangered (facing an extremely high risk of extinction in the near future) by the IUCN and listed on Appendix II of CITES. Countrywide surveys in Belize in 2010 confirmed D. mawaii populations are severely reduced in most areas, particularly in habitats more accessible to humans. These findings prompted the formation of the National Hicatee Conservation and Monitoring Network (NHCMN) in Belize in 2011. Through the NHCMN, plans were outlined for increased research, conservation, and educational efforts countrywide. Currently, relatively few data are available regarding several aspects of the life history of D. mawaii, including growth, behavioral and feeding ecology, and reproductive biology. The purpose of this working bibliography is to provide a comprehensive list of literature pertaining to D. mawaii with the goal of increasing scientific research, regulatory law enforcement, educational awareness, and species conservation. Introduction Dermatemys mawii (Central American River Turtle), locally known in Belize as the “Hicatee” and in Guatemala and Mexico as Tortuga Blanca, is a large, highly aquatic freshwater turtle (Figs. 1, 2) found along the coastal lowlands of southern Mexico, northern Guatemala, and Belize (Alvarez del Toro 1982; Ernst and Barbour 1989; Iverson 1992; Iverson and Mittermeier 1980; Lee 1996; Legler and Vogt 2013; TTWG 2014, 2017; Vogt et al. 2011) and is the only extant representative of the family Dermatemydidae (Iverson and Mittermeier 1980). Currently D. mawii is classified as critically endangered by the International Union for Conservation of Nature (IUCN 2016), listed as endangered under the provisions of the US 1University of Florida, Fort Lauderdale Research and Education Center, 3205 College Avenue, Fort Lauderdale, FL 33314, USA. 2North Carolina Zoo, 4401 Zoo Parkway, Asheboro, NC 27205, USA. 3Tom Yawkey Wildlife Center and Belle W. Baruch Institute of Coastal Ecology and Forest Science, Clemson University, PO Box 596, Georgetown, SC 29442, USA. *Corresponding author - vsbriggs@ufl.edu. Manuscript Editor: Don Moll Endangered and Threatened Species of the Caribbean Region 2018 CARIBBEAN NATURALIST Special Issue No. 2:1–22 Caribbean Naturalist V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 2 Figure 1. Female Dermatemys mawii (Hicatee) captured and released in Northern Belize, Lamanai Field Research Center. Figure 2. Male Dermatemys mawii (Hicatee) in breeding color, captured and released in Central Belize. Caribbean Naturalist 3 V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 Endangered Species Act (USFWS 1983), and listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES 2017). It is currently considered one of the top 25 most endangered turtle species in the world (Turtle Conservation Coalition 2011). Dermatemys mawaii has been and continues to be intensely exploited throughout its restricted range for its meat and eggs (Moll 1986; Polisar 1994, 1995), with populations having collapsed over recent decades and a very slow recovery, if any, in some protected areas (IUCN 2016). Local populations have become extirpated in Mexico and Guatemala, and Belize remains the only stronghold for the species (Campbell 1998, Rainwater et al. 2012). However, with a minimum 10-year generation time and a conservative annual take of 5%, a drastic population decline of 81.5% within 3 generations has been estimated (IUCN 2016). Current harvesting rates which now include commercial take, particularly of reproductive adults, will likely reduce the population even further and drive this species to extinction more rapidly (Polisar 1997, Rainwater et al. 2012, IUCN 2016). In Belize, a comprehensive survey of D. mawaii was conducted in 1983 and 1984 (Moll 1986) that found the species was still common to abundant in areas sparsely populated by humans but declining in more-developed areas where turtles were more accessible to hunters. Further research conducted from 1989 to 1991 indicated that exploitation of D. mawaii persisted in the more populated areas of northern Belize and that the level of harvesting was not sustainable (Polisar 1992, 1994, 1995, 1997; Polisar and Horwich 1994). As a result, in 1993 the Belize Fisheries Department drafted nationwide comprehensive legislation for the protection and management of D. mawaii that included year-round possession limits, a brief closed (non-hunting) season, a complete prohibition on selling and purchasing D. mawaii, and a series of protected zones in the major waterways of northern Belize (Polisar 1994, 1995, 1997; Polisar and Horwich 1994). In 2010, another countrywide survey of D. mawaii found populations were severely reduced in most areas, but small populations existed in a few remote areas where some level of protection exists and human activity was minimal (Rainwater et al. 2012). In several localities where D. mawaii once existed in large numbers, these turtles were by then uncommon or rare, and continued hunting still targeted large, reproductive adults, further endangering existing populations (Rainwater et al. 2012). These findings prompted the formation of the National Hicatee Conservation and Monitoring Network (NHCMN) in Belize in 2011. The NHCMN consists of government and non-governmental agencies, scientists, students, and naturalists with the goals of education and outreach, legislation and enforcement, and science. As a result, over the last 6 years additional surveys have been conducted across the country by local research centers, such as the Lamanai Field Research Center and Toledo Institute for Development and the Environment, and field studies have been initiated (V. Briggs-Gonzalez et al., unpubl. data; Requeña et al. 2015, Smith 2015). Regular surveys of local markets have been implemented to detect and deter illegal sales of D. mawaii. Law enforcement has increased, resulting in several arrests and turtle confiscations (Fig. 3). Additionally, annual countrywide educational awareness campaigns have been launched (Fig. 4; McLoughlin 2013), particularly during Caribbean Naturalist V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 4 Figure 3. Eight live Dermatemys mawii (Hicatee) confiscated by Belize Fisheries Department officers. Figure 4. Hicatee (Dermatemys mawii) sticker distributed nationwide to promote educational awareness in Belize. Caribbean Naturalist 5 V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 times of the year when Hicatee are harvested and consumed as part of specific traditional and cultural events. A Hicatee Conservation Research Center has been established to examine the reproductive biology of Hicatee as well as various aspects of the species’ captive husbandry (Rainwater et al. 2011). Since 2011, legislation has been revised to reflect the initiative of the NHCMN to include a closed season from May 1st to 31st, a reduced harvest limit of 3 turtles per person or 5 per vehicle, catch size limits of females no greater than 43 cm and no less than 38 cm, and the prohibition to sell or purchase D. mawaii (Government of Belize 2011). Furthermore, nets are prohibited for use in capture, and Hicatee captures are prohibited in a list of specified areas that include parts of the Belize River, New River, Sibun River, Rio Bravo, Cox and Mucklehany Lagoons, headwaters of Mussel Creek, Northern and Southern Lagoons and tributaries (Government of Belize 2011). Poaching still occurs, but with increased enforcement efforts and greater local involvement, there seems to have been a decrease (McLoughlin 2013; R. Quintana, Belize Fisheries Department, Belize City, Belize, pers. comm.; Rainwater et al. 2011, 2012). The purpose of this working bibliography is to provide a comprehensive list of literature pertaining to Dermatemys with the goal of increasing scientific research, regulatory law enforcement, educational awareness, and species conservation. Published works on D. mawaii are limited, but information is also available in reports and pilot studies that exist in gray literature. We conducted exhaustive literature searches using Google Scholar and Web of Science with “Dermatemys” and “turtle” used as keywords, in addition to Google searches for “Hicatee” and “tortuga blanca”; particularly for gray literature. Authors of more recently published work were contacted to provide additional papers that were not accessible. Papers, theses, and reports are organized under relevant categories. In several instances, D. mawaii is not the target species, but important information concerning this species is included in the reference. Hicatee Research Bibliography Taxonomy and Species Description Alvarez del Toro, M. 1982. Los Reptiles de Chiapas, 3rd Edition. Instituto de Historia Natural del Estado de Chiapas. Tuxtla Gutíerrez, Chiapas, México. Bienz, A. 1896. Dermatemys mawii Gray, eine osteologische Studie mit Beitragen zur Kenntnis der Schildkroten. Revue Suisse de Zoologie 3:61–135. Bonin, F., B. Devaux, and A. Dupré. 2006. Turtles of the World. P. Pritchard (Transl.). The Johns Hopkins University Press, Baltimore, MD, USA. Boulenger, G.A. 1889. Dermatemydidae. Pp. 27–30, In G.A. Boulenger (Ed.) Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum of Natural History, London, UK. Bourque, J., J. Hutchison, P. Holroyd, and J. Bloch. 2008. A new kinosternoid (Testudines: Dermatemydidae) from the Paleocene–Eocene boundary of the Bighorn Basin, Wyoming, and its paleoclimatological implications. Journal of Vertebrate Paleontology 28 (3) (Supplement):55A. Caribbean Naturalist V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 6 Campbell, J. 1998. Amphibians and Reptiles of Northern Guatemala, the Yucatan, and Belize. University of Oklahoma Press, Norman, OK, USA. Cope, E.D. 1865. Third contribution to the herpetology of tropical America. Proceedings of the Academy of Natural Sciences of Philadelphia 17:185–198. de Borre, A.P. 1869. Description d'un jeune individu de la Dermatemys mawii, espèce Américaine de la famille des Élodites. Académie Royale de Belgique 28:177–121. Duméril, A.M.C., and G. Bibron. 1851. Emys areolata, Emys berardii, Cinosternon leucostomum, Cinosternon cruentatum. In A.M.C. Duméril and A.H.A. Duméril (Eds.) Catalogue Methodique de la Collection des Reptiles (Museum d’Histoire Naturelle de Paris). Gide and Baudry, Paris, France. Ernst, C.H., and R.W. Barbour. 1989. Turtles of the World. Smithsonian Institution Press, Washington, DC, USA. Frair, W. 1979. Taxonomic relations among sea turtles elucidated by serological tests. Herpetologica 35:239–244. Gray, J.E. 1847. Description of a new genus of Emydidae. Proceedings of the Zoological Society of London 1847:55–56. Gray, J.E. 1855. Description of a new genus and some new species of tortoises. Annals and Magazine of Natural History 15:67–69. Gray, J.E. 1856b [“1855”]. Catalogue of the Shield Reptiles in the Collection of the British Museum. Part I. Testudinata (Tortoises). British Museum, London, UK. Gray, J.E. 1864. Additional observations on Dermatemys: A genus of Emydidae from Central America. Annals and Magazine of Natural History 14:391–392. Gray, J.E. 1870. On the family Dermatemydae, and a description of a living species in the gardens of the Society. Proceedings of the Zoological Society of London 1870:711–716. Gray, J.E. 1872. Appendix to the Catalogue of Shield Reptiles in the Collection of the British Museum. Part II. Emydisaurians, Rhynchocephalia, and Amphisbaenians. British Museum, London, UK. Gray, J.E. 1873. Hand-list of the specimens of shield reptiles in the British Museum. British Museum, London, UK. Holman, A.J. 1963. Observations on dermatemydid and staurotypine turtles from Veracruz, Mexico. Herpetologica 19:277-279. Hutchison, J., and D. Bramble. 1981. Homology of the plastral scales of the Kinosternidae and related turtles. Herpetologica 37:73–85. Iverson, J.B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. J.B. Iverson Publishers, Richmond, IN, USA. Iverson, J.B., and R.A. Mittermeier. 1980. Dermatemydidae, Dermatemys. Catalogue of American Amphibians and Reptiles 237:1–4. Iverson, J.B., R.M. Brown, T.S. Akre, T.J. Near, M. Le, R.C. Thomson, and D.E. Starkey. 2007. In search of the tree of life for turtles. Pp. 85–106, In H.B. Shafer, N.N. FitzSimmons, A. Georges, and A.G.H. Rhodin (Eds.). Defining turtle diversity: Proceedings of a workshop on genetics, ethics, and taxonomy of freshwater turtles and tortoises. Chelonian Research Monographs No. 4. Caribbean Naturalist 7 V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 Knauss, G.E., W.G. Joyce, T.R. Lyson, and D. Pearson. 2011. A new kinosternoid from the Late Cretaceous Hell Creek formation of North Dakota and Montana and the origin of the Dermatemys mawii lineage. Paläontologische Zeitschrift 85:125–142. Köhler, G. 2008. Reptiles of Central America, 2nd Revised Edition. Herpeton Verlag, Germany. Lee, J.C. 1996. The Amphibians and Reptiles of the Yucatan Peninsula. Cornell University Press, Ithaca, NY, USA. Lee, J. 2000. A Field Guide to the Amphibians and Reptiles of the Maya World: The Lowlands of Mexico, Northern Guatemala, and Belize. Cornell University Press, Ithaca, NY, USA. Lee, R.C. 1969. Observing the Tortuga Blanca. International Turtle and Tortoise Society Journal 3:32–34. Legler, J.M., and R.C. Vogt. 2013. The Turtles of Mexico: Land and Freshwater Forms. University of California Press, Berkeley, CA, USA. McDowell, S.B. 1964. Partition of the genus Clemmys and related problems in the taxonomy of aquatic Testudinidae. Proceedings of the Zoological Society of London 143:239–279. Méndez, M.O. 2012. Distribución, caracterización de hábitat, e historia natural para la tortuga blanca (Dermatemys mawii) en el Área de Usos Múltiples de Río Sarstún, Izabal. Tesis de Licenciatura en Biología. Universidad del Valle de Guatemala, Guatemala City, Guatemala. Mittermeier, R. 1970. Turtles in Central American markets. International Turtle and Tortoise Society Journal 4:20–26. Neill, W.T., and R. Allen. 1959. Studies on the amphibians and reptiles of British Honduras. Publications of the Research Division, Ross Allen’s Reptile Institute 2:1–76. Polisar, J. 1990. An inventory of the turtle fauna of the Rio Bravo Conservation and Management Area: A preliminary report. Programme for Belize, Belize City, Belize. 12 pp. Pritchard, P. 1979. Encyclopedia of Turtles. TFH Publications, Inc., Neptune, NJ, USA. Savage, J.M. 1982. The enigma of the Central American herpetofauna: Dispersal or vicariance? Annals of the Missouri Botanical Garden 69:464–547. Schmidt, K.P. 1941. The Amphibians and Reptiles of British Honduras. Field Museum of Natural History, Chicago, IL. Shaffer, H.B., N.N. FitzSimmons, A. Georges, and A.G.J. Rhodin. (Eds.). 2007. Defining turtle diversity: Proceedings of a workshop on genetics, ethics, and taxonomy of freshwater turtles and tortoises. Chelonian Research Monographs No. 4. Smith, H.M., and L.F. James. 1958. The taxonomic significance of cloacal bursae in turtles. Transactions of the Kansas Academy of Sciences 61:86–96. Smith, H.M., and R.B. Smith. 1979. Synopsis of the Herpetofauna of Mexico: Vol. 6. Guide to the Mexican Turtles. John Johnson, North Bennington, VT, USA. Caribbean Naturalist V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 8 Smith, H.M., and E.H. Taylor. 1950. Type localities of Mexican reptiles and amphibians. University of Kansas Science Bulletin 33:313–380. Stafford, P.J., and J.R. Meyer. 2000. A Guide to the Reptiles of Belize. Academic Press, San Diego, CA, USA. Stuart, L.C. 1963. A checklist of the herpetofauna of Guatemala. Miscellaneous Publications of the Museum of Zoology, University of Michigan, No. 122, Ann Arbor, MI, USA. 150 pp. Stuart, L.C. 1963. A study of the herpetofauna of the Uaxactun-Tikal area of northern El Petén, Guatemala. Contributions from the Laboratory of Vertebrate Biology, University of Michigan 75:1–30. Turtle Taxonomy Working Group [J.W. Bickham, J.B. Iverson, J.F. Parham, H.D. Philippen, A.G.J. Rhodin, H.B. Shaffer, P.Q. Spinks, and P.P. van Dijk.]. 2007. 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Shaffer, and R. Bour]. 2014. Turtles of the World, 7th Edition: Annotated checklist of taxonomy, synonymy, distribution with maps, and conservations status. Pp. 329–479, In A.G.J. Rhodin, P.C.H. Pritchard, P.P. van Dijk, R.A. Saumure, K.A. Buhlmann, J.B. Iverson, and R.A. Mittermeier (Eds.).Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/ SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs No. 5. Turtle Taxonomy Working Group [A.G.J. Rhodin, J.B. Iverson, R. Bour, U. Fritz, A. Georges, H.B. Shaffer, and P.P. van Dijk]. 2017. Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status (8th Edition.). Pp. 1-292, In: A.G.J. Rhodin, P.C.H. Pritchard, P.P. van Dijk, R.A. Saumure, K.A. Buhlmann, J.B. Iverson, and R.A. Mittermeier (Eds.). Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs 7. Vogt, R.C., J.L.B. Villareal, and G. Perez-Higareda. 1997. Lista anotada de anfibios y reptiles. Pp. 507–522, In S.E. Gonzalez, R. Dirzo, and R.C. Vogt (Eds.).Historia natural de Los Tuxtlas. Universidad Nacional Autonoma de México, México. Caribbean Naturalist 9 V. Briggs-Gonzalez, S.C. Gonzalez, D. Smith, K. Allen, T.R. Rainwater, and F.J. Mazzotti 2018 Special Issue No. 2 Vogt, R.C., G.P. Gonzalez-Porter, and P.P. van Dijk. 2006. Dermatemys mawii. (errata version published in 2016) IUCN Red List of Threatened Species 2006:e. T6493A97409830. Vogt, R.C., J.R. Polisar, D. Moll, and G. Gonzalez-Porter. 2011. Dermatemys mawii Gray 1847—Central American River Turtle, Tortuga Blanca, Hickatee. Pp. 058.1–058.12, In A.G.J. Rhodin, P.C.H. Pritchard, P.P. van Dijk, R.A. Saumure, K.A. Buhlmann, J.B. Iverson, and R.A. 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