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(Caiman Crocodilus) in Puerto Rico Management of Biological Invasions (2016) Volume 7, Issue 3: 287–296 DOI: http://dx.doi.org/10.3391/mbi.2016.7.3.08 Open Access © 2016 The Author(s). Journal compilation © 2016 REABIC Research Article Diet of the non-native spectacled caiman (Caiman crocodilus) in Puerto Rico Damien R. Bontemps1,*, Elvira Cuevas1, Eileen Ortiz2, Joseph M. Wunderle, Jr.3 and Rafael L. Joglar1 1Department of Biology, University of Puerto Rico—Río Piedras Campus, PO Box 23360, San Juan, Puerto Rico 2Departamento de Recursos Naturales y Ambientales, PO Box 366147 San Juan, Puerto Rico 3International Institute of Tropical Forestry, USDA Forest Service, Sabana Field Research Station, HC 02, Box 6205, Luquillo, Puerto Rico E-mail addresses: [email protected] (DRB), [email protected] (EC), [email protected] (EO), [email protected] (JMW), [email protected] (RLJ) *Corresponding author Received: 1 September 2015 / Accepted: 28 January 2016 / Published online: 18 March 2016 Handling editor: Matthew A. Barnes Abstract The spectacled caiman (Caiman crocodilus) was introduced to Puerto Rico over 50 years ago with the Tortuguero Lagoon Natural Reserve (TLNR) as its epicenter, where it is now established as an apex predator. Although concerns have been raised regarding the potential impact of this naturalized predator on Puerto Rico’s native fauna, little was known of the caiman’s diet on the island. Therefore this study was conducted to determine the diet of the spectacled caiman and its potential impact on island animals. For this study, measurements were obtained from 138 caimans across all life stages (12–94 cm snout–vent length; SVL) from October 2014 to May 2015 within the TLNR. Stomach contents were retrieved and analyzed based on prey category occurrence frequency. In addition, caiman muscle samples were obtained to determine their nitrogen and carbon isotopic signature. Insects were the most abundant prey items encountered with 90.7% and 68.8% in hatchling (SVL < 20 cm) and juvenile (SVL = 20–59.9 cm) stomach respectively. In adult (SVL > 60 cm) caimans, fish remains were the most significant prey items with 38.3% frequency of occurrence. Fish, insects, and gastropods were the only categories of ten designated prey categories to show significant variation among the three caiman age classes. This study provides novel information on dietary habits of spectacled caimans in Puerto Rico relevant to the design of management strategies. Key words: diet analysis, stomach content, stable isotopes, exotic apex predator, crocodilian Introduction of effects on prey in other geographical locations, correlative studies of predator/prey population The introduction and successful establishment of dynamics, experimental removal, and dietary studies apex predators has been widely documented as (Park 2004). The latter method has been widely an important cause of biodiversity loss. Invasive employed by means of food extraction and top predators have been found to disrupt food stomach content analysis (SCA) across a variety webs at multiple trophic levels, modify habitats, of taxa (Michael Anthony et al. 2000; Pierce et out-compete native species for resources or to al. 2004; Kidera 2008; Somaweera et al. 2011). serve as vectors for disease and parasite transmission Although these methods contribute to our (Work et al. 2002; Levy et al. 2008; Mortensen understanding of a predator’s diet, they do have et al. 2008; Dorcas et al. 2011). The lack of co- some disadvantages. For example, the extensive evolutionary history with non-native species and sample size, labor, cost, and time required to limited ability to geographically disperse render obtain dietary information via SCA makes it a some native species from insular ecosystems challenging technique to use, particularly when particularly vulnerable to the effects of introduced studying a predator feeding on a wide variety of species (D’Antonio and Dudley 1998; Simberloff prey (Pethybridge et al. 2011; Bowen and Iverson 2000; Wiles et al. 2003). 2013). Time is of the essence when dealing with Common methods to assess potential ecological recent introductions. In spite of failure to prevent impacts of introduced predators include observations the spread of an exotic, an early detection of the 287 D.R. Bontemps et al. potential effects can play a major role in the within these areas (Joglar et al. 2010). This study careful design of management strategies and revealed a drastic range expansion since its first establishing priorities for mitigating the threat. appearance in the TLNR and presented numerous Advances in the field of stable isotope analysis reports on caiman sightings and captures at (SIA) have proven to be advantageous in the various localities within Puerto Rico’s coastal prediction of foraging impacts in invasive top plain and its satellite island of Vieques. predators (Bodey et al. 2011). Whereas stomach Caiman crocodilus has been extensively studied content analysis (SCA) reveals recently ingested in its native range including population dynamics, prey items or a snapshot of the diet, SIA provides diet, breeding, and harvesting studies (Staton and long-term dietary habits. It can be performed in a Dixon 1977; Gorzula 1978; Ayarzagüena 1983; timely fashion in order to quickly respond to the Thorbjarnarson 1993; Allsteadt 1994; Thorbjarnarson threat. Albeit SIA is deficient in providing high and Velasco 1999; Moreno-Arias et al. 2012; Laverty and Dobson 2013). In Puerto Rico, two taxonomic resolutions of consumed prey species, unpublished population studies have been conducted it reveals the relative diet source proportion with (Santos Reyes 1988; Joglar et al. 2010). However, a priori knowledge of the prey species inhabiting despite having been established on the island for the novel environment. Especially in the context over 50 years, there have been no thorough of a generalist predator, where prey categories ecological studies of the caiman’s effects on the rather than specific prey species are to be native fauna. Therefore, our study was initiated assessed, SIA provides relevant information to to address the need for new information on the predict direct dietary effects. introduced caiman’s foraging behavior in Puerto The spectacled caiman Caiman crocodilus Rico. The objectives of our study were to use (Linnaeus, 1758), also referred to as common stomach content analysis to provide baseline data caiman, is a medium-sized crocodilian considered on caimans’ diet in Puerto Rico and to make an opportunistic omni-carnivorous species as its recommendations for caiman management, based diet includes a wide array of invertebrates, fish, on the findings of the diet analysis. amphibians, reptiles, birds, and even mammals for larger specimens (Thorbjarnarson 1993). Less Methods commonly known, part of their diet may include plant material such as seeds, leaves, or even fruits Study area (Platt et al. 2013). The spectacled caiman has been introduced and is considered fully established in Caiman crocodilus proliferation in Puerto Rico Florida (USA) and Isla de la Juventud (Cuba) (Ellis appears to have initiated from the Tortuguero 1980; Estrada and Ruibal 1999). In the 1960’s, it Lagoon which was designed as a natural reserve was introduced in Puerto Rico principally associated in 1971 and currently under the management of with the legal caiman pet trade during these the DNER (Schwartz and Henderson 1991; Thomas times (Thomas and Joglar 1996). A combination and Joglar 1996). It is located between the munici- of deliberate and accidental release from caiman palities of Vega Baja and Manatí (N 18.46328 pet owners is likely to be the cause of introduction and W -66.43962; Figure 1). The reserve contains to the natural environment. Thomas and Joglar the largest freshwater lagoon in Puerto Rico (1996) documented the presence of Caiman covering an approximate surface area of 2.43 km2. crocodilus in the northern coastal plains of the The lagoon holds a water volume of approxi- island with the Tortuguero Lagoon Natural Reserve mately 2.68 million m3 with a mean depth of 1.2 m. (TLNR) as its source of proliferation. It is fed by subterranean, mainly fresh water sources In 1985, after an increasing number of sightings in addition to precipitation (approx. 1,600 mm/year) within the TLNR and surrounding areas, the and flushed about 7.5 times per year through a Department of Natural and Environmental Resources canal built in 1940 leading to the sea on the north- (DNER) implemented a management plan to east side (Quiñones-Márquez and Fusté 1978). Other eradicate the species from the island. Since then, output includes the estimated evapotranspiration the frequency of caiman sightings and encounters rate of 1,274 mm per year. Water temperatures has increased far beyond the TLNR. Authorities oscillate from 24 ºC to 31 ºC in accord with and inhabitants of the San Juan Bay Estuary seasonal changes with the absence of vertical became concerned with the caiman’s presence. As temperature gradients. The wet and dry season a result, a rapid assessment survey was performed occur from May to November and from December to estimate population abundance and distribution to April respectively. 288 Diet of the non-native spectacled caiman Figure 1. Map of Puerto Rico displaying the location of the Tortuguero Lagoon (in red) including a satellite photographic image. The lagoon’s surrounding habitat composition 59.9 cm), sub-adults/adult females (Size Class III includes swamps, marshes, coastal shrubs, and hills. = 60–89.9 cm), and adult males (Size Class IV ≥ Among the 717 plants species recorded from the 90 cm). Spring scales of 5 kg, 10 kg, and 20 kg site are 144 endemic and/or endangered species. maximums were used to record the weight of Consequently it is considered the fourth most larger individuals. A Denver Instrument XL-3100D important flora in Puerto Rico. As one of the only electric scale (accurate to 0.01g) was used to two natural reservoirs of the island, the Tortuguero measure the mass of smaller caimans, mainly Lagoon is of great ecological importance due to hatchlings.
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