Caretta caretta (North West Atlantic subpopulation)
|Scientific Name:||Caretta caretta (North West Atlantic subpopulation)|
|Species Authority:||(Linnaeus, 1758)|
See Caretta caretta
|Red List Category & Criteria:||Least Concern ver 3.1|
|Assessor(s):||Ceriani, S.A. & Meylan, A.B.|
|Reviewer(s):||Casale, P., Wallace, B.P. & Pilcher, N.J.|
|Contributor(s):||Godfrey, M., Pate, M., Dodd, M., Brost, B., Ingram, D., Zurita, J.C., Tzee, M., Herrera, R., Arenas, A., Witherington, B.E., Casale, P., Owens, D., Eckert, K., Epperly, S., Guada, H., Musick, J., Pilcher, N.J. & Wallace, B.P.|
The North West Atlantic subpopulation breeds mostly in the southeastern U.S. and on the Yucatán Peninsula in Mexico. Its marine habitats extend throughout all the north and central Atlantic Ocean (northwest, northeast, central west and central east) (Fig. 2 in the Supplementary Material).NMFS and USFWS (2008) identified five recovery units within the North West Atlantic subpopulation based on genetic differences and a combination of geographic distribution of nesting densities, geographic separation, and geopolitical boundaries. The first four recovery units represent nesting assemblages in the southeast U.S.: the Northern (southern Virginia to Florida/Georgia border); Peninsular Florida (Florida/Georgia border south and then north through Pinellas County on the west coast, excluding the islands west of Key West, Florida); Dry Tortugas (islands west of Key West, Florida); and the Northern Gulf of Mexico (Franklin County, Florida, west through Texas). The fifth recovery unit (the Greater Caribbean) includes all other nesting assemblages within the Northwest Atlantic (Mexico through French Guiana, the Bahamas, and Lesser and Greater Antilles); nesting is mostly concentrated on the Yucatán peninsula of Mexico. Even though several genetic nesting stocks have been identified within this subpopulation (NMFS andUSFWS 2008; Shamblin et al. 2012), metapopulation dynamics support its designation as a single subpopulation, or regional management unit (Wallace et al. 2010),and a single distinct population segment under the US Endangered Species Act (Conant et al. 2009).
The available long-term series of annual nest counts (used as an index of population abundance) shows an overall increase over the past three generations for the North West Atlantic Loggerhead subpopulation. Moreover, both geographic distribution and population size are large. Therefore, the North West Atlantic Loggerhead subpopulation is considered Least Concern under current IUCN Red List criteria. This “Least Concern” status should be considered as entirely conservation-dependent, because the current population is the result of decades of intense conservation programs, especially at nesting sites (e.g., NMFS and USFWS 2008, TEWG 2009, Ehrhart et al. 2014) and in some parts of its marine range, through the use of Turtle Excluder Devices, and the cessation of these programs would likely be followed by a population decrease. However, due to the long maturation time of Loggerheads and the available indices of abundance (nest counts), the subpopulation would probably not qualify for a threatened category within five years following the cessation of conservation programs, as would be needed to trigger the Near Threatened category (IUCN 2014).
The analysis of time series datasets with ≥10 years of annual nest counts at 86 nesting sites (representing four of the five Recovery Units identified by NMFS and USFWS 2008) revealed different nesting trends within the Northwest Atlantic Loggerhead subpopulation. We examined nesting trends for the overall North West Atlantic Loggerhead subpopulation and for each of the four Recovery units that had sufficient data for the present analysis. An adequate time series was not available for the Dry Tortugas Recovery Unit; however, this Recovery Unit has been estimated to account for less than 1% of the North West Atlantic Loggerhead subpopulation (NMFS and USFWS 2008). We found an overall subpopulation increase relative to subpopulation size three generations ago (criterion A2; Table 1 in the Supplementary Material). At the Recovery unit level, the data indicated an overall nesting trend increase for the Northern and Greater Caribbean Recovery Units and a slight decrease for the Florida Peninsular and the Northern Gulf of Mexico Recovery Units (Table 1 in the Supplementary Material). We also applied criteria B, C, and D to the North West Atlantic subpopulation, but the extent of occurrence (EOO) and area of occupancy (AOO) values exceeded the thresholds of criterion B, and the subpopulation did not trigger any threatened category of criteria C and D.
No population viability analysis (criterion E) was available so the North West Atlantic Loggerhead subpopulation assessment was conducted by applying criteria A-D.
For marine turtles, annual counts of nesting females and their nesting activities (more often the latter) are the most frequently recorded and reported abundance metric across index monitoring sites, species, and geographic regions (National Research Council 2010). To apply Criterion A, three generations (or a minimum of ten years, whichever is longer) of abundance data are required (IUCN 2014). In the case of the Loggerhead, we conservatively estimated its generation time as 45 years (see the Habitats & Ecology section). For criteria A1-A2, data from three generations ago (~135 yrs) are necessary to estimate population declines beginning three generations ago to the present (i.e., assessment) year. The challenges of this requirement for long-lived species like marine turtles—with generation lengths of 30 yrs or more—are obvious (see Seminoff and Shanker 2008 for a review). Abundance data from ~135 yrs ago are not available for Loggerheads anywhere in the world. Extrapolating backward using population trends based on current datasets was considered inappropriate because estimates produced would be biologically unrealistic and unsubstantiated. In the absence of better information, we assumed that population abundance three generations ago (~135 years, one generation estimated 45 yrs; see teh Habitats & Ecology section) was similar to the first observed abundance rather than assuming that the population has always been in a decline (or increase) of the same magnitude as in the current generation. A similar approach was used in the Red List assessment of another marine turtle, the Leatherback Turtle Dermochelys coriacea (Wallace et al. 2013), and of another long-lived, geographically widespread taxon, the African Elephant (Blanc 2008). Thus, to apply criterion A we assumed that the abundance at the beginning of an available time series dataset had not changed significantly in the preceding three generations, and therefore used the same abundance value in trend calculations. For the Loggerhead global and subpopulation assessments we only considered time series datasets of ≥10 yrs.
For the North West Atlantic Loggerhead subpopulation, we included datasets of ≥10 yrs from 86 index nesting beaches in two countries (United States and Mexico; Table 1 in the Supplementary Material). Data from Mexico were limited to the state of Quintana Roo (Yucatán), which, together with the U.S. data, serves as a good representation of the Northwest Atlantic subpopulation because breeding mostly occurs in the southeastern U.S. and on the Yucatán Peninsula in Mexico (NMFS and USFWS 2008). The number of nests in the datasets used in the trend analysis (86 index nesting sites) represented 63.4% of the estimated total number of nests for the whole subpopulation. The total number of nests from these index sites was 53,043 nests yr-1 in the most recent counts, while the total number of nests―from index sites as well as sites without long-term datasets―in the North West Atlantic subpopulation is estimated to have been >83,717 nests yr-1. The latter estimate was based on the 5-year (2009-2013) average of total nest numbers recorded in the southeast U.S. and Mexico (North Carolina Wildlife Research Commission (NCWRC) unpublished data; South Carolina Department of Natural Resources (SCDNR) unpublished data; Georgia Department of Natural Resources (GDNR) unpublished data; Florida Fish and Wildlife Conservation Commission/Fish and Wildlife Research Institute (FFWC/FWRI) unpublished data; U.S. Fish and Wildlife Service (USFWS) unpublished data; Sea Turtle Protection Committee of Quintana Roo, Mexico (CPTMQROO) unpublished data). Data on nest numbers from Cancun (Quintana Roo, Mexico; approximately 3.7% of nests in Quintana Roo) were not available for 2013 and were not included in the calculation of the 5-year average of total nest numbers for the North West Atlantic subpopulation. Given the relatively high level of geographic coverage represented by the 86 index beaches with long-term datasets, we think the data support a robust evaluation of the status of this subpopulation based on IUCN Red List criteria. However, we urge caution in the interpretation of the results for reasons discussed in the section Sources of Uncertainty.
We calculated annual and overall trends using past and present abundance estimates representing the annual average of 5-yr nest counts for each of the 86 index nesting beaches within the subpopulation, as well as for each of the Recovery Units included (four of five). We then calculated the overall subpopulation past trend. The most recent year for which abundance data were available across all index nesting sites (except one) was 2013. Time series data ended in 2008 for Pritchard Island and, therefore, we estimated population size at this site through 2013 based on the population trend for existing years.
Trends ranged from negative to positive across the different index nesting beaches, with most negative trends observed in the Florida Peninsular Recovery Unit (35% of the 23 Florida index nesting beaches included in the analysis showed negative trends). At the Recovery unit level – i.e., all nests at index sites within the unit combined – the data indicated an overall increase of 35% for the Northern Recovery Unit, a 2% decrease for the Florida Peninsular Recovery Unit, a 1% decrease for the Northern Gulf of Mexico Recovery Unit, and a 53% increase for the Greater Caribbean Recovery Unit (Table 1). The overall trend for the North West Atlantic subpopulation was positive (+2%).
Thus, with no overall observed decline on the index beaches included in the analysis, the North West Atlantic Loggerhead subpopulation did not qualify for any threatened category under criterion A. It should be noted, however, that declines in loggerhead nesting populations are suspected to have occurred elsewhere in the Caribbean (e.g., Venezuela, Guada et al. 2014, Colombia, Amorocho 2003).
Since the subpopulation area is the entire North West Atlantic, the EOO exceeds the threat category threshold (20,000 km²). The AOO for marine turtles is identified as the nesting beach habitat, which represents the smallest habitat for a critical life stage. The total length of known Loggerhead nesting beaches in the North West Atlantic is estimated to be >2,610 km (NCWRC unpublished data; SCDNR unpublished data; GDNR unpublished data; FFWC/FWRI unpublished data; USFWS unpublished data; CPTMQROO unpublished data). This value is an underestimation of the total length of available nesting habitat as it does not include the Bahamas, Cuba and other islands in the Greater Caribbean. Since the appropriate scale for AOO is a grid 2x2 km, the above linear measure was converted to 5,220 km². This value exceeded the threatened category threshold (2,000 km²). In conclusion, the subpopulation did not trigger any of the thresholds and options for a threatened category under criterion B.
To apply criterion C, we used the population estimate of Richards et al. (2011) for the whole North West Atlantic subpopulation, which was based on a sophisticated modelling approach incorporating available nest counts (2001-2010) and demographic reproductive parameters (clutch frequency, breeding interval, adult female survival probability). The nest count data used by Richards et al. (2011) included many of the same 86 index nesting beaches used in our criterion A analysis. Their population estimation technique was designed to incorporate uncertainty in the parameters used to extrapolate nests to adult females. To be conservative, we used their minimum estimate for the Northwest Atlantic adult female Loggerhead subpopulation over the decade from 2001 to 2010, which was 30,096 adult females. Because there is a lack of accurate data on adult sex ratios of Loggerheads, we decided not to extrapolate to the total mature population. In any case, the estimated number of adult females in the Northwest Atlantic subpopulation is well above 10,000 individuals and, therefore, criterion C was not met. The North West Atlantic subpopulation did not meet any other parameter threshold, i.e., continuing decline, number of mature individuals in one subpopulation, and extreme fluctuations in the number of mature individuals. In conclusion, the subpopulation did not trigger any of the thresholds and options for a threatened category under criterion C.
The number of mature individuals (see criterion C) and AOO value (see criterion B) exceeded the respective thresholds. In conclusion, the subpopulation did not trigger any of the thresholds and options for a threatened category under criterion D.
Sources of Uncertainty
Although marine turtle nest counts are the most common metric recorded and reported across sites and species globally, there are several disadvantages to using them as a proxy in determining overall population dynamics (National Research Council 2010). First, because nesting females are a very small proportion of a marine turtle population, using their abundance and nesting activities as proxies for overall population abundance and trends requires knowledge of other key demographic parameters to allow proper interpretation (National Research Council 2010). However, there is still considerable uncertainty about many of these fundamental demographic parameters for Loggerheads, including age at maturity, generation length, survivorship across life stages, adult and hatchling sex ratios, and conversion factors among reproductive parameters (clutch frequency, nesting success, remigration intervals, etc). These values can vary among subpopulations and recovery units, further complicating the process of combining subpopulation abundance and trend estimates to obtain global population abundance and trend estimates, and contributing to the uncertainty in these estimates. Second, despite the wide availability of nesting abundance data for marine turtles, monitoring effort and methodologies can vary within and across study sites, complicating comparison of nesting count data. However, we have reduced uncertainty by including in the analyses only datasets obtained through highly standardized monitoring. Another source of uncertainty is that the Red List trend evaluation methods followed here involve simple comparisons of early and later 5-yr-average annual nest counts. Which 5-yr intervals are used in the analyses can have a large effect on the outcome. For example, had the assessment of the North West Atlantic Loggerhead subpopulation been undertaken in 2008, after nine consecutive years of declining nest counts on Florida beaches, the trend analysis would likely have yielded different results. The reasons for the steep decline on Florida beaches, and for the subsequent reversal of this trend, remain unknown, adding to the uncertainty associated with the prediction of trends in sea turtle populations. Ideally, sophisticated models should be applied to have a higher chance of detecting complex and cryptic trends. The time interval used for trend evaluation also varies widely among beaches because of the differences in the monitoring history. Lastly, the assumption implicit in criteria A1 and A2 that population abundance three generations ago (135 yrs) was similar to the first observed abundance might not be valid, given the nearly complete lack of historical abundance data on this subpopulation and the evidence that other marine turtle species in this region have undergone significant declines during the past century (Meylan and Donnelly 1999).
Recognizing these caveats, the North West Atlantic Loggerhead subpopulation is very well monitored with data from 86 index nesting beaches available for use in this subpopulation assessment and has contributed a great amount of information about the biology of the species. The dataset represents 63.4%of the total estimated annual nests of the subpopulation. Therefore, we believe that the results of the present analysis are representative of the nesting beaches as a whole and, hence, the overall trend.
For further reading on sources of uncertainty in marine turtle Red List assessments, see Seminoff and Shanker (2008).
We are extremely grateful to the following people who provided nesting data on behalf of their agencies and organizations: Matthew Godfrey, North Carolina Wildlife Research Commission (NCWRC); Michelle Pate, South Carolina Department of Natural Resources (SCDNR), Mark Dodd, Georgia Department of Natural Resources (GDNR); Beth Brost, Florida Fish and Wildlife Conservation Commission/Fish and Wildlife Research Institute (FWCC/FWRI); Dianne Ingram, U.S. Fish and Wildlife Service, Alabama (USFWS); Julio Zurita, Miriam Tzee, Roberto Herrera, and Alejandro Arenas, Sea Turtle Protection Committee of Quintana Roo (CPTMQROO), Mexico.The collection of the nesting data included in these analyses would not be possible without the dedication and hard work of thousands of permit holders and volunteers who carried out the beach surveys.We also thank Blair Witherington and Paolo Casale for their guidance on the Red List process and David Owens for lending his expertise on sex ratios of Northwest Atlantic loggerhead populations. We appreciate the constructive comments received from the following colleagues: K. Eckert, S. Epperly, H. Guada, J. Musick, N. Pilcher and B. Wallace.
|Range Description:||The Loggerhead Turtle has a worldwide distribution across subtropical and temperate regions of the Mediterranean Sea and Pacific, Indian, and Atlantic oceans (Wallace et al. 2010) (Figure 1 in the Supplementary Material). Collectively, the North West Atlantic Ocean hosts the most significant nesting assemblage of Loggerheads in the Western Hemisphere and is one of the two largest Loggerhead nesting assemblages in the world (Ehrhart et al. 2003). The North West Atlantic subpopulation breeds mostly along the coast of the United States and Mexico and its marine habitats extend throughout all the north and central Atlantic Ocean (northwest, northeast, central west and central east).|
Native:Anguilla; Aruba; Bahamas; Barbados; Belize; Bermuda; Bonaire, Sint Eustatius and Saba; Cayman Islands; Colombia; Costa Rica; Cuba; Curaçao; Dominican Republic; French Guiana; Grenada; Guadeloupe; Guatemala; Haiti; Honduras; Jamaica; Mexico; Montserrat; Nicaragua; Panama; Puerto Rico; Saint Barthélemy; Saint Kitts and Nevis; Saint Lucia; Saint Vincent and the Grenadines; Suriname; Trinidad and Tobago; Turks and Caicos Islands; United States; Venezuela, Bolivarian Republic of; Virgin Islands, British; Virgin Islands, U.S.
|FAO Marine Fishing Areas:|
Atlantic – western central; Atlantic – northwest; Atlantic – northeast; Atlantic – eastern central
|Range Map:||Click here to open the map viewer and explore range.|
|Population:||Loggerheads are a single species globally comprising 10 biologically described regional management units (RMUs: Wallace et al. 2010), which describe biologically and geographically explicit population segments by integrating information from nesting sites, mitochondrial and nuclear DNA studies, movements and habitat use by all life stages. RMUs are functionally equivalent to IUCN subpopulations, thus providing the appropriate demographic unit for Red List assessments. There are 10 Loggerhead RMUs (hereafter subpopulations): North West Atlantic Ocean, North East Atlantic Ocean, South West Atlantic Ocean, Mediterranean Sea, North East Indian Ocean, North West Indian Ocean, South East Indian Ocean, South West Indian Ocean, North Pacific Ocean, and South Pacific Ocean (Figure 2 in the Supplementary Material). Multiple genetic stocks have been defined according to geographically disparate nesting areas around the world and are included within RMU delineations (Wallace et al. 2010) (shapefiles can be viewed and downloaded at: http://seamap.env.duke.edu/swot).|
Nesting occurs within the Northwest Atlantic along the coasts of North America, Central America, northern South America, the Antilles, and the Bahamas, but is concentrated in the southeastern U.S. and on the Yucatan Peninsula in Mexico (Ehrhart et al. 2003, Dow et al. 2007, NMFS and USFWS 2008). NMFS and USFWS (2008) identified five recovery units within the Northwest Atlantic subpopulation: the Northern (southern Virginia to Florida/Georgia border); Peninsular Florida (Florida/Georgia border south and then north through Pinellas County on the west coast, excluding the islands west of Key West, Florida); Dry Tortugas (islands west of Key West, Florida); Northern Gulf of Mexico (Franklin County, Florida, west through Texas) and the Greater Caribbean (Mexico through French Guiana, the Bahamas, and Lesser and Greater Antilles). The Peninsular Florida Recovery Unit and the Northern Recovery Unit represent ~87% and ~10%, respectively, of all nesting effort in the Northwest Atlantic Loggerhead subpopulation (Ehrhart et al. 2003, NMFS and USFWS 2008).
|Current Population Trend:||Increasing|
|Habitat and Ecology:||The Loggerhead Turtle nests on insular and mainland sandy beaches throughout temperate and subtropical regions. Like most marine turtles, Loggerhead Turtles are highly migratory and use a wide range of broadly separated localities and habitats during their lifetimes (Bolten and Witherington 2003). Upon leaving the nesting beach, hatchlings begin an oceanic phase, perhaps floating passively in major current systems (gyres) that serve as open-ocean developmental grounds (Bolten and Witherington 2003). After 4-19 years in the oceanic zone, Loggerheads recruit to neritic developmental areas rich in benthic prey or epipelagic prey where they forage and grow until maturity at 10-39 years (Avens and Snover 2013). Upon attaining sexual maturity, Loggerhead Turtles undertake breeding migrations between foraging grounds and nesting areas at remigration intervals of one to several years with a mean of 2.5-3 years for females (Schroeder et al. 2003) while males would have a shorter remigration interval (e.g., Hays et al. 2010, Wibbels et al. 1990). Migrations are carried out by both males and females and may traverse oceanic zones spanning hundreds to thousands of kilometres (Plotkin 2003). During non-breeding periods adults reside at coastal neritic feeding areas that sometimes coincide with juvenile developmental habitats (Bolten and Witherington 2003), although recent studies indicate that a proportion of adults forage pelagically (Hatase et al. 2002, 2013; Hawkes et al. 2006).|
Like other species of marine turtles, Loggerheads contribute to marine and coastal food webs and transport nutrients within the oceans (Bouchard and Bjorndal 2000). The nest remains of unhatched eggs or shells offer oceanic nutrient inputs that stabilize terrestrial dune vegetation. Loggerheads inhabit estuaries, nearshore and offshore reef environments, and feed on a wide variety of benthic invertebrates, especially crustaceans and molluscs.
The IUCN Red List Criteria (IUCN 2014) define generation length to be the average age of parents in a population (i.e., older than the age at maturity and younger than the oldest mature individual). Although different subpopulations may have different generation lengths, we adopted the same value for all the subpopulations because of limited data, taking care to avoid underestimation as recommended by IUCN (2014).
Loggerheads attain maturity at 10-39 years (Avens and Snover 2013), and we considered here 30 years to be equal or greater than the average age at maturity. Data on reproductive longevity in Loggerheads are limited, but are becoming available with increasing numbers of intensively monitored, long-term projects on protected beaches. Tagging studies have documented reproductive histories up to 28 years in the North Western Atlantic Ocean (Mote Marine Laboratory, unpubl. data), up to 18 years in the South Western Indian Ocean (Nel et al. 2013), up to 32 years in the South Western Atlantic Ocean (Projeto Tamar unpubl. data), and up to 37 years in the South Western Pacific Ocean, where females nesting for 20-25 years are common (C. Limpus, pers. comm). We considered 15 years to be equal or greater than the average reproductive longevity. Therefore, we considered here 45 years to be equal or greater than the average generation length, therefore avoiding underestimation as recommended by IUCN (IUCN Standards and Petitions Subcommittee 2014).
|Generation Length (years):||45|
Loggerhead turtles, like other marine turtle species, are particularly susceptible to population declines because of their vulnerability to anthropogenic impacts during all life-stages from eggs to adults and in both marine and terrestrial environments.
Threats to Loggerheads vary in time and space, and in relative impact to populations. Threats affecting marine turtles, including Loggerheads, were described by Wallace et al. (2011) as:
Fisheries bycatch has been identified as the major threat for the recovery of the Northwest Atlantic Loggerhead subpopulation (Witherington et al. 2009, Bolten et al. 2010), followed by pollution and possible food resource decline (Witherington et al. 2009). See also NMFS and USFWS (2008) for detailed description of threats.
Loggerhead Turtles are afforded legislative protection under a number of treaties and laws (Wold 2002). Annex II of the SPAW Protocol to the Cartagena Convention (a protocol concerning specially protected areas and wildlife); Appendix I of CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora); and Appendices I and II of the Convention on Migratory Species (CMS). A partial list of the International Instruments that benefit Loggerhead Turtles includes the Inter-American Convention for the Protection and Conservation of Sea Turtles, the Memorandum of Understanding on the Conservation and Management of Marine Turtles and their Habitats of the Indian Ocean and South-East Asia (IOSEA), the Memorandum of Understanding on ASEAN Sea Turtle Conservation and Protection, the Memorandum of Agreement on the Turtle Islands Heritage Protected Area (TIHPA), and the Memorandum of Understanding Concerning Conservation Measures for Marine Turtles of the Atlantic Coast of Africa.
As a result of these designations and agreements, many of the intentional impacts directed at marine turtles have been lessened. Harvest of eggs and adults has been reduced or eliminated at several nesting areas through nesting beach conservation efforts. An increasing number of community-based initiatives are in place to reduce the take of turtles in foraging areas. In regard to incidental take, the implementation of Turtle Excluder Devices has proved to be beneficial in some areas, primarily in the United States and South and Central America (National Research Council 1990). Guidelines are available to reduce marine turtle mortality in fishing operations in coastal and high seas fisheries (FAO 2009). However, despite these advances, human impacts continue throughout the world. The lack of effective monitoring and law enforcement in pelagic and near-shore fisheries operations still allow substantial direct and indirect mortality, and the uncontrolled development of coastal and marine habitats threatens to destroy the supporting ecosystems of long-lived Loggerhead Turtles.
Most countries where Northwest Atlantic Loggerheads are found, including those with major nesting and foraging habitats, are signatories to international conventions for the conservation of species, including marine turtles, and have national laws for the protection of marine turtles. Moreover, many countries have marine turtle conservation projects run by governmental bodies or NGOs. Special attention has been given to the protection of major nesting sites, particularly in the southeastern United States leading to the recent definition and protection of Loggerhead Critical Habitat in the Northwest Atlantic and Gulf of Mexico (NMFS 2014). The NOAA-designated marine critical habitat included some nearshore reproductive areas directly off of nesting beaches, breeding and wintering habitats, migratory corridors and Sargassum habitat. The USFWS-designated terrestrial critical habitat areas included 88 nesting beaches in the southeastern U.S. which account for about 84% of the documented numbers of nests in the southeastern U.S. For a detailed description of the conservation efforts targeting the Northwest Atlantic Loggerhead population see NMFS and USFWS (2008).
|Citation:||Ceriani, S.A. & Meylan, A.B. 2015. Caretta caretta (North West Atlantic subpopulation). The IUCN Red List of Threatened Species 2015: e.T84131194A84131608.Downloaded on 20 February 2017.|
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