No. 48
Caribbean Naturalist
2018
Invasive Rats (Rattus sp.), but not always Mice (Mus musculus), are Ubiquitous at All Elevations and Habitats within the Caribbean National Forest, Puerto Rico Aaron B. Shiels and Gabriela E. Ramírez de Arellano
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Cover Photograph: A Black Rat (Rattus rattus) consuming seeds of the native tree Guarea guidonia in the Caribbean National Forest, PR, USA. Image captured at night using a trail camera. © Aaron B. Shiels and Gabriela E. Ramírez de Arellano.
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CARIBBEAN NATURALIST
James D. Ackerman, Department of Biology, University of Puerto Rico at Río Piedras, USA Alfonso Aguilar-Perera, Department of Marine Biology, Universidad Autónoma de Yucatán, Mexico Wayne J. Arendt, International Institute of Tropical Forestry, Luquillo, Puerto Rico, USA Rüdiger Bieler, Field Museum of Natural History, Chicago, IL, USA Christopher P. Bloch, Department of Biological Sciences, Bridgewater State University, Bridgewater, MA, USA William R. Buck, Institute of Systematic Botany, New York Botanical Garden, Bronx, NY, USA Leo Douglas, Department of Geography/Geology, University of the West Indies, Mona, Jamaica Robert Erdman, Department of Biological Sciences, Florida Gulf Coast University, Fort Myers, FL, USA Keith Goldfarb, GoldRush Science Services, Steuben, ME, USA ... Editor-in-Chief Grizelle González, International Institute of Tropical Forestry, San Juan, Puerto Rico, USA Gary R. Graves, Department of Vertebrate Zoology, Smithsonian Institution, Washington, DC, USA Scott Jones, Smithsonian Institution, Caribbean Coral Reef Ecosystems, Carrie Bow Cay, Belize Heather Judkins, Department of Biological Sciences, University of South Florida, St. Petersburg, FL, USA Craig A. Layman, Department of Applied Ecology, North Carolina State University, Raleigh, NC, USA John Leavengood, USDA-APHIS-PPQ, Tampa Bay, FL Joerg-Henner Lotze, Eagle Hill Institute, Steuben, ME, USA ... Publisher Antonio A. Mignucci-Giannoni, Manatee Conservation Center, Inter American University, Bayamón, Puerto Rico, USA Gregg Moore, Department of Biological Sciences, Jackson Estuarine Laboratory, University of New Hampshire, Durham, NH, USA Dawn Phillip, Department of Life Sciences, University of The West Indies, St. Augustine, Trinidad and Tobago... Managing Editor Robert Powell, Department of Biological Sciences, Avila University, Kansas City, MO, USA Chris Rimmer, Vermont Center for Ecostudies, Norwich, VT, USA Noris Salazar Allen, Smithsonian Tropical Research Institute, Panama Amy Siuda, Collegium of Natural Sciences, Eckerd College, St. Petersburg, FL, USA J. Angel Soto-Centeno, Rutgers University, Department of Biological Sciences, Newark, NJ, USA David W. Steadman, Florida Museum of Natural History, Gainesville, FL, USA Kathleen Sullivan Sealey, Department of Biology, University of Miami, Coral Gables, FL, USA Jarrod M. Thaxton, Department of Biology, University of Puerto at Mayagüez, USA Jason M. Townsend, Hamilton College, Biology Department, Clinton, NY, USA Jill Weber, Eagle Hill Institute, Steuben, ME, USA ... Production Editor Byron Wilson, Department of Life Sciences, University of the West Indies at Mona, Kingston, Jamaica Graham A. J. Worthy, Department of Biology, University of Central Florida, Orlando, FL, USA Joseph M. Wunderle, International Institute of Tropical Forestry, University of Puerto Rico at Río Píedras, USA
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2018 2018
Caribbean Naturalist CARIBBEAN NATURALIST A.B. Shiels and G.E. Ramírez de Arellano
No. 48 No. 48:1–14
Invasive Rats (Rattus sp.), but not always Mice (Mus musculus), are Ubiquitous at All Elevations and Habitats within the Caribbean National Forest, Puerto Rico Aaron B. Shiels1,* and Gabriela E. Ramírez de Arellano2 Abstract - Invasive rodents, particularly rats (Rattus spp.), occupy >80% of the world’s islands and are among the greatest threats to native biodiversity and agriculture on islands. At the time of their introduction in the 1500s, there was at least 1 native rat species in Puerto Rico. Today there are no native rodents remaining in Puerto Rico, but R. norvegicus (Norway Rat) may be found in urban settings, and R. rattus (Black Rat) are the most common rat across the island including within natural areas, and invasive Mus musculus (House Mouse) may also be found in urban and non-urban habitats. The Caribbean National Forest (CNF; locally El Yunque) in northeastern Puerto Rico has some native and endangered species vulnerable to rat predation. The objective of our study was to determine the presence and distribution of invasive rodents (rats and mice) across elevations and habitats within the CNF. We used 104 tracking tunnels, which are baited ink cards placed in tunnels so that foot prints of animal visitors could be identified, to determine presence of invasive rodent species. We placed 3 tracking tunnels at each 50-m elevation-gain (n = 66 total tunnels), on the edge of forest habitat from sea level to 1070 m at El Yunque peak along the main road (Highway 191) through the CNF. We established additional tracking tunnels (n = 38) in the major habitats in the CNF, including treefall and hurricane gaps, landslides, stream edges, and continuous forest. House Mice had not been previously reported in the CNF, and were found only at the forest edge along Highway 191 at elevations of 50–150 m and 300–1070 m, whereas rats (Rattus sp.) were found at all elevations and in all habitat types sampled. Logistic regressions revealed that mice and rat presence each increases with elevation (mice: P = 0.0352, rat: P = 0.0019), though total rodent presence did not. Knowledge of the habitat types and elevations that these invasive rodents occupy can inform management strategies for rodent control and native species protection.
Introduction Rodents (Rattus spp. and Mus musculus L. [House Mouse]) have been introduced to many ecosystems worldwide and are among the most widespread and problematic invasive animals affecting islands (Angel et al. 2009, Shiels et al. 2014, Towns et al. 2006). Through mostly unintentional introductions by humans, these rodents occupy >80% of the islands worldwide (Atkinson 1985, Towns 2009). In addition to having negative impacts on agricultural resources, invasive rodents are a threat to many native species on islands. Invasive House Mice are known to USDA, National Wildlife Research Center, 4101 LaPorte Avenue, Fort Collins, CO 80521, USA. 2University of Puerto Rico at Mayagüez, Biology Department, Mayagüez, PR 00681, USA. *Corresponding author -
[email protected].
1
Manuscript Editor: Byron Wilson 1
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Caribbean Naturalist A.B. Shiels and G.E. Ramírez de Arellano
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damage and depredate plants and insects, whereas invasive rats R. rattus L. (Black Rat), and R. norvegicus Berkenhout (Norway Rat) are known to consume seabirds, forest birds, snails, arthropods, and a variety of plants including large-seeded species (Shiels and Drake 2011, Shiels et al. 2013, Towns et al. 2006). Due to their high fecundity, lack of effective predators, and generalist diets, invasive rodent populations rapidly expand on islands, and few habitats and ecosystems in urban and natural areas are free of at least 1 of these rodent species (Drake and Hunt 2009, Shiels 2010). Puerto Rico currently has 3 invasive rodent species: Black Rats, Norway Rats, and House Mice. At the time of non-native rodent introductions in the 1500s, there was at least 1 native rat species in Puerto Rico: Heteropsomys insulans Anthony (Insular Cave Rat or Spiny Rat). Turvey et al. (2007) and others hypothesize that extinctions of a native shrew and the native Spiny Rat were driven by predation and/ or competition with introduced Black Rats in Puerto Rico, rather than massive-scale deforestation for sugarcane (not commencing until the late 1700s) or the introduction of Herpestes javanicus (É. Geoffroy Saint-Hilaire) (Small Indian Mongoose), which did not occur until 1877. In addition to the Spiny Rat, there were at least 2 much larger native rodent species in Puerto Rico: Elasmodontomys obliquus Anthony (Platetoothed Giant Hutia), which has been estimated to have had a mean body weight of 13.7 kg (McFarlane 1999), and a newly described large arboreal species (Tainotherium valei; Turvey et al. 2006). Isolobodon portoricensis J.A. Allen (Puerto Rican Hutia) was another rodent species that has been described in the prehistoric fauna, and this rodent was introduced to Puerto Rico, presumably as a food source, from nearby Hispanola by pre-Columbian Amerindians (Turvey et al. 2007). All of these native or non-European rodent species are extinct. However, the presence of native rodents in Puerto Rico may have been beneficial to the contemporary native flora and fauna, as theory predicts that islands that lacked native rodents have native biota that are more likely to suffer herbivory and predation from introduced rodents than are biota on islands with native rodents (Drake and Hunt 2009). Black Rats are currently documented as the most widespread invasive rodent species in Puerto Rico, as they occupy urban, suburban, and natural areas (De Leön 1964, Miranda-Castro et al. 2000, Weinbren et al. 1970). In the Caribbean National Forest (CNF) in northeastern Puerto Rico, Black Rats previously have been the only rodent species documented, and this rodent is a threat to many native species including the endangered Amazona vittata Boddaert (Puerto Rican Parrot; Engeman et al. 2006, Snyder et al. 1987, Zwank and Layton 1989). Norway Rats are generally restricted to urban areas and possibly agricultural settings in Puerto Rico (De Leön 1964, Miranda-Castro et al. 2000), whereas House Mice distributions are not well known in Puerto Rico but have been documented in and around suburban areas (Tamsitt and Fox 1970). The CNF is often described as a patchwork of forest habitats that reflect different types of disturbances (e.g., treefalls, hurricanes, landslides, flooding, drought) in varying states of recovery (Brokaw et al. 2012, Shiels and Walker 2013). In addition to gaining a better understanding of how invasive rodents are distributed 2
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along the elevational gradient in the CNF, there is additional interest in discovering rodent establishment and activity within the many habitat types in the CNF. Prior to our study, Black Rats, which are the most common rodent species in island forests (Shiels et al. 2014), were documented in the CNF within continuous forest at ~300– 600 m elevation (Engeman et al. 2006, Weinbren et al. 1970). Rodents are often in areas of relatively high plant cover presumably to limit their exposure to predators (Arthur et al. 2005, Cox et al. 2000, King et al. 1996); therefore, older gaps with established vegetation ground-cover rather than young (e.g.,
