Endangered and Threatened Species; Proposed Listing of Nine Distinct Population Segments of Loggerhead Sea Turtles as Endangered or Threatened

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DEPARTMENT OF THE INTERIOR Fish and Wildlife Service 50 CFR Part 17 DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration 50 CFR Parts 223 and 224 [Docket No. 100104003-0004-01] RIN 0648-AY49 Endangered and Threatened Species; Proposed Listing of Nine Distinct Population Segments of Loggerhead Sea Turtles as Endangered or Threatened AGENCY:

National Marine Fisheries Service (NMFS), National Oceanic and Atmospheric Administration (NOAA), Commerce; United States Fish and Wildlife Service (USFWS), Interior.

ACTION:

Proposed rules; 12-month petition findings; request for comments.

SUMMARY:

We (NMFS and USFWS; also collectively referred to as the Services) have determined that the loggerhead sea turtle ( Caretta caretta ) is composed of nine distinct population segments (DPSs) that qualify as “species” for listing as endangered or threatened under the Endangered Species Act (ESA), and we propose to list two as threatened and seven as endangered. This also constitutes the 12-month findings on a petition to reclassify loggerhead turtles in the North Pacific Ocean as a DPS with endangered status and designate critical habitat, and a petition to reclassify loggerhead turtles in the Northwest Atlantic as a DPS with endangered status and designate critical habitat. We will propose to designate critical habitat, if found to be prudent and determinable, for the two loggerhead sea turtle DPSs occurring within the United States in a subsequent Federal Register notice.

DATES:

Comments on this proposal must be received by June 14, 2010. Public hearing requests must be received by June 1, 2010.

ADDRESSES:

You may submit comments, identified by the RIN 0648-AY49, by any of the following methods:

• Electronic Submissions: Submit all electronic public comments via the Federal eRulemaking Portal

the United States in a subsequent Federal Register notice.

DATES:

Comments on this proposal must be received by June 14, 2010. Public hearing requests must be received by June 1, 2010.

ADDRESSES:

You may submit comments, identified by the RIN 0648-AY49, by any of the following methods:

• Electronic Submissions: Submit all electronic public comments via the Federal eRulemaking Portal.

• Mail: NMFS National Sea Turtle Coordinator, Attn: Loggerhead Proposed Listing Rule, Office of Protected Resources, National Marine Fisheries Service, 1315 East-West Highway, Room 13657, Silver Spring, MD 20910 or USFWS National Sea Turtle Coordinator, U.S. Fish and Wildlife Service, 7915 Baymeadows Way, Suite 200, Jacksonville, FL 32256.

• Fax: To the attention of NMFS National Sea Turtle Coordinator at 301-713-0376 or USFWS National Sea Turtle Coordinator at 904-731-3045.

Instructions: All comments received are a part of the public record and will generally be posted to http://www.regulations.gov without change. All Personal Identifying Information (for example, name, address, etc. ) voluntarily submitted by the commenter may be publicly accessible. Do not submit Confidential Business Information or otherwise sensitive or protected information.

NMFS and USFWS will accept anonymous comments (enter N/A in the required fields, if you wish to remain anonymous). Attachments to electronic comments will be accepted in Microsoft Word, Excel, WordPerfect, or Adobe PDF file formats only. The proposed rule is available electronically at http://www.nmfs.noaa.gov/pr.

FOR FURTHER INFORMATION CONTACT:

v/pr.

FOR FURTHER INFORMATION CONTACT:

Barbara Schroeder, NMFS (ph. 301-713-1401, fax 301-713-0376, e-mail barbara.schroeder@noaa.gov ), Sandy MacPherson, USFWS (ph. 904-731-3336, e-mail sandy_macpherson@fws.gov ), Marta Nammack, NMFS (ph. 301-713-1401, fax 301-713-0376, e-mail marta_nammack@noaa.gov ), or Emily Bizwell, USFWS (ph. 404-679-7149, fax 404-679-7081, e-mail emily_bizwell@fws.gov ). Persons who use a Telecommunications device for the deaf (TDD) may call the Federal Information Relay Service (FIRS) at 1-800-877-8339, 24 hours a day, 7 days a week.

SUPPLEMENTARY INFORMATION:

Public Comments Solicited

We solicit public comment on this proposed listing determination. We intend that any final action resulting from this proposal will be as accurate and as effective as possible and informed by the best available scientific and commercial information. Therefore, we request comments or information from the public, other concerned governmental agencies, the scientific community, industry, or any other interested party concerning this proposed rule. We are seeking information and comments on whether the nine proposed loggerhead sea turtle DPSs qualify as DPSs and, if so, whether they should be classified as threatened or endangered as described in the “Listing Determinations Under the ESA” section provided below. Specifically, we are soliciting information in the following areas relative to loggerhead turtles within the nine proposed DPSs: (1) Historical and current population status and trends, (2) historical and current distribution, (3) migratory movements and behavior, (4) genetic population structure, (5) current or planned activities that may adversely impact loggerhead turtles, and (6) ongoing efforts to protect loggerhead turtles. We are also soliciting information and comment on the status and effectiveness of conservation efforts and the approach that should be used to weigh the risk of extinction of each DPS

ribution, (3) migratory movements and behavior, (4) genetic population structure, (5) current or planned activities that may adversely impact loggerhead turtles, and (6) ongoing efforts to protect loggerhead turtles. We are also soliciting information and comment on the status and effectiveness of conservation efforts and the approach that should be used to weigh the risk of extinction of each DPS. Comments and new information will be considered in making final determinations whether listing of each DPS is warranted and if so whether it is threatened or endangered. We request that all data, information, and comments be accompanied by supporting documentation such as maps, bibliographic references, or reprints of pertinent publications.

Background

We issued a final rule listing the loggerhead sea turtle as threatened throughout its worldwide range on July 28, 1978 (43 FR 32800). On July 12, 2007, we received a petition to list the “North Pacific populations of loggerhead sea turtle” as an endangered species under the ESA. NMFS published a notice in the Federal Register on November 16, 2007 (72 FR 64585), concluding that the petitioners (Center for Biological Diversity and Turtle Island Restoration Network) presented substantial scientific information indicating that the petitioned action may be warranted. Also, on November 15, 2007, we received a petition to list the “Western North Atlantic populations of loggerhead sea turtle” as an endangered species under the ESA. NMFS published a notice in the Federal Register on March 5, 2008 (73 FR 11849), concluding that the petitioners (Center for Biological Diversity and Oceana) presented substantial scientific information indicating that the petitioned action may be warranted.

luding that the petitioners (Center for Biological Diversity and Oceana) presented substantial scientific information indicating that the petitioned action may be warranted.

On March 12, 2009, the petitioners (Center for Biological Diversity, Turtle Island Restoration Network, and Oceana) sent a 60-day notice of intent to sue to the Services for failure to make 12-month findings on the petitions. The statutory deadlines for the 12-month findings were July 16, 2008, for the North Pacific petition and November 16, 2008, for the Northwest Atlantic petition. On May 28, 2009, the petitioners filed a Complaint for Federal Register a 12-month finding on the two petitions on or before February 19, 2010. On February 16, 2010, the United States District Court for the Northern District of California modified the February 19, 2010 deadline to March 8, 2010.

In early 2008, NMFS assembled a Loggerhead Biological Review Team (BRT) to complete a status review of the loggerhead sea turtle. The BRT was composed of biologists from NMFS, USFWS, the Florida Fish and Wildlife Conservation Commission, and the North Carolina Wildlife Resources Commission. The BRT was charged with reviewing and evaluating all relevant scientific information relating to loggerhead population structure globally to determine whether DPSs exist and, if so, to assess the status of each DPS. The findings of the BRT, which are detailed in the “Loggerhead Sea Turtle ( Caretta caretta ) 2009 Status Review under the U.S. Endangered Species Act” (Conant et al., 2009; hereinafter referred to as the Status Review), addressed DPS delineations, extinction risks to the species, and threats to the species. The Status Review underwent independent peer review by nine scientists with expertise in loggerhead sea turtle biology, genetics, and modeling. The Status Review is available electronically at http://www.nmfs.noaa.gov/pr/species/statusreviews.htm

ant et al., 2009; hereinafter referred to as the Status Review), addressed DPS delineations, extinction risks to the species, and threats to the species. The Status Review underwent independent peer review by nine scientists with expertise in loggerhead sea turtle biology, genetics, and modeling. The Status Review is available electronically at http://www.nmfs.noaa.gov/pr/species/statusreviews.htm.

This Federal Register document announces 12-month findings on the petitions to list the North Pacific populations and the Northwest Atlantic populations of the loggerhead sea turtle as DPSs with endangered status and includes a proposed rule to designate nine loggerhead DPSs worldwide.

Policies for Delineating Species Under the ESA

Section 3 of the ESA defines “species” as including “any subspecies of fish or wildlife or plants, and any distinct population segment of any species of vertebrate fish or wildlife which interbreeds when mature.” The term “distinct population segment” is not recognized in the scientific literature. Therefore, the Services adopted a joint policy for recognizing DPSs under the ESA (DPS Policy; 61 FR 4722) on February 7, 1996. Congress has instructed the Secretary of the Interior or of Commerce to exercise this authority with regard to DPSs “* * * sparingly and only when the biological evidence indicates such action is warranted.” The DPS Policy requires the consideration of two elements when evaluating whether a vertebrate population segment qualifies as a DPS under the ESA: (1) The discreteness of the population segment in relation to the remainder of the species or subspecies to which it belongs; and (2) the significance of the population segment to the species or subspecies to which it belongs

such action is warranted.” The DPS Policy requires the consideration of two elements when evaluating whether a vertebrate population segment qualifies as a DPS under the ESA: (1) The discreteness of the population segment in relation to the remainder of the species or subspecies to which it belongs; and (2) the significance of the population segment to the species or subspecies to which it belongs.

A population segment of a vertebrate species may be considered discrete if it satisfies either one of the following conditions: (1) It is markedly separated from other populations of the same taxon (an organism or group of organisms) as a consequence of physical, ecological, or behavioral factors. Quantitative measures of genetic or morphological discontinuity may provide evidence of this separation; or (2) it is delimited by international governmental boundaries within which differences in control of exploitation, management of habitat, conservation status, or regulatory mechanisms exist that are significant in light of section 4(a)(1)(D) of the ESA ( i.e., inadequate regulatory mechanisms).

If a population segment is found to be discrete under one or both of the above conditions, its biological and ecological significance to the taxon to which it belongs is evaluated. This consideration may include, but is not limited to: (1) Persistence of the discrete population segment in an ecological setting unusual or unique for the taxon; (2) evidence that loss of the discrete population segment would result in a significant gap in the range of a taxon; (3) evidence that the discrete population segment represents the only surviving natural occurrence of a taxon that may be more abundant elsewhere as an introduced population outside its historic range; or (4) evidence that the discrete population segment differs markedly from other population segments of the species in its genetic characteristics

esult in a significant gap in the range of a taxon; (3) evidence that the discrete population segment represents the only surviving natural occurrence of a taxon that may be more abundant elsewhere as an introduced population outside its historic range; or (4) evidence that the discrete population segment differs markedly from other population segments of the species in its genetic characteristics.

Listing Determinations Under the ESA

The ESA defines an endangered species as one that is in danger of extinction throughout all or a significant portion of its range, and a threatened species as one that is likely to become endangered in the foreseeable future throughout all or a significant portion of its range (sections 3(6) and 3(20), respectively). The statute requires us to determine whether any species is endangered or threatened because of any of the following five factors: (1) The present or threatened destruction, modification, or curtailment of its habitat or range; (2) overutilization for commercial, recreational, scientific, or educational purposes; (3) disease or predation; (4) the inadequacy of existing regulatory mechanisms; or (5) other natural or manmade factors affecting its continued existence (section 4(a)(1)(A-E)). We are to make this determination based solely on the best available scientific and commercial data available after conducting a review of the status of the species and taking into account any efforts being made by States or foreign governments to protect the species.

Biology and Life History of Loggerhead Turtles

A thorough account of loggerhead biology and life history may be found in the Status Review, which is incorporated here by reference. The following is a succinct summary of that information.

of loggerhead biology and life history may be found in the Status Review, which is incorporated here by reference. The following is a succinct summary of that information.

The loggerhead occurs throughout the temperate and tropical regions of the Atlantic, Pacific, and Indian Oceans (Dodd, 1988). However, the majority of loggerhead nesting is at the western rims of the Atlantic and Indian Oceans. The most recent reviews show that only two loggerhead nesting aggregations have greater than 10,000 females nesting per year: Peninsular Florida, United States, and Masirah Island, Oman (Baldwin et al., 2003; Ehrhart et al., 2003; Kamezaki et al., 2003; Limpus and Limpus, 2003; Margaritoulis et al., 2003). Nesting aggregations with 1,000 to 9,999 females nesting annually are Georgia through North Carolina (United States), Quintana Roo and Yucatan (Mexico), Brazil, Cape Verde Islands (Cape Verde), Western Australia (Australia), and Japan. Smaller nesting aggregations with 100 to 999 nesting females annually occur in the Northern Gulf of Mexico (United States), Dry Tortugas (United States), Cay Sal Bank (The Bahamas), Tongaland (South Africa), Mozambique, Arabian Sea Coast (Oman), Halaniyat Islands (Oman), Cyprus, Peloponnesus (Greece), Zakynthos (Greece), Crete (Greece), Turkey, and Queensland (Australia). In contrast to determining population size on nesting beaches, determining population size in the marine environment has been very localized. A summary of information on distribution and habitat by ocean basin follows.

Pacific Ocean

Loggerheads can be found throughout tropical to temperate waters in the Pacific; however, their breeding grounds include a restricted number of sites in the North Pacific and South Pacific. Within the North Pacific, loggerhead nesting has been documented only in Japan (Kamezaki et al., 2003), although et al., 2007). In the South Pacific, nesting beaches are restricted to eastern Australia and New Caledonia and, to a much lesser extent, Vanuatu and Tokelau (Limpus and Limpus, 2003)

ver, their breeding grounds include a restricted number of sites in the North Pacific and South Pacific. Within the North Pacific, loggerhead nesting has been documented only in Japan (Kamezaki et al., 2003), although et al., 2007). In the South Pacific, nesting beaches are restricted to eastern Australia and New Caledonia and, to a much lesser extent, Vanuatu and Tokelau (Limpus and Limpus, 2003).

Based on tag-recapture studies, the East China Sea has been identified as the major habitat for post-nesting adult females (Iwamoto et al., 1985; Kamezaki et al., 1997; Balazs, 2006), while satellite tracking of juvenile loggerheads indicates the Kuroshio Extension Bifurcation Region to be an important pelagic foraging area for juvenile loggerheads (Polovina et al., 2006). Other important juvenile turtle foraging areas have been identified off the coast of Baja California Sur, Mexico (Pitman, 1990; Peckham and Nichols, 2006).

Nesting females tagged on the coast of eastern Australia have been recorded foraging in New Caledonia; Queensland, New South Wales, and Northern Territory, Australia; Solomon Islands; Papua New Guinea; and Indonesia (Limpus and Limpus, 2003). Foraging Pacific loggerheads originating from nesting beaches in Australia are known to migrate to Chile and Peru (Alfaro-Shigueto et al., 2004, 2008a; Donoso and Dutton, 2006; Boyle et al., 2009).

Indian Ocean

In the North Indian Ocean, Oman hosts the vast majority of loggerhead nesting. The majority of the nesting in Oman occurs on Masirah Island, on the Al Halaniyat Islands, and on mainland beaches south of Masirah Island all the way to the Oman-Yemen border (IUCN—The World Conservation Union, 1989a, 1989b; Salm, 1991; Salm and Salm, 1991). In addition, nesting probably occurs on the mainland of Yemen on the Arabian Sea coast, and nesting has been confirmed on Socotra, an island off the coast of Yemen (Pilcher and Saad, 2000)

sirah Island, on the Al Halaniyat Islands, and on mainland beaches south of Masirah Island all the way to the Oman-Yemen border (IUCN—The World Conservation Union, 1989a, 1989b; Salm, 1991; Salm and Salm, 1991). In addition, nesting probably occurs on the mainland of Yemen on the Arabian Sea coast, and nesting has been confirmed on Socotra, an island off the coast of Yemen (Pilcher and Saad, 2000). Limited information exists on the foraging habitats of North Indian Ocean loggerheads; however, foraging individuals have been reported off the southern coastline of Oman (Salm et al., 1993). Satellite telemetry studies of post-nesting migrations of loggerheads nesting on Masirah Island, Oman, have revealed extensive use of the waters off the Arabian Peninsula, with the majority of telemetered turtles traveling southwest, following the shoreline of southern Oman and Yemen, and circling well offshore in nearby oceanic waters (Environment Society of Oman and Ministry of Environment and Climate Change, Oman, unpublished data). A minority traveled north as far as the western Persian (Arabian) Gulf or followed the shoreline of southern Oman and Yemen as far west as the Gulf of Aden and the Bab-el-Mandab.

The only verified nesting beaches for loggerheads on the Indian subcontinent are found in Sri Lanka. A small number of nesting females use the beaches of Sri Lanka every year (Deraniyagala, 1939; Kar and Bhaskar, 1982; Dodd, 1988); however, there are no records indicating that Sri Lanka has ever been a major nesting area for loggerheads (Kapurusinghe, 2006). No confirmed nesting occurs on the mainland of India (Tripathy, 2005; Kapurusinghe, 2006). The Gulf of Mannar provides foraging habitat for juvenile and post-nesting adult turtles (Tripathy, 2005; Kapurusinghe, 2006).

In the East Indian Ocean, western Australia hosts all known loggerhead nesting (Dodd, 1988)

that Sri Lanka has ever been a major nesting area for loggerheads (Kapurusinghe, 2006). No confirmed nesting occurs on the mainland of India (Tripathy, 2005; Kapurusinghe, 2006). The Gulf of Mannar provides foraging habitat for juvenile and post-nesting adult turtles (Tripathy, 2005; Kapurusinghe, 2006).

In the East Indian Ocean, western Australia hosts all known loggerhead nesting (Dodd, 1988). Nesting distributions in western Australia span from the Shark Bay World Heritage Area northward through the Ningaloo Marine Park coast to the North West Cape and to the nearby Muiron Islands (Baldwin et al., 2003). Nesting individuals from Dirk Hartog Island have been recorded foraging within Shark Bay and Exmouth Gulf, while other adults range much farther (Baldwin et al., 2003).

In the Southwest Indian Ocean, loggerhead nesting occurs on the southeastern coast of Africa, from the Paradise Islands in Mozambique southward to St. Lucia in South Africa, and on the south and southwestern coasts of Madagascar (Baldwin et al., 2003). Foraging habitats are only known for post-nesting females from Tongaland, South Africa; tagging data show these loggerheads migrating eastward to Madagascar, northward to Mozambique, Tanzania, and Kenya, and southward to Cape Agulhas at the southernmost point of Africa (Baldwin et al., 2003; Luschi et al., 2006).

Atlantic Ocean

In the Northwest Atlantic, the majority of loggerhead nesting is concentrated along the coasts of the United States from southern Virginia through Alabama. Additional nesting beaches are found along the northern and western Gulf of Mexico, eastern Yucatan Peninsula, at Cay Sal Bank in the eastern Bahamas (Addison and Morford, 1996; Addison, 1997), on the southwestern coast of Cuba (F. Moncada-Gavilan, personal communication, cited in Ehrhart et al., 2003), and along the coasts of Central America, Colombia, Venezuela, and the eastern Caribbean Islands. In the Southwest Atlantic, loggerheads nest in significant numbers only in Brazil

eastern Yucatan Peninsula, at Cay Sal Bank in the eastern Bahamas (Addison and Morford, 1996; Addison, 1997), on the southwestern coast of Cuba (F. Moncada-Gavilan, personal communication, cited in Ehrhart et al., 2003), and along the coasts of Central America, Colombia, Venezuela, and the eastern Caribbean Islands. In the Southwest Atlantic, loggerheads nest in significant numbers only in Brazil. In the eastern Atlantic, the largest nesting population of loggerheads is in the Cape Verde Islands (L.F. Lopez-Jurado, personal communication, cited in Ehrhart et al., 2003), and some nesting occurs along the West African coast (Fretey, 2001).

As post-hatchlings, Northwest Atlantic loggerheads use the North Atlantic Gyre and enter Northeast Atlantic waters (Carr, 1987). They are also found in the Mediterranean Sea (Carreras et al., 2006; Eckert et al., 2008). In these areas, they overlap with animals originating from the Northeast Atlantic and the Mediterranean Sea (Laurent et al., 1993, 1998; Bolten et al., 1998; LaCasella et al., 2005; Carreras et al., 2006; Monzon-Arguello et al., 2006; Revelles et al., 2007; Eckert et al., 2008). The oceanic juvenile stage in the North Atlantic has been primarily studied in the waters around the Azores and Madeira (Bolten, 2003). In Azorean waters, satellite telemetry data and flipper tag returns suggest a long period of residency (Bolten, 2003), whereas turtles appear to be moving through Madeiran waters (Dellinger and Freitas, 2000). Preliminary genetic analyses indicate that juvenile loggerheads found in Moroccan waters are of western Atlantic origin (M. Tiwari, NMFS, and A. Bolten, University of Florida, unpublished data). Other concentrations of oceanic juvenile turtles exist in the Atlantic ( e.g., in the region of the Grand Banks off Newfoundland)

appear to be moving through Madeiran waters (Dellinger and Freitas, 2000). Preliminary genetic analyses indicate that juvenile loggerheads found in Moroccan waters are of western Atlantic origin (M. Tiwari, NMFS, and A. Bolten, University of Florida, unpublished data). Other concentrations of oceanic juvenile turtles exist in the Atlantic ( e.g., in the region of the Grand Banks off Newfoundland). Genetic information indicates the Grand Banks are foraging grounds for a mixture of loggerheads from all the North Atlantic rookeries (LaCasella et al., 2005; Bowen et al., 2005), and a large size range is represented (Watson et al., 2004, 2005).

After departing the oceanic zone, neritic juvenile loggerheads in the Northwest Atlantic inhabit continental shelf waters from Cape Cod Bay, Massachusetts, south through Florida, The Bahamas, Cuba, and the Gulf of Mexico (neritic refers to the inshore marine environment from the surface to the sea floor where water depths do not exceed 200 meters).

Habitat preferences of Northwest Atlantic non-nesting adult loggerheads in the neritic zone differ from the juvenile stage in that relatively enclosed, shallow water estuarine habitats with limited ocean access are less frequently used. Areas such as Pamlico Sound and the Indian River Lagoon in the United States, regularly used by juvenile loggerheads, are only rarely frequented by adults. In comparison, estuarine areas with more open ocean access, such as Chesapeake Bay in the U.S. mid-Atlantic, are also regularly used by juvenile loggerheads, as well as by adults primarily during et al. (in press) reported the recapture in Cuban waters of five adult female loggerheads originally flipper tagged in Quintana Roo, Mexico, indicating that Cuban shelf waters likely also provide foraging habitat for adult females that nest in Mexico

ocean access, such as Chesapeake Bay in the U.S. mid-Atlantic, are also regularly used by juvenile loggerheads, as well as by adults primarily during et al. (in press) reported the recapture in Cuban waters of five adult female loggerheads originally flipper tagged in Quintana Roo, Mexico, indicating that Cuban shelf waters likely also provide foraging habitat for adult females that nest in Mexico.

In the Northeast Atlantic, satellite telemetry studies of post-nesting females from Cape Verde identified two distinct dispersal patterns; larger individuals migrated to benthic foraging areas off the northwest Africa coast and smaller individuals foraged primarily oceanically off the northwest Africa coast (Hawkes et al., 2006). Monzon-Arguello et al. (2009) conducted a mixed stock analysis of juvenile loggerheads sampled from foraging areas in the Canary Islands, Madeira, Azores, and Andalusia and concluded that while juvenile loggerheads from the Cape Verde population were distributed among these four sites, a large proportion of Cape Verde juvenile turtles appear to inhabit as yet unidentified foraging areas.

In the South Atlantic, relatively little is known about the at-sea behavior of loggerheads originating from nesting beaches in Brazil. Recaptures of tagged juvenile turtles and nesting females have shown movement of animals up and down the coast of South America (Almeida et al., 2000; Marcovaldi et al., 2000; Laporta and Lopez, 2003; Almeida et al., 2007). Juvenile loggerheads, presumably of Brazilian origin, have also been captured on the high seas of the South Atlantic (Kotas et al., 2004; Pinedo and Polacheck, 2004) and off the coast of Atlantic Africa (Bal et al., 2007; Petersen, 2005; Petersen et al., 2007) suggesting that loggerheads of the South Atlantic may undertake transoceanic developmental migrations (Bolten et al., 1998; Peckham et al., 2007).

Mediterranean Sea

Loggerhead turtles are widely distributed in the Mediterranean Sea

f the South Atlantic (Kotas et al., 2004; Pinedo and Polacheck, 2004) and off the coast of Atlantic Africa (Bal et al., 2007; Petersen, 2005; Petersen et al., 2007) suggesting that loggerheads of the South Atlantic may undertake transoceanic developmental migrations (Bolten et al., 1998; Peckham et al., 2007).

Mediterranean Sea

Loggerhead turtles are widely distributed in the Mediterranean Sea. However, nesting is almost entirely confined to the eastern Mediterranean basin, with the main nesting concentrations in Cyprus, Greece, and Turkey (Margaritoulis et al., 2003). Preliminary surveys in Libya suggested nesting activity comparable to Greece and Turkey, although a better quantification is needed (Laurent et al., 1999). Minimal to moderate nesting also occurs in other countries throughout the Mediterranean including Egypt, Israel, Italy (southern coasts and islands), Lebanon, Syria, and Tunisia (Margaritoulis et al., 2003). Recently, isolated nesting events have been recorded in the western Mediterranean basin, namely in Spain, Corsica (France), and in the Tyrrhenian Sea (Italy) (Tomas et al., 2002; Delaugerre and Cesarini, 2004; Bentivegna et al., 2005).

Important neritic habitats have been suggested for the large continental shelves of: (1) Tunisia-Libya, (2) northern Adriatic Sea, (3) Egypt, and (4) Spain (Margaritoulis, 1988; Argano et al., 1992; Laurent and Lescure, 1994; Lazar et al., 2000; Gomez de Segura et al., 2006; Broderick et al., 2007; Casale et al., 2007b; Nada and Casale, 2008). At least the first three constitute shallow benthic habitats for adults (including post-nesting females). Some other neritic foraging areas include Amvrakikos Bay in western Greece, Lakonikos Bay in southern Greece, and southern Turkey. Oceanic foraging areas for small juvenile loggerheads have been identified in the south Adriatic Sea (Casale et al., 2005b), Ionian Sea (Deflorio et al., 2005), Sicily Strait (Casale et al., 2007b), and western Mediterranean (Spain) ( e.g., Camiñas et al., 2006)

ales). Some other neritic foraging areas include Amvrakikos Bay in western Greece, Lakonikos Bay in southern Greece, and southern Turkey. Oceanic foraging areas for small juvenile loggerheads have been identified in the south Adriatic Sea (Casale et al., 2005b), Ionian Sea (Deflorio et al., 2005), Sicily Strait (Casale et al., 2007b), and western Mediterranean (Spain) ( e.g., Camiñas et al., 2006). In addition, tagged juvenile loggerheads have been recorded crossing the Mediterranean from the eastern to the western basin and vice versa, as well as in the Eastern Atlantic (Argano et al., 1992; Casale et al., 2007b).

Reproductive migrations have been confirmed by flipper tagging and satellite telemetry. Female loggerheads, after nesting in Greece, migrate primarily to the Gulf of Gabès and the northern Adriatic (Margaritoulis, 1988; Margaritoulis et al., 2003; Lazar et al., 2004; Zbinden et al., 2008). Loggerheads nesting in Cyprus migrate to Egypt and Libya, exhibiting fidelity in following the same migration route during subsequent nesting seasons (Broderick et al., 2007). In addition, directed movements of juvenile loggerheads have been confirmed through flipper tagging (Argano et al., 1992; Casale et al., 2007b) and satellite tracking (Rees and Margaritoulis, 2009).

Overview of Information Used To Identify DPSs

In the Status Review, the BRT considered a vast array of information to assess whether there are any loggerhead population segments that satisfy the DPS criteria of both discreteness and significance. First, the BRT examined whether there were any loggerhead population segments that were discrete. Data relevant to the discreteness question included physical, ecological, behavioral, and genetic data. Given the physical separation of ocean basins by continents, the BRT evaluated these data by ocean basin (Pacific Ocean, Indian Ocean, and Atlantic Ocean). This was not to preclude any larger or smaller DPS delineation, but to aid in data organization and assessment

segments that were discrete. Data relevant to the discreteness question included physical, ecological, behavioral, and genetic data. Given the physical separation of ocean basins by continents, the BRT evaluated these data by ocean basin (Pacific Ocean, Indian Ocean, and Atlantic Ocean). This was not to preclude any larger or smaller DPS delineation, but to aid in data organization and assessment. The BRT then evaluated genetic information by ocean basin. The genetic data consisted of results from studies using maternally inherited mitochondrial DNA (mtDNA) and biparentally inherited nuclear DNA microsatellite markers. Next, tagging data (both flipper and PIT tags) and telemetry data were reviewed. Additional information, such as potential differences in morphology, was also evaluated. Finally, the BRT considered whether the available information on loggerhead population segments was bounded by any oceanographic features ( e.g., current systems) or geographic features ( e.g., land masses).

In accordance with the DPS policy, the BRT also reviewed whether the population segments identified in the discreteness analysis were significant. If a population segment is considered discrete, its biological and ecological significance must then be considered. NMFS and USFWS must consider available scientific evidence of the discrete segment's importance to the taxon to which it belongs. Data relevant to the significance question include morphological, ecological, behavioral, and genetic data, as described above

analysis were significant. If a population segment is considered discrete, its biological and ecological significance must then be considered. NMFS and USFWS must consider available scientific evidence of the discrete segment's importance to the taxon to which it belongs. Data relevant to the significance question include morphological, ecological, behavioral, and genetic data, as described above. The BRT considered the following factors, listed in the DPS policy, in determining whether the discrete population segments were significant: (a) Persistence of the discrete segment in an ecological setting unusual or unique for the taxon; (b) evidence that loss of the discrete segment would result in a significant gap in the range of the taxon; (c) evidence that the discrete segment represents the only surviving natural occurrence of a taxon that may be more abundant elsewhere as an introduced

A discrete population segment needs to satisfy only one of these criteria to be considered significant. The DPS policy also allows for consideration of other factors if they are appropriate to the biology or ecology of the species. As described below, the BRT evaluated the available information and considered items (a), (b) and (d), as noted above, to be most applicable to loggerheads.

Discreteness Determination

As described in the Status Review, the loggerhead sea turtle is present in all tropical and temperate ocean basins, and has a life history that involves nesting on coastal beaches and foraging in neritic and oceanic habitats, as well as long-distance migrations between and within these areas. As with other globally distributed marine species, today's global loggerhead population has been shaped by a sequence of isolation events created by tectonic and oceanographic shifts over geologic time scales, the result of which is population substructuring in many areas (Bowen et al., 1994; Bowen, 2003)

tic and oceanic habitats, as well as long-distance migrations between and within these areas. As with other globally distributed marine species, today's global loggerhead population has been shaped by a sequence of isolation events created by tectonic and oceanographic shifts over geologic time scales, the result of which is population substructuring in many areas (Bowen et al., 1994; Bowen, 2003). Globally, loggerhead turtles comprise a mosaic of populations, each with unique nesting sites and in many cases possessing disparate demographic features ( e.g., mean body size, age at first reproduction) (Dodd, 1988). However, despite these differences, loggerheads from different nesting populations often mix in common foraging areas during certain life stages (Bolten and Witherington, 2003), thus creating unique challenges when attempting to delineate distinct population segments for management or listing purposes.

Bowen et al. (1994) examined the mtDNA sequence diversity of loggerheads across their global distribution and found a separation of loggerheads in the Atlantic-Mediterranean basins from those in the Indo-Pacific basins since the Pleistocene period. The divergence between these two primary lineages corresponds to approximately three million years (2 percent per million years; Dutton et al., 1996; Encalada et al., 1996). Geography and climate appear to have shaped the evolution of these two matriarchal lineages with the onset of glacial cycles, the appearance of the Panama Isthmus creating a land barrier between the Atlantic and eastern Pacific, and upwelling of cold water off southern Africa creating an oceanographic barrier between the Atlantic and Indian Oceans (Bowen, 2003). Recent warm temperatures during interglacial periods allowed bi-directional invasion by the temperate-adapted loggerheads into the respective basins (Bowen et al., 1994; J.S. Reece, Washington University, personal communication, 2008)

Atlantic and eastern Pacific, and upwelling of cold water off southern Africa creating an oceanographic barrier between the Atlantic and Indian Oceans (Bowen, 2003). Recent warm temperatures during interglacial periods allowed bi-directional invasion by the temperate-adapted loggerheads into the respective basins (Bowen et al., 1994; J.S. Reece, Washington University, personal communication, 2008). Today, it appears that loggerheads within a basin are effectively isolated from populations in the other basin, but some dispersal from the Tongaland rookery in the Indian Ocean into feeding and developmental habitat in the South Atlantic is possible via the Agulhas Current (G.R. Hughes, unpublished data, cited in Bowen et al., 1994). In the Pacific, extensive mtDNA studies show that the northern loggerhead populations are isolated from the southern Pacific populations, and that juvenile loggerheads from these distinct genetic populations do not disperse across the equator (Hatase et al., 2002a; Dutton, 2007, unpublished data).

Mitochondrial DNA data indicate that regional turtle rookeries within an ocean basin have been strongly isolated from one another over ecological timescales (Bowen et al., 1994; Bowen and Karl, 2007). These same data indicate strong female natal homing and suggest that each regional nesting population is an independent demographic unit (Bowen and Karl, 2007). It is difficult to determine the precise boundaries of these demographically independent populations in regions, such as the eastern U.S. coast, where rookeries are close to each other and range along large areas of a continental coastline. There appear to be varying levels of connectivity between proximate rookeries facilitated by imprecise natal homing and male mediated gene flow (Pearce, 2001; Bowen, 2003; Bowen et al., 2005). Regional genetic populations often are characterized by allelic frequency differences rather than fixed genetic differences

ies are close to each other and range along large areas of a continental coastline. There appear to be varying levels of connectivity between proximate rookeries facilitated by imprecise natal homing and male mediated gene flow (Pearce, 2001; Bowen, 2003; Bowen et al., 2005). Regional genetic populations often are characterized by allelic frequency differences rather than fixed genetic differences.

Through the evaluation of genetic data, tagging data, telemetry, and demography, the BRT determined that there are at least nine discrete population segments of loggerhead sea turtles globally. These discrete population segments are markedly separated from each other as a consequence of physical, ecological, behavioral, and oceanographic factors, and given the genetic evidence, the BRT concluded that each regional population identified is discrete from other populations of loggerheads. Information considered by the BRT in its delineation of discrete population segments is presented below by ocean basin.

Pacific Ocean

In the North Pacific Ocean, the primary loggerhead nesting areas are found along the southern Japanese coastline and Ryukyu Archipelago (Kamezaki et al., 2003), although low level nesting may occur outside Japan in areas surrounding the South China Sea (Chan et al., 2007). Loggerhead turtles hatching on Japanese beaches undertake extensive developmental migrations using the Kuroshio and North Pacific Currents (Balazs, 2006; Kobayashi et al., 2008), and some turtles reach the vicinity of Baja California in the eastern Pacific (Uchida and Teruya, 1988; Bowen et al., 1995; Peckham et al., 2007). After spending years foraging in the central and eastern Pacific, loggerheads return to their natal beaches for reproduction (Resendiz et al., 1998; Nichols et al., 2000) and remain in the western Pacific for the remainder of their life cycle (Iwamoto et al., 1985; Kamezaki et al., 1997; Sakamoto et al., 1997; Hatase et al., 2002c)

chida and Teruya, 1988; Bowen et al., 1995; Peckham et al., 2007). After spending years foraging in the central and eastern Pacific, loggerheads return to their natal beaches for reproduction (Resendiz et al., 1998; Nichols et al., 2000) and remain in the western Pacific for the remainder of their life cycle (Iwamoto et al., 1985; Kamezaki et al., 1997; Sakamoto et al., 1997; Hatase et al., 2002c).

Despite the long-distance developmental movements of loggerheads in the North Pacific, current scientific evidence, based on genetic analysis, flipper tag recoveries, and satellite telemetry, indicates that individuals originating from Japan remain in the North Pacific for their entire life cycle, never crossing the equator or mixing with individuals from the South Pacific (Hatase et al., 2002a; LeRoux and Dutton, 2006; Dutton, 2007, unpublished data). This apparent, almost complete separation of two adjacent populations most likely results from: (1) The presence of two distinct Northern and Southern Gyre (current flow) systems in the Pacific (Briggs, 1974), (2) near-passive movements of post-hatchlings in these gyres that initially move them farther away from areas of potential mixing among the two populations along the equator, and (3) the nest-site fidelity of adult turtles that prevents turtles from returning to non-natal nesting areas.

Pacific loggerheads are further partitioned evolutionarily from other loggerheads throughout the world based on additional analyses of mtDNA. The haplotypes (a haplotype refers to the genetic signature, coded in mtDNA, of an individual) from both North and South Pacific loggerheads are distinguished by a minimum genetic distance (d) equal to 0.017 from other conspecifics, which indicates isolation of approximately one million years (Bowen, 2003).

Within the Pacific, Bowen et al

roughout the world based on additional analyses of mtDNA. The haplotypes (a haplotype refers to the genetic signature, coded in mtDNA, of an individual) from both North and South Pacific loggerheads are distinguished by a minimum genetic distance (d) equal to 0.017 from other conspecifics, which indicates isolation of approximately one million years (Bowen, 2003).

Within the Pacific, Bowen et al. (1995) used mtDNA to identify two genetically distinct nesting populations in the Pacific—a northern hemisphere population nesting in Japan and a southern hemisphere population nesting primarily in Australia. This study also et al., 1995). However, more extensive mtDNA rookery data from Japan (Hatase et al., 2002a) taken together with preliminary results from microsatellite (nuclear) analysis confirms that loggerheads inhabiting the North Pacific actually originate from nesting beaches in Japan (P. Dutton, NMFS, unpublished data). LeRoux et al. (2008) reported additional genetic variation in North Pacific loggerheads based on analyses using new mtDNA primers designed to target longer mtDNA sequences, and suggested finer scale population structure in North Pacific loggerheads may be present.

Although these studies indicate genetic distinctness between loggerheads nesting in Japan versus those nesting in Australia, Bowen et al. (1995) did identify individuals with the common Australian haplotype at foraging areas in the North Pacific, based on a few individuals sampled as bycatch in the North Pacific. More recently, Hatase et al. (2002a) detected this common haplotype at very low frequency at Japanese nesting beaches. However, the presence of the common Australian haplotype does not preclude the genetic distinctiveness of Japanese and Australian nesting populations, and is likely the result of rare gene flow events occurring over geologic time scales.

The discrete status of loggerheads in the North Pacific is further supported by results from flipper tagging in the North Pacific

y at Japanese nesting beaches. However, the presence of the common Australian haplotype does not preclude the genetic distinctiveness of Japanese and Australian nesting populations, and is likely the result of rare gene flow events occurring over geologic time scales.

The discrete status of loggerheads in the North Pacific is further supported by results from flipper tagging in the North Pacific. Flipper tagging of loggerheads has been widespread throughout this region, occurring on adults nesting in Japan and bycaught in the coastal pound net fishery (Y. Matsuzawa, Sea Turtle Association of Japan, personal communication, 2006), juvenile turtles reared and released in Japan (Uchida and Teruya, 1988; Hatase et al., 2002a), juvenile turtles foraging near Baja California, Mexico (Nichols, 2003; Seminoff et al., 2004), and juvenile and adult loggerheads captured in and tagged from commercial fisheries platforms in the North Pacific high seas (NMFS, unpublished data). To date, there have been at least three transPacific tag recoveries showing east-west and west-east movements (Uchida and Teruya, 1988; Resendiz et al., 1998; W.J. Nichols, Ocean Conservancy, and H. Peckham, Pro Peninsula, unpublished data) and several recoveries of adults in the western Pacific (Iwamoto et al., 1985; Kamezaki et al., 1997). However, despite the more than 30,000 marked individuals, not a single tag recovery has been reported outside the North Pacific.

A lack of movements by loggerheads south across the equator has also been supported by extensive satellite telemetry

and H. Peckham, Pro Peninsula, unpublished data) and several recoveries of adults in the western Pacific (Iwamoto et al., 1985; Kamezaki et al., 1997). However, despite the more than 30,000 marked individuals, not a single tag recovery has been reported outside the North Pacific.

A lack of movements by loggerheads south across the equator has also been supported by extensive satellite telemetry. As with flipper tagging, satellite telemetry has been conducted widely in the North Pacific, with satellite transmitters being placed on adult turtles departing nesting beaches (Sakamoto et al., 1997; Japan Fisheries Resource Conservation Association, 1999; Hatase et al., 2002b, 2002c), on adult and juvenile turtles bycaught in pound nets off the coast of Japan (Sea Turtle Association of Japan, unpublished data), on headstarted juvenile turtles released in Japan (Balazs, 2006), on juvenile and adult turtles bycaught in the eastern and central North Pacific ( e.g., Kobayashi et al., 2008), and on juvenile turtles foraging in the eastern Pacific (Nichols, 2003; Peckham et al., 2007; J. Seminoff, NMFS, unpublished data). Of the nearly 200 loggerheads tracked using satellite telemetry in the North Pacific, none have moved south of the equator. These studies have demonstrated the strong association loggerheads show with oceanographic mesoscale features such as the Transition Zone Chlorophyll Front or the Kuroshio Current Bifurcation Region (Polovina et al., 2000, 2001, 2004, 2006; Etnoyer et al., 2006; Kobayashi et al., 2008). Kobayashi et al. (2008) demonstrated that loggerheads strongly track these zones even as they shift in location, suggesting that strong habitat specificity during the oceanic stage also contributes to the lack of mixing. Telemetry studies in foraging areas of the eastern Pacific, near Baja California, Mexico (Nichols, 2003; Peckham et al., 2007; H. Peckham, Pro Peninsula, unpublished data) and Peru (J

yashi et al. (2008) demonstrated that loggerheads strongly track these zones even as they shift in location, suggesting that strong habitat specificity during the oceanic stage also contributes to the lack of mixing. Telemetry studies in foraging areas of the eastern Pacific, near Baja California, Mexico (Nichols, 2003; Peckham et al., 2007; H. Peckham, Pro Peninsula, unpublished data) and Peru (J. Mangel, Pro Delphinus, unpublished data) similarly showed a complete lack of long distance north or south movements.

The North Pacific population of loggerheads appears to occupy an ecological setting distinct from other loggerheads, including those of the South Pacific population. This is the only known population of loggerheads to be found north of the equator in the Pacific Ocean, foraging in the eastern Pacific as far south as Baja California Sur, Mexico (Seminoff et al., 2004; Peckham et al., 2007) and in the western Pacific as far south as the Philippines (Limpus, 2009) and the mouth of Mekong River, Vietnam (Sadoyama et al., 1996). Pelagic juvenile turtles spend much of their time foraging in the central and eastern North Pacific Ocean. The Kuroshio Extension Current, lying west of the international date line, serves as the dominant physical and biological habitat in the North Pacific and is highly productive, likely due to unique features such as eddies and meanders that concentrate prey and support food webs. Juvenile loggerheads originating from nesting beaches in Japan exhibit high site fidelity to an area referred to as the Kuroshio Extension Bifurcation Region, an area with extensive meanders and mesoscale eddies (Polovina et al., 2006). Juvenile turtles also were found to correlate strongly with areas of surface chlorophyll a levels in an area known as the Transition Zone Chlorophyll Front, an area concentrating surface prey for loggerheads (Polovina et al., 2001; Parker et al., 2005; Kobayashi et al., 2008)

as the Kuroshio Extension Bifurcation Region, an area with extensive meanders and mesoscale eddies (Polovina et al., 2006). Juvenile turtles also were found to correlate strongly with areas of surface chlorophyll a levels in an area known as the Transition Zone Chlorophyll Front, an area concentrating surface prey for loggerheads (Polovina et al., 2001; Parker et al., 2005; Kobayashi et al., 2008). Another area found ecologically unique to the North Pacific population of loggerheads, likely because of the high density of pelagic red crabs ( Pleuronocodes planipes ), is located off the Pacific coast of the Baja California Peninsula, Mexico, where researchers have documented a foraging area for juvenile turtles based on aerial surveys and satellite telemetry (Seminoff et al., 2006; Peckham et al., 2007). Tag returns show post-nesting females migrating into the East China Sea off South Korea, China, and the Philippines, and the nearby coastal waters of Japan (Iwamoto et al., 1985; Kamezaki et al., 1997, 2003). Clearly, the North Pacific population of loggerheads is uniquely adapted to the ecological setting of the North Pacific Ocean and serves as an important part of the ecosystem it inhabits.

In summary, loggerheads inhabiting the North Pacific Ocean are derived primarily, if not entirely, from Japanese beaches (although low level nesting may occur outside Japan in areas surrounding the South China Sea), with the possible exception of rare waifs over evolutionary time scales. Further, nesting colonies of Japanese loggerheads are found to be genetically distinct based on mtDNA analyses, and when compared to much larger and more genetically diverse loggerhead populations in the Atlantic and Mediterranean, Pacific loggerheads have likely experienced critical bottlenecks (in Hatase et al., 2002a), underscoring the importance of conservation and management to retain this genetically distinct population

Japanese loggerheads are found to be genetically distinct based on mtDNA analyses, and when compared to much larger and more genetically diverse loggerhead populations in the Atlantic and Mediterranean, Pacific loggerheads have likely experienced critical bottlenecks (in Hatase et al., 2002a), underscoring the importance of conservation and management to retain this genetically distinct population.

In the South Pacific Ocean, loggerhead turtles nest primarily in Queensland, Australia, and, to a lesser extent, New Caledonia and Vanuatu (Limpus and Limpus, 2003; Limpus et al., 2006; Limpus, 2009). Loggerheads from these rookeries undertake an oceanic developmental migration, et al., 2004; Donoso and Dutton, 2006, 2007; Boyle et al., 2009). Large juvenile and adult loggerheads generally remain in the western South Pacific, inhabiting neritic and oceanic foraging sites during non-nesting periods (Limpus et al., 1994; Limpus, 2009).

Loggerheads from Australia and New Caledonia apparently do not travel north of the equator. Flipper tag recoveries from nesting females have been found throughout the western Pacific, including sites north of Australia, the Torres Straight, and the Gulf of Carpentaria (Limpus, 2009). Of approximately 1,000 (adult and juvenile; male and female) loggerheads that have been tagged in eastern Australian feeding areas, only two have been recorded nesting outside of Australia; both traveled to New Caledonia (Limpus, 2009). Flipper tagging programs in Peru and Chile tagged approximately 500 loggerheads from 1999 to 2006, none of which have been reported from outside of the southeastern Pacific (Alfaro-Shigueto et al., 2008a; S. Kelez, Duke University Marine Laboratory, unpublished data; M. Donoso, ONG Pacifico Laud—Chile, unpublished data). Limited satellite telemetry data from 12 turtles in the area show a similar trend (J. Mangel, Pro Delphinus, unpublished data)

approximately 500 loggerheads from 1999 to 2006, none of which have been reported from outside of the southeastern Pacific (Alfaro-Shigueto et al., 2008a; S. Kelez, Duke University Marine Laboratory, unpublished data; M. Donoso, ONG Pacifico Laud—Chile, unpublished data). Limited satellite telemetry data from 12 turtles in the area show a similar trend (J. Mangel, Pro Delphinus, unpublished data).

The spatial separation between the North Pacific and South Pacific loggerhead populations has contributed to substantial differences in the genetic profiles of the nesting populations in these two regions. Whereas the dominant mtDNA haplotypes among loggerheads nesting in Japan are CCP2 and CCP3 (equivalent to B and C respectively in Bowen et al., 1995 and Hatase et al., 2002a; LeRoux et al., 2008; P. Dutton, NMFS, unpublished data), loggerheads nesting in eastern Australia have a third haplotype (CCP1, previously A) which is dominant (98 percent of nesting females) (Bowen et al., 1994; FitzSimmons et al., 1996; Boyle et al., 2009). Further, preliminary genetic analysis using microsatellite markers (nuclear DNA) indicates genetic distinctiveness between nesting populations in the North versus South Pacific (P. Dutton, NMFS, personal communication, 2008).

The separateness between nesting populations in eastern Australia (in the South Pacific Ocean) and western Australia (in the East Indian Ocean) is less clear, although these too are considered to be genetically distinct from one another (Limpus, 2009). For example, mtDNA haplotype CCP1, which is the overwhelmingly dominant haplotype among eastern Australia nesting females (98 percent), is also found in western Australia, although at much lower frequency (33 percent) (FitzSimmons et al., 1996, 2003). The remaining haplotype for both regions was the CCP5 haplotype. Further, FitzSimmons (University of Canberra, unpublished data) found significant differences in nuclear DNA microsatellite loci from females nesting in these two regions

eastern Australia nesting females (98 percent), is also found in western Australia, although at much lower frequency (33 percent) (FitzSimmons et al., 1996, 2003). The remaining haplotype for both regions was the CCP5 haplotype. Further, FitzSimmons (University of Canberra, unpublished data) found significant differences in nuclear DNA microsatellite loci from females nesting in these two regions. Estimates of gene flow between eastern and western Australian populations was an order of magnitude less than gene flow within regions. These preliminary results based on nuclear DNA indicate that male-mediated gene flow between eastern and western Australia may be insignificant, which, when considered in light of the substantial disparity in mtDNA haplotype frequencies between these two regions, provides further evidence of population separation.

At present, there is no indication from genetic studies that the loggerhead turtles nesting in eastern Australia are distinct from those nesting in New Caledonia. Of 27 turtles sequenced from New Caledonia, 93 percent carried the CCP1 haplotype and the remaining had the CCP5 haplotype; similar to eastern Australia (Boyle et al., 2009).

The South Pacific population of loggerheads occupies an ecological setting distinct from other loggerheads, including the North Pacific population; however, less is known about the ecosystem on which South Pacific oceanic juvenile and adult loggerheads depend. Sea surface temperature and chlorophyll frontal zones in the South Pacific have been shown to dramatically affect the movements of green turtles, Chelonia mydas (Seminoff et al., 2008) and leatherback turtles, Dermochelys coriacea (Shillinger et al., 2008), and it is likely that loggerhead distributions are also affected by these mesoscale oceanographic features.

Loggerheads in the South Pacific are substantially impacted by periodic environmental perturbations such as the El Niño Southern Oscillation (ENSO)

he movements of green turtles, Chelonia mydas (Seminoff et al., 2008) and leatherback turtles, Dermochelys coriacea (Shillinger et al., 2008), and it is likely that loggerhead distributions are also affected by these mesoscale oceanographic features.

Loggerheads in the South Pacific are substantially impacted by periodic environmental perturbations such as the El Niño Southern Oscillation (ENSO). This 3- to 6-year cycle within the coupled ocean-atmosphere system of the tropical Pacific brings increased surface water temperatures and lower primary productivity, both of which have profound biological consequences (Chavez et al., 1999). Loggerheads are presumably adversely impacted by the reduced food availability that often results from ENSO events, although data on this subject are lacking. Although ENSO may last for only short periods and thus not have a long-term effect on loggerheads in the region, recent studies by Chaloupka et al. (2008) suggested that long-term increases in sea surface temperature within the South Pacific may influence the ability of the Australian nesting population to recover from historic population declines.

Loggerheads originating from nesting beaches in the western South Pacific are the only population of loggerheads to be found south of the equator in the Pacific Ocean. As post-hatchlings, they are generally swept south by the East Australian Current (Limpus et al., 1994), spend a large portion of time foraging in the oceanic South Pacific Ocean, and some migrate to the southeastern Pacific Ocean off the coasts of Peru and Chile as juvenile turtles (Alfaro-Shigueto et al., 2004; Donoso et al., 2000; Boyle et al., 2009). As large juveniles and adults, these loggerheads' foraging range encompasses the eastern Arafura Sea, Gulf of Carpentaria, Torres Strait, Gulf of Papua, Coral Sea, and western Tasman Sea to southern New South Wales including the Great Barrier Reef, Hervey Bay, and Moreton Bay

f the coasts of Peru and Chile as juvenile turtles (Alfaro-Shigueto et al., 2004; Donoso et al., 2000; Boyle et al., 2009). As large juveniles and adults, these loggerheads' foraging range encompasses the eastern Arafura Sea, Gulf of Carpentaria, Torres Strait, Gulf of Papua, Coral Sea, and western Tasman Sea to southern New South Wales including the Great Barrier Reef, Hervey Bay, and Moreton Bay. The outer extent of this range includes the coastal waters off eastern Indonesia northeastern Papua New Guinea, northeastern Solomon Islands, and New Caledonia (in Limpus, 2009).

In summary, all loggerheads inhabiting the South Pacific Ocean are derived from beaches in eastern Australia and a lesser known number of beaches in southern New Caledonia, Vanuatu, and Tokelau (Limpus and Limpus, 2003; Limpus, 2009). Furthermore, nesting colonies of the South Pacific population of loggerheads are found to be genetically distinct from loggerheads in the North Pacific and Indian Ocean.

Given the information presented above, the BRT concluded, and we concur, that two discrete population segments exist in the Pacific Ocean: (1) North Pacific Ocean and (2) South Pacific Ocean. These two population segments are markedly separated from et al., 2002a; LeRoux and Dutton, 2006; Dutton, 2007, unpublished data). This apparent, almost complete separation most likely results from: (1) The presence of two distinct Northern and Southern Gyre (current flow) systems in the Pacific (Briggs, 1974), (2) near-passive movements of post-hatchlings in these gyres that initially move them farther away from areas of potential mixing along the equator, and (3) the nest-site fidelity of adult turtles that prevents turtles from returning to non-natal nesting areas. The separation of the Pacific Ocean population segments from population segments within the Indian Ocean and Atlantic Ocean basins is believed to be the result of land barriers and oceanographic barriers. Based on mtDNA analysis, Bowen et al

areas of potential mixing along the equator, and (3) the nest-site fidelity of adult turtles that prevents turtles from returning to non-natal nesting areas. The separation of the Pacific Ocean population segments from population segments within the Indian Ocean and Atlantic Ocean basins is believed to be the result of land barriers and oceanographic barriers. Based on mtDNA analysis, Bowen et al. (1994) found a separation of loggerheads in the Atlantic-Mediterranean basins from those in the Indo-Pacific basins since the Pleistocene period. Geography and climate appear to have shaped the evolution of these two matriarchal lineages with the onset of glacial cycles, the appearance of the Panama Isthmus creating a land barrier between the Atlantic and eastern Pacific, and upwelling of cold water off southern Africa creating an oceanographic barrier between the Atlantic and Indian Oceans (Bowen, 2003).

Indian Ocean

Similar to loggerheads in the Pacific and Atlantic, loggerheads in the Indian Ocean nest on coastal beaches, forage in neritic and oceanic habitats, and undertake long-distance migrations between and within these areas. The distribution of loggerheads in the Indian Ocean is limited by the Asian landmass to the north (approximately 30° N latitude); distributions east and west are not restricted by landmasses south of approximately 38° S latitude.

Historical accounts of loggerhead turtles in the Indian Ocean are found in Smith (1849), who described the species in South Africa, and Deraniyagala (1933, 1939) who described Indian Ocean loggerheads within the subspecies C. c. gigas. Hughes (1974) argued that there was little justification for this separation.

South Africa, and Deraniyagala (1933, 1939) who described Indian Ocean loggerheads within the subspecies C. c. gigas. Hughes (1974) argued that there was little justification for this separation.

In the North Indian Ocean, Oman hosts the vast majority of loggerhead nesting. The largest nesting assemblage is at Masirah Island, Oman, in the northern tropics at 21° N latitude (Baldwin et al., 2003). Other key nesting assemblages occur on the Al Halaniyat Islands, Oman (17° S latitude) and on Oman's Arabian Sea mainland beaches south of Masirah Island to the Oman-Yemen border (17-20° S latitude) (IUCN—The World Conservation Union, 1989a, 1989b; Salm, 1991; Salm and Salm, 1991; Baldwin et al., 2003). In addition, nesting probably occurs on the mainland of Yemen on the Arabian Sea coast, and nesting has been confirmed on Socotra, an island off the coast of Yemen (Pilcher and Saad, 2000).

Outside of Oman, loggerhead nesting is rare in the North Indian Ocean. The only verified nesting beaches for loggerheads on the Indian subcontinent are found in Sri Lanka (Deraniyagala, 1939; Kar and Bhaskar, 1982; Dodd, 1988; Kapurusinghe, 2006). Reports of regular loggerhead nesting on the Indian mainland are likely misidentifications of olive ridleys ( Lepidochelys olivacea ) (Tripathy, 2005; Kapurusinghe, 2006). Although loggerheads have been reported nesting in low numbers in Myanmar, these data may not be reliable because of misidentification of species (Thorbjarnarson et al., 2000).

n low numbers in Myanmar, these data may not be reliable because of misidentification of species (Thorbjarnarson et al., 2000).

Limited information exists on foraging locations of North Indian Ocean loggerheads. Foraging individuals have been reported off the southern coastline of Oman (Salm et al., 1993) and in the Gulf of Mannar, between Sri Lanka and India (Tripathy, 2005; Kapurusinghe, 2006). Satellite telemetry studies of post-nesting migrations of loggerheads nesting on Masirah Island, Oman, have revealed extensive use of the waters off the Arabian Peninsula, with the majority of telemetered turtles (15 of 20) traveling southwest, following the shoreline of southern Oman and Yemen, and circling well offshore in nearby oceanic waters (Environment Society of Oman and Ministry of Environment and Climate Change, Oman, unpublished data). A minority traveled north as far as the western Persian (Arabian) Gulf (3 of 20) or followed the shoreline of southern Oman and Yemen as far west as the Gulf of Aden and the Bab-el-Mandab (2 of 20). These preliminary data suggest that post-nesting migrations and adult female foraging areas may be centered within the region (Environment Society of Oman and Ministry of Environment and Climate Change, Oman, unpublished data). No tag returns or satellite tracks indicated that loggerheads nesting in Oman traveled south of the equator.

North West Cape and to the nearby Muiron Islands (Baldwin et al., 2003). Nesting individuals from Dirk Hartog Island have been recorded foraging within Shark Bay and Exmouth Gulf, while other adults range into the Gulf of Carpentaria (Baldwin et al., 2003). At the eastern extent of this apparent range, there is possible overlap with loggerheads that nest on Australia's Pacific coast (Limpus, 2009). However, despite extensive tagging at principal nesting beaches on Australia's Indian Ocean and Pacific coasts, no exchange of females between nesting beaches has been observed (Limpus, 2009).

Loggerhead nesting in the Southwest Indian Ocean includes the southeastern coast of Africa from the Paradise Islands in Mozambique southward to St. Lucia in South Africa, and on the south and southwestern coasts of Madagascar (Baldwin et al., 2003). Foraging habitats are only known for the Tongaland, South Africa, adult female loggerheads. Returns of flipper tags describe a range that extends eastward to Madagascar, northward to Mozambique, Tanzania, and Kenya, and southward to Cape Agulhas at the southernmost point of Africa (Baldwin et al., 2003). Four post-nesting loggerheads satellite tracked by Luschi et al. (2006) migrated northward, hugging the Mozambique coast and remained in shallow shelf waters off Mozambique for more than 2 months. Only one post-nesting female from the Southwest Indian Ocean population (South Africa) has been documented migrating north of the equator (to southern Somalia) (Hughes and Bartholomew, 1996).

omew, 1996).

The available genetic information relates to connectivity and broad evolutionary relationships between ocean basins. There is a lack of genetic information on population structure among rookeries within the Indian Ocean. Bowen et al. (1994) described mtDNA sequence diversity among eight loggerhead nesting assemblages and found one of two principal branches in the Indo-Pacific basins. Using additional published and unpublished data, Bowen (2003) estimated divergence between these two lineages to be approximately three million years. Bowen pointed out evidence for more recent colonizations (12,000-250,000 years ago) between the Indian Ocean and the Atlantic- et al. (1994) at Masirah Island, Oman, is known from the Atlantic and suggests some exchange between oceans some 250,000 years ago. The other principal Indian Ocean haplotype reported by Bowen et al. (1994) was seen in all loggerheads sampled (n=15) from Natal, South Africa. Encalada et al. (1998) reported that this haplotype was common throughout the North Atlantic and Mediterranean, thus suggesting a similar exchange between the Atlantic and Indian Oceans as recently as 12,000 years ago (Bowen et al., 1994). Bowen (2003) speculated that Indian-Atlantic Ocean exchanges took place via the temperate waters south of South Africa and became rare as the ocean shifted to cold temperate conditions in this region.

To estimate loggerhead gene flow in and out of the Indian Ocean, J.S. Reece (Washington University, personal communication, 2008) examined 100 samples from Masirah Island, 249 from Atlantic rookeries (from Encalada et al., 1998), and 311 from Pacific rookeries (from Hatase et al., 2002a and Bowen et al., 1995). Reece estimated that gene flow, expressed as number of effective migrants, or exchanges of breeding females between Indian Ocean rookeries and those from the Atlantic or Pacific occurred at the rate of less than 0.1 migrant per generation

from Masirah Island, 249 from Atlantic rookeries (from Encalada et al., 1998), and 311 from Pacific rookeries (from Hatase et al., 2002a and Bowen et al., 1995). Reece estimated that gene flow, expressed as number of effective migrants, or exchanges of breeding females between Indian Ocean rookeries and those from the Atlantic or Pacific occurred at the rate of less than 0.1 migrant per generation. Reece estimated gene flow based on coalescence of combined mtDNA and nuclear DNA data to be approximately 0.5 migrants per generation. These unpublished results, while somewhat theoretical, may indicate that there is restricted gene flow into and out of the Indian Ocean. The low level of gene flow most likely reflects the historical connectivity over geological timescales rather than any contemporary migration, and is consistent with Bowen's hypothesis that exchange occurred most recently over 12,000-3,000,000 years ago, and has been restricted over recent ecological timescales.

The discrete status of three loggerhead populations in the Indian Ocean is primarily supported by observations of tag returns and satellite telemetry. The genetic information currently available based on mtDNA sequences does not allow for a comprehensive analysis of genetic population structure analysis for Indian Ocean rookeries, although Bowen et al. (1994) indicated the Oman and South African rookeries are genetically distinct, and once sequencing studies are completed for these rookeries, it is likely that they will also be genetically distinct from the rookeries in western Australia. Based on multiple lines of evidence, discrete status is supported for the North Indian Ocean, Southeast Indo-Pacific Ocean, and Southwest Indian Ocean loggerhead populations. Although there is not a sufficiently clear picture of gene flow between these regions, significant vicariant barriers likely exist between these three Indian Ocean populations that would prevent migration of individuals on a time scale relative to management and conservation efforts

pported for the North Indian Ocean, Southeast Indo-Pacific Ocean, and Southwest Indian Ocean loggerhead populations. Although there is not a sufficiently clear picture of gene flow between these regions, significant vicariant barriers likely exist between these three Indian Ocean populations that would prevent migration of individuals on a time scale relative to management and conservation efforts. These vicariant barriers are the oceanographic phenomena associated with Indian Ocean equatorial waters, and the large expanse between continents in the South Indian Ocean without suitable benthic foraging habitat.

Given the information presented above, the BRT concluded, and we concur, that three discrete population segments exist in the Indian Ocean: (1) North Indian Ocean, (2) Southeast Indo-Pacific Ocean, and (3) Southwest Indian Ocean. These three population segments are markedly separated from each other and from population segments within the Pacific Ocean and Atlantic Ocean basins as a consequence of physical, ecological, behavioral, and oceanographic factors. Information supporting this conclusion is primarily based on observations of tag returns and satellite telemetry. The genetic information currently available based on mtDNA sequences does not allow for a comprehensive analysis of genetic population structure for Indian Ocean rookeries; however, the Oman and South African rookeries are genetically distinct, and once sequencing studies are completed for these rookeries, it is likely that they will also be determined genetically distinct from the rookeries in western Australia (Bowen et al. 1994). Furthermore, significant vicariant barriers ( i.e., oceanographic phenomena associated with Indian Ocean equatorial waters, and the large expanse between continents in the South Indian Ocean without suitable benthic foraging habitat) likely exist between these three Indian Ocean populations that would prevent migration of individuals on a time scale relative to management and conservation efforts

Furthermore, significant vicariant barriers ( i.e., oceanographic phenomena associated with Indian Ocean equatorial waters, and the large expanse between continents in the South Indian Ocean without suitable benthic foraging habitat) likely exist between these three Indian Ocean populations that would prevent migration of individuals on a time scale relative to management and conservation efforts. The separation of the Indian Ocean population segments from population segments within the Pacific Ocean and Atlantic Ocean basins is believed to be the result of land barriers and oceanographic barriers. Based on mtDNA analysis, Bowen et al. (1994) found a separation of loggerheads in the Atlantic-Mediterranean basins from those in the Indo-Pacific basins since the Pleistocene period. Geography and climate appear to have shaped the evolution of these two matriarchal lineages with the onset of glacial cycles, the appearance of the Panama Isthmus creating a land barrier between the Atlantic and eastern Pacific, and upwelling of cold water off southern Africa creating an oceanographic barrier between the Atlantic and Indian Oceans (Bowen, 2003). In the East Indian Ocean, although there is possible overlap with loggerheads that nest on Australia's Indian Ocean and Pacific Ocean coasts, extensive tagging at the principal nesting beaches on both coasts has revealed no exchange of females between these nesting beaches (Limpus, 2009).

Atlantic Ocean and Mediterranean Sea

Within the Atlantic Ocean, loss and re-colonization of nesting beaches over evolutionary time scales has been influenced by climate, natal homing, and rare dispersal events (Encalada et al., 1998; Bowen and Karl, 2007). At times, temperate beaches were too cool to incubate eggs and nesting could have succeeded only on tropical beaches. Thus, the contemporary distribution of nesting is the product of colonization events from the tropical refugia during the last 12,000 years

volutionary time scales has been influenced by climate, natal homing, and rare dispersal events (Encalada et al., 1998; Bowen and Karl, 2007). At times, temperate beaches were too cool to incubate eggs and nesting could have succeeded only on tropical beaches. Thus, the contemporary distribution of nesting is the product of colonization events from the tropical refugia during the last 12,000 years. Apparently, turtles from the Northwest Atlantic colonized the Mediterranean and at least two matrilines were involved (Schroth et al., 1996); these rookeries became isolated from the Atlantic populations in the last 10,000 years (Encalada et al., 1998). A similar colonization event appears to have populated the Northeast Atlantic (C. Monzon-Arguello, Instituto Canario de Ciencias Marinas—Spain, personal communication, 2008).

Nesting in the western South Atlantic occurs primarily along the mainland coast of Brazil from Sergipe south to Rio de Janeiro, with peak concentrations in northern Bahia, Espírito Santo, and northern Rio de Janeiro (Marcovaldi and Chaloupka, 2007). In the eastern South Atlantic, diffuse nesting may occur along the mainland coast of Africa (Fretey, 2001), with more than 200 loggerhead nests reported for Rio Longa beach in central Angola in 2005 (Brian, 2007). However, other researchers have been unable to confirm nesting by loggerheads in the last decade anywhere along the south Atlantic coast of Africa, including Angola (Fretey, 2001; Weir et al., 2007). There is the possibility that reports of nesting loggerheads from Angola and Namibia (Márquez M., 1990; Brian, 2007) may have arisen from misidentified olive ridley turtles (Brongersma, 1982; Fretey, 2001). At the current time, it is not possible to confirm that regular, if any, nesting of

Genetic surveys of loggerheads have revealed that the Brazilian rookeries have a unique mtDNA haplotype (Encalada et al., 1998; Pearce, 2001)

rts of nesting loggerheads from Angola and Namibia (Márquez M., 1990; Brian, 2007) may have arisen from misidentified olive ridley turtles (Brongersma, 1982; Fretey, 2001). At the current time, it is not possible to confirm that regular, if any, nesting of

Genetic surveys of loggerheads have revealed that the Brazilian rookeries have a unique mtDNA haplotype (Encalada et al., 1998; Pearce, 2001). The Brazilian mtDNA haplotype, relative to North Atlantic haplotypes, indicates isolation of South Atlantic loggerheads from North Atlantic loggerheads on a scale of 250,000-500,000 years ago, and microsatellite DNA results show divergence on the same time scale (Bowen, 2003). Brazil's unique haplotype has been found only in low numbers in foraging populations of juvenile loggerheads of the North Atlantic (Bass et al., 2004). Other lines of evidence support a deep division between loggerheads from the South Atlantic and from the North Atlantic, including: (1) A nesting season in Brazil that peaks in the austral summer around December-January (Marcovaldi and Laurent, 1996), as opposed to the April-September nesting season in the southeastern United States in the northern hemisphere (Witherington et al., 2009); and (2) no observations of tagged loggerheads moving across the equator in the Atlantic, except a single case of a captive-reared animal that was released as a juvenile from Espírito Santo and was recaptured 3 years later in the Azores (Bolten et al., 1990). Post-nesting females from Espírito Santo, Brazil, moved either north or south along the coast, but remained between 10° S latitude and 30° S latitude (Projeto TAMAR, unpublished data).

latitude and 30° S latitude (Projeto TAMAR, unpublished data).

Relatively little is known about the at-sea behavior of loggerheads originating from nesting beaches in Brazil. Recaptures of tagged juvenile turtles and nesting females have shown movement of animals up and down the coast of South America (Almeida et al., 2000; Marcovaldi et al., 2000; Laporta and Lopez, 2003; Almeida et al., 2007). Juvenile loggerheads, presumably of Brazilian origin, have also been captured on the high seas of the South Atlantic (Kotas et al., 2004; Pinedo and Polacheck, 2004) and off the coast of Atlantic Africa (Petersen, 2005; Petersen et al., 2007; Weir et al., 2007) suggesting that, like their North Pacific and Northwest Atlantic counterparts, loggerheads of the South Atlantic may undertake transoceanic developmental migrations (Bolten et al., 1998; Peckham et al., 2007).

The mean size of reproductive female loggerheads in Brazil is 92.9 cm straight carapace length (SCL), which is comparable to the size of nesting females in the Northwest Atlantic, but larger than nesting females in the Northeast Atlantic and Mediterranean (Tiwari and Bjorndal, 2000; Margaritoulis et al., 2003; Varo Cruz et al., 2007). Egg size and mass of Brazilian loggerheads are smaller than those from the Northwest Atlantic, but larger than those of the Mediterranean (Tiwari and Bjorndal, 2000).

Within the Northwest Atlantic, the majority of nesting activity occurs from April through September, with a peak in June and July (Williams-Walls et al., 1983; Dodd, 1988; Weishampel et al., 2006). 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 United States and on the Yucatan Peninsula in Mexico (Sternberg, 1981; Ehrhart, 1989; Ehrhart et al., 2003; NMFS and USFWS, 2008). Many nesting beaches within the Northwest Atlantic have yet to be sampled for genetic analysis

thin 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 United States and on the Yucatan Peninsula in Mexico (Sternberg, 1981; Ehrhart, 1989; Ehrhart et al., 2003; NMFS and USFWS, 2008). Many nesting beaches within the Northwest Atlantic have yet to be sampled for genetic analysis. Five recovery units (subpopulations) have been identified based on genetic differences and a combination of geographic distribution of nesting densities and geographic separation. These recovery units are: Northern Recovery Unit (Florida/Georgia border through southern Virginia), Peninsular Florida Recovery Unit (Florida/Georgia border through Pinellas County, Florida), Northern Gulf of Mexico Recovery Unit (Franklin County, Florida, through Texas), Greater Caribbean Recovery Unit (Mexico through French Guiana, The Bahamas, Lesser Antilles, and Greater Antilles), and Dry Tortugas Recovery Unit (islands located west of Key West, Florida) (NMFS and USFWS, 2008). There is limited exchange of nesting females among these recovery units (Encalada et al., 1998; Foote et al., 2000; J. Richardson personal communication cited in NMFS, 2001; Hawkes et al., 2005). Based on the number of haplotypes, the highest level of loggerhead mtDNA genetic diversity in the Atlantic has been observed in females of the Greater Caribbean Recovery Unit that nest at Quintana Roo, Mexico (Encalada et al., 1999; Nielsen et al., in press). However, genetic diversity should be evaluated further using haplotype and nucleotide diversity calculated similarly for each recovery unit. Genetic data are not available for all the nesting assemblages in the region, including a key nesting assemblage in Cuba

ved in females of the Greater Caribbean Recovery Unit that nest at Quintana Roo, Mexico (Encalada et al., 1999; Nielsen et al., in press). However, genetic diversity should be evaluated further using haplotype and nucleotide diversity calculated similarly for each recovery unit. Genetic data are not available for all the nesting assemblages in the region, including a key nesting assemblage in Cuba. New genetic markers have recently been developed, including primers that produce additional mtDNA sequence data (Abreu-Grobois et al., 2006; LeRoux et al., 2008), and an array of microsatellite markers (Shamblin et al., 2008) that will enable finer resolution of population boundaries.

Loggerheads in the Northwest Atlantic display complex population structure based on life history stages. Based on mtDNA, oceanic juveniles show no structure, neritic juveniles show moderate structure, and nesting colonies show strong structure (Bowen et al., 2005). In contrast, a survey using microsatellite (nuclear DNA) markers showed no significant population structure among nesting populations (Bowen et al., 2005), indicating that while females exhibit strong philopatry, males may provide an avenue of gene flow between nesting colonies in this region. However, the power to detect structure with the nuclear markers used in this study may have been limited due to the few markers used and small sample sizes. Nevertheless, Bowen et al. (2005) argued that male-mediated gene flow within the Northwest Atlantic does not detract from the classification of breeding areas as independent populations ( e.g., recovery units) because the production of progeny depends on female nesting success. All Northwest Atlantic recovery units are reproductively isolated from populations within the Northeast Atlantic, South Atlantic, and Mediterranean Sea.

f progeny depends on female nesting success. All Northwest Atlantic recovery units are reproductively isolated from populations within the Northeast Atlantic, South Atlantic, and Mediterranean Sea.

As oceanic juveniles, loggerheads from the Northwest Atlantic use the North Atlantic Gyre and often are associated with Sargassum communities (Carr, 1987). They also are found in the Mediterranean Sea. In these areas, they overlap with animals originating from the Northeast Atlantic and the Mediterranean Sea (Laurent et al., 1993, 1998; Bolten et al., 1998; Bowen et al., 2005; LaCasella et al., 2005; Carreras et al., 2006; Monzon-Arguello et al., 2006; Revelles et al., 2007). In the western Mediterranean, they tend to be associated with the waters off the northern African coast and the northeastern Balearic Archipelago, areas generally not inhabited by turtles of Mediterranean origin (Carreras et al., 2006; Revelles et al., 2007; Eckert et al., 2008). As larger neritic juveniles, they show more structure and tend to inhabit areas closer to their natal origins (Bowen et al., 2004), but some do move to and from oceanic foraging grounds throughout this life stage (Mansfield, 2006; McClellan and Read, 2007), and some continue to use the Mediterranean Sea (Casale et al., 2008a; Eckert et al., 2008). Adult populations are highly structured with no overlap in distribution among adult loggerheads from the Northwest Atlantic, Northeast Atlantic, South Atlantic, and et al., 1983; Foley et al., 2008; Turtle Expert Working Group, 2009). Adult females from the Northern Gulf of Mexico Recovery Unit remained in the Gulf of Mexico, including off the Yucatan Peninsula of Mexico, based on satellite telemetry and flipper tag returns (Foley et al., 2008; Turtle Expert Working Group, 2009; M. Lamont, Florida Cooperative Fish and Wildlife Research Unit, personal communication, 2009; M. Nicholas, National Park Service, personal communication, 2009)

males from the Northern Gulf of Mexico Recovery Unit remained in the Gulf of Mexico, including off the Yucatan Peninsula of Mexico, based on satellite telemetry and flipper tag returns (Foley et al., 2008; Turtle Expert Working Group, 2009; M. Lamont, Florida Cooperative Fish and Wildlife Research Unit, personal communication, 2009; M. Nicholas, National Park Service, personal communication, 2009).

Nesting in the Northeast Atlantic is concentrated in the Cape Verde Archipelago, with some nesting occurring on most of the islands, and the highest concentration on the beaches of Boa Vista Island (Lopez-Jurado et al., 2000; Varo Cruz et al., 2007; Loureiro, 2008). On mainland Africa, there is minor nesting on the coasts of Mauritania to Senegal (Brongersma, 1982; Arvy et al., 2000; Fretey, 2001). Earlier reports of loggerhead nesting in Morocco (Pasteur and Bons, 1960) have not been confirmed in recent years (Tiwari et al., 2001). Nesting has not been reported from Macaronesia (Azores, Madeira Archipelago, The Selvagens Islands, and the Canary Islands), other than in the Cape Verde Archipelago (Brongersma, 1982). In Cape Verde, nesting begins in mid June and extends into October (Cejudo et al., 2000), which is somewhat later than when nesting occurs in the Northwest Atlantic.

Based on an analysis of mtDNA of 196 nesting females from Boa Vista Island, the Cape Verde nesting assemblage is genetically distinct from other studied rookeries (C. Monzon-Arguello, Instituto Canario de Ciencias Marinas—Spain, personal communication, 2008; Monzon-Arguello et al., 2009). The results also indicate that despite the close proximity of the Mediterranean, the Boa Vista rookery is most closely related to the rookeries of the Northwest Atlantic.

The distribution of juvenile loggerheads from the Northeast Atlantic is largely unknown but they have been found on the oceanic foraging grounds of the North Atlantic (A

sonal communication, 2008; Monzon-Arguello et al., 2009). The results also indicate that despite the close proximity of the Mediterranean, the Boa Vista rookery is most closely related to the rookeries of the Northwest Atlantic.

The distribution of juvenile loggerheads from the Northeast Atlantic is largely unknown but they have been found on the oceanic foraging grounds of the North Atlantic (A. Bolten, University of Florida, personal communication, 2008, based on Bolten et al., 1998 and LaCasella et al., 2005; Monzon-Arguello et al., 2009; M. Tiwari, NMFS, and A. Bolten, University of Florida, unpublished data) and in the western and central Mediterranean (A. Bolten, University of Florida, personal communication, 2008, based on Carreras et al., 2006), along with small juvenile loggerheads from the Northwest Atlantic. The size of nesting females in the Northeast Atlantic is comparable to those in the Mediterranean (average 72-80 cm SCL; Margaritoulis et al., 2003) and smaller than those in the Northwest Atlantic or the South Atlantic; 91 percent of the nesting turtles are less than 86.5 cm curved carapace length (CCL) (Hawkes et al., 2006) and nesting females average 77.1 cm SCL (Cejudo et al., 2000). Satellite-tagged, post-nesting females from Cape Verde foraged in coastal waters along northwest Africa or foraged oceanically, mostly between Cape Verde and the African shelf from Mauritania to Guinea Bissau (Hawkes et al., 2006).

In the Mediterranean, nesting occurs throughout the central and eastern basins on the shores of Italy, Greece, Cyprus, Turkey, Syria, Lebanon, Israel, the Sinai, Egypt, Libya, and Tunisia (Sternberg, 1981; Margaritoulis et al., 2003; SWOT, 2007). Sporadic nesting also has been reported in the western Mediterranean on Corsica (Delaugerre and Cesarini, 2004), southwestern Italy (Bentivegna et al., 2005), and on the Spanish Mediterranean coast (Tomas et al., 2003, 2008). Nesting in the Mediterranean is concentrated between June and early August (Margaritoulis et al., 2003)

and Tunisia (Sternberg, 1981; Margaritoulis et al., 2003; SWOT, 2007). Sporadic nesting also has been reported in the western Mediterranean on Corsica (Delaugerre and Cesarini, 2004), southwestern Italy (Bentivegna et al., 2005), and on the Spanish Mediterranean coast (Tomas et al., 2003, 2008). Nesting in the Mediterranean is concentrated between June and early August (Margaritoulis et al., 2003).

Within the Mediterranean, a recent study of mitochondrial and nuclear DNA in nesting assemblages from Greece to Israel indicated genetic structuring, philopatry by both females and males, and limited gene flow between assemblages (Carreras et al., 2007). Genetic differentiation based on mtDNA indicated that there are at least four independent nesting subpopulations within the Mediterranean and usually they are characterized by a single haplotype: (1) Mainland Greece and the adjoining Ionian Islands, (2) eastern Turkey, (3) Israel, and (4) Cyprus. There is no evidence of adult female exchange among these four subpopulations (Carreras et al., 2006). In studies of the foraging grounds in the western and central Mediterranean, seven of the 17 distinct haplotypes detected had not yet been described, indicating that nesting beach data to describe the natal origins of juveniles exploiting the western Mediterranean Sea are incomplete (Carreras et al., 2006; Casale et al., 2008a). Gene flow among the Mediterranean rookeries estimated from nuclear DNA was significantly higher than that calculated from mtDNA, consistent with the scenario of female philopatry maintaining isolation between rookeries, offset by male-mediated gene flow. Nevertheless, the nuclear data show there was a higher degree of substructuring among Mediterranean rookeries compared to those in the Northwest Atlantic (Bowen et al., 2005; Carreras et al., 2007)

d from nuclear DNA was significantly higher than that calculated from mtDNA, consistent with the scenario of female philopatry maintaining isolation between rookeries, offset by male-mediated gene flow. Nevertheless, the nuclear data show there was a higher degree of substructuring among Mediterranean rookeries compared to those in the Northwest Atlantic (Bowen et al., 2005; Carreras et al., 2007).

Small oceanic juveniles from the Mediterranean Sea use the eastern basin (defined as inclusive of the central Mediterranean, Ionian, Adriatic, and Aegean Seas) and the western basin (defined as inclusive of the Tyrrhenian Sea) along the European coast (Laurent et al., 1998; Margaritoulis et al., 2003; Carreras et al., 2006; Revelles et al., 2007). Larger juveniles also use the eastern Atlantic and the eastern Mediterranean, especially the Tunisia-Libya shelf and the Adriatic Sea (Laurent et al., 1993; Margaritoulis et al., 2003; Monzón-Argüllo et al., 2006; Revelles et al., 2007). Adults appear to forage closer to the nesting beaches in the eastern basin; most tag recoveries from females nesting in Greece have occurred in the Adriatic Sea and off Tunisia (Margaritoulis et al., 2003; Lazar et al., 2004).

Loggerheads nesting in the Mediterranean were significantly smaller than loggerheads nesting in the Northwest Atlantic and the South Atlantic. Within the Mediterranean, straight carapace lengths ranged from 58 to 95 cm SCL (Margaritoulis et al., 2003). Greece's loggerheads averaged 77-80 cm SCL (Tiwari and Bjorndal, 2000; Margaritoulis et al., 2003), whereas Turkey's loggerheads averaged 72-73 cm SCL (Margaritoulis et al., 2003). The Greece turtles also produced larger clutches (relative to body size) than those produced by Florida or Brazil nesters (Tiwari and Bjorndal, 2000). The authors suggested that sea turtles in the Mediterranean encounter environmental conditions significantly different from those experienced by populations elsewhere in the Atlantic Ocean basin

ds averaged 72-73 cm SCL (Margaritoulis et al., 2003). The Greece turtles also produced larger clutches (relative to body size) than those produced by Florida or Brazil nesters (Tiwari and Bjorndal, 2000). The authors suggested that sea turtles in the Mediterranean encounter environmental conditions significantly different from those experienced by populations elsewhere in the Atlantic Ocean basin.

Given the information presented above, the BRT concluded, and we concur, that four discrete population et al., 1994; Encalada et al., 1998; Pearce, 2001; Carerras et al., 2007; C. Monzon-Arguello, Instituto Canario de Ciencias Marinas-Spain, personal communication, 2008; Monzon-Arguello et al., 2009). Although loggerheads from the Northwest Atlantic, Northeast Atlantic, and Mediterranean Sea population segments may comingle on oceanic foraging grounds as juveniles, adults are apparently isolated from each other; they also differ demographically. Data from satellite telemetry studies and flipper tag returns have shown that nesting females from the Northwest Atlantic return to the same nesting areas; they reveal no evidence of movement of adults south of the equator or east of 40° W longitude. Similarly, there is no evidence of movement of Northeast Atlantic adults south of the equator, west of 40° W longitude, or east of the Strait of Gibraltar, a narrow strait that connects the Atlantic Ocean to the Mediterranean Sea. Also, there is no evidence of movement of adult Mediterranean Sea loggerheads west of the Strait of Gibraltar. With regard to South Atlantic loggerheads, there have been no observations of tagged loggerheads moving across the equator in the Atlantic, except a single case of a captive-reared animal that was released as a juvenile from Espírito Santo and was recaptured 3 years later in the Azores (Bolten et al., 1990)

vidence of movement of adult Mediterranean Sea loggerheads west of the Strait of Gibraltar. With regard to South Atlantic loggerheads, there have been no observations of tagged loggerheads moving across the equator in the Atlantic, except a single case of a captive-reared animal that was released as a juvenile from Espírito Santo and was recaptured 3 years later in the Azores (Bolten et al., 1990). The separation of the Atlantic Ocean/Mediterranean Sea population segments from population segments within the Indian Ocean and Pacific Ocean basins is believed to be the result of land barriers and oceanographic barriers. Based on mtDNA analysis, Bowen et al. (1994) found a separation of loggerheads in the Atlantic-Mediterranean basins from those in the Indo-Pacific basins since the Pleistocene period. Geography and climate appear to have shaped the evolution of these two matriarchal lineages with the onset of glacial cycles, the appearance of the Panama Isthmus creating a land barrier between the Atlantic and eastern Pacific, and upwelling of cold water off southern Africa creating an oceanographic barrier between the Atlantic and Indian Oceans (Bowen, 2003).

Significance Determination

As stated in the preceding section, the BRT identified nine discrete population segments. As described below by ocean basin, the BRT found that each of the nine discrete population segments is biologically and ecologically significant. They each represent a large portion of the species range, sometimes encompassing an entire hemispheric ocean basin. The range of each discrete population segment represents a unique ecosystem, influenced by local ecological and physical factors. The loss of any individual discrete population segment would result in a significant gap in the loggerhead's range

logically and ecologically significant. They each represent a large portion of the species range, sometimes encompassing an entire hemispheric ocean basin. The range of each discrete population segment represents a unique ecosystem, influenced by local ecological and physical factors. The loss of any individual discrete population segment would result in a significant gap in the loggerhead's range. Each discrete population segment is genetically unique, often identified by unique mtDNA haplotypes, and the BRT indicated that these unique haplotypes could represent adaptive differences; the loss of any one discrete population segment would represent a significant loss of genetic diversity. Therefore, the BRT concluded, and we concur, that these nine population segments are both discrete from other conspecific population segments and significant to the species to which they belong, Caretta caretta.

The geographic delineations given below for each discrete population segment were determined primarily based on nesting beach locations, genetic evidence, oceanographic features, thermal tolerance, fishery bycatch data, and information on loggerhead distribution and migrations from satellite telemetry and flipper tagging studies. With rare exception, adults from discrete population segments remain within the delineated boundaries. In some cases, juvenile turtles from two or more discrete population segments may mix on foraging areas and therefore, their distribution and migrations may extend beyond the geographic boundaries delineated below for each discrete population segment ( e.g., juvenile turtles from the Northwest Atlantic Ocean, Northeast Atlantic Ocean, and Mediterranean Sea discrete population segments share foraging habitat in the western Mediterranean Sea)

more discrete population segments may mix on foraging areas and therefore, their distribution and migrations may extend beyond the geographic boundaries delineated below for each discrete population segment ( e.g., juvenile turtles from the Northwest Atlantic Ocean, Northeast Atlantic Ocean, and Mediterranean Sea discrete population segments share foraging habitat in the western Mediterranean Sea).

Pacific Ocean

The BRT considered 60° N latitude and the equator as the north and south boundaries, respectively, of the North Pacific Ocean population segment based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the North Pacific Ocean discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The North Pacific Ocean population segment encompasses an entire hemispheric ocean basin and its loss would result in a significant gap in the range of the taxon. There is no evidence or reason to believe that female loggerheads from South Pacific nesting beaches would repopulate the North Pacific nesting beaches should those nesting assemblages be lost (Bowen et al., 1994; Bowen, 2003). Tagging studies show that the vast majority of nesting females return to the same nesting area. As summarized by Hatase et al. (2002a), of 2,219 tagged nesting females from Japan, only five females relocated their nesting sites. In addition, flipper tag and satellite telemetry research, as described in detail in the Discreteness Determination section above, has shown no evidence of north-south movement of loggerheads across the equator

rity of nesting females return to the same nesting area. As summarized by Hatase et al. (2002a), of 2,219 tagged nesting females from Japan, only five females relocated their nesting sites. In addition, flipper tag and satellite telemetry research, as described in detail in the Discreteness Determination section above, has shown no evidence of north-south movement of loggerheads across the equator. This discrete population segment is genetically unique ( see Discreteness Determination section above) and the BRT indicated that these unique haplotypes could represent adaptive differences; thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the North Pacific Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

The BRT considered the equator and 60° S latitude as the north and south boundaries, respectively, and 67° W longitude and 139° E longitude as the east and west boundaries, respectively, of the South Pacific Ocean population segment based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the South Pacific Ocean discrete population segment is biologically and ecologically significant because the loss et al., 1994; Bowen, 2003). In addition, flipper tag and satellite telemetry research, as described in detail in the Discreteness Determination section above, has shown no evidence of north-south movement of loggerheads across the equator. The BRT also stated that it does not expect that recolonization from Indian Ocean loggerheads would occur in eastern Australia within ecological time frames

oss et al., 1994; Bowen, 2003). In addition, flipper tag and satellite telemetry research, as described in detail in the Discreteness Determination section above, has shown no evidence of north-south movement of loggerheads across the equator. The BRT also stated that it does not expect that recolonization from Indian Ocean loggerheads would occur in eastern Australia within ecological time frames. Despite evidence of foraging in the Gulf of Carpentaria by adult loggerheads from the nesting populations in eastern Australia (South Pacific Ocean population segment) and western Australia (Southeast Indo-Pacific Ocean population segment), the nesting females from these two regions are considered to be genetically distinct from one another (Limpus, 2009). In addition to a substantial disparity in mtDNA haplotype frequencies between these two populations, FitzSimmons (University of Canberra, unpublished data) found significant differences in nuclear DNA microsatellite loci between females nesting in these two regions, indicating separation between the South Pacific Ocean and the Southeast Indo-Pacific Ocean population segments. Long-term studies show a high degree of site fidelity by adult females in the South Pacific, with most females returning to the same beach within a nesting season and in successive nesting seasons (Limpus, 1985, 2009; Limpus et al., 1994). This has been documented as characteristic of loggerheads from various rookeries throughout the world (Schroeder et al., 2003). This discrete population segment is genetically unique and the BRT indicated that these unique haplotypes could represent adaptive differences. Thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the South Pacific Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy

e unique haplotypes could represent adaptive differences. Thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the South Pacific Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

Indian Ocean

The BRT considered 30° N latitude and the equator as the north and south boundaries, respectively, of the North Indian Ocean population segment based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the North Indian Ocean discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The North Indian Ocean population segment encompasses an entire hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon. Genetic information currently available for Indian Ocean populations indicates that the Oman rookery in the North Indian Ocean and the South African rookery in the Southwest Indian Ocean are genetically distinct, and once sequencing studies are completed for these rookeries, it is likely that they will also be determined to be genetically distinct from the western Australia rookeries in the Southeast Indo-Pacific Ocean (Bowen et al., 1994)

Indian Ocean populations indicates that the Oman rookery in the North Indian Ocean and the South African rookery in the Southwest Indian Ocean are genetically distinct, and once sequencing studies are completed for these rookeries, it is likely that they will also be determined to be genetically distinct from the western Australia rookeries in the Southeast Indo-Pacific Ocean (Bowen et al., 1994). In addition, oceanographic phenomena associated with Indian Ocean equatorial waters exist between the North Indian Ocean population segment and the two population segments in the South Indian Ocean, which likely prevent migration of individuals across the equator on a time scale relative to management and conservation efforts (Conant et al., 2009). Therefore, there is no evidence or reason to believe that female loggerheads from the Southwest Indian Ocean or Southeast Indo-Pacific Ocean would repopulate the North Indian Ocean nesting beaches should those populations be lost (Bowen et al., 1994; Bowen, 2003). Based on this information, the BRT concluded, and we concur, that the North Indian Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

The BRT considered the equator and 60° S latitude as the north and south boundaries, respectively, and 20° E longitude at Cape Agulhas on the southern tip of Africa and 80° E longitude as the east and west boundaries, respectively, of the Southwest Indian Ocean population segment based on oceanographic features, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies

as the north and south boundaries, respectively, and 20° E longitude at Cape Agulhas on the southern tip of Africa and 80° E longitude as the east and west boundaries, respectively, of the Southwest Indian Ocean population segment based on oceanographic features, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the Southwest Indian Ocean discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The Southwest Indian Ocean population segment encompasses half of an hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon. Genetic information currently available for Indian Ocean populations indicates that the Oman rookery in the North Indian Ocean and the South African rookery in the Southwest Indian Ocean are genetically distinct, and once sequencing studies are completed for these rookeries, it is likely that they will also be determined to be genetically distinct from the western Australia rookeries in the Southeast Indo-Pacific Ocean (Bowen et al., 1994). In addition, vicariant barriers ( i.e., oceanographic phenomena associated with Indian Ocean equatorial waters, and the large expanse between continents in the South Indian Ocean without suitable benthic foraging habitat) likely exist between the three Indian Ocean populations that would prevent migration of individuals between populations on a time scale relative to management and conservation efforts (Conant et al., 2009)

iers ( i.e., oceanographic phenomena associated with Indian Ocean equatorial waters, and the large expanse between continents in the South Indian Ocean without suitable benthic foraging habitat) likely exist between the three Indian Ocean populations that would prevent migration of individuals between populations on a time scale relative to management and conservation efforts (Conant et al., 2009). Therefore, there is no evidence or reason to believe that female loggerheads from the North Indian Ocean or Southeast Indo-Pacific Ocean would repopulate the Southwest Indian Ocean nesting beaches should those populations be lost (Bowen et al., 1994; Bowen, 2003). There is also no evidence of movement of adult Southwest Indian Ocean loggerheads west of 20° E longitude at Cape Agulhas, the southernmost point on the African continent, or east of 80° E longitude within the Indian Ocean. Based on this information, the BRT concluded, and we concur, that the Southwest Indian Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

The BRT considered the equator and 60° S latitude as the north and south boundaries, respectively, and 139° E et al., 1994). In addition, vicariant barriers ( i.e., oceanographic phenomena associated with Indian Ocean equatorial waters, and the large expanse between continents in the South Indian Ocean without suitable benthic foraging habitat) likely exist between the three Indian Ocean populations that would prevent migration of individuals between populations on a time scale relative to management and conservation efforts (Conant et al., 2009). Therefore, there is no evidence or reason to believe that female loggerheads from the North Indian Ocean or Southwest Indian Ocean would repopulate the Southeast Indo-Pacific Ocean nesting beaches should those populations be lost (Bowen et al., 1994; Bowen, 2003)

vent migration of individuals between populations on a time scale relative to management and conservation efforts (Conant et al., 2009). Therefore, there is no evidence or reason to believe that female loggerheads from the North Indian Ocean or Southwest Indian Ocean would repopulate the Southeast Indo-Pacific Ocean nesting beaches should those populations be lost (Bowen et al., 1994; Bowen, 2003). There is also no evidence of movement of adult Southeast Indo-Pacific Ocean loggerheads west of 80° E longitude within the Indian Ocean. Despite evidence of foraging in the Gulf of Carpentaria by adult loggerheads from the nesting populations in eastern Australia (South Pacific Ocean population segment) and western Australia (Southeast Indo-Pacific Ocean population segment), the nesting females from these two regions are considered to be genetically distinct from one another (Limpus, 2009). In addition to a substantial disparity in mtDNA haplotype frequencies between these two regions, FitzSimmons (University of Canberra, unpublished data) found significant differences in nuclear DNA microsatellite loci from females nesting in these two regions, indicating separation between the South Pacific Ocean population segment and the Southeast Indo-Pacific Ocean population segment. Based on this information, the BRT concluded, and we concur, that the Southeast Indo-Pacific Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

Atlantic Ocean and Mediterranean Sea

diterranean Sea

The BRT considered 60° N latitude and the equator as the north and south boundaries, respectively, and 40° W longitude as the east boundary of the Northwest Atlantic Ocean population segment based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the Northwest Atlantic Ocean discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The Northwest Atlantic Ocean population segment encompasses half of an hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon. Genetic studies have shown that adult populations are highly structured with no overlap in distribution among adult loggerheads from the Northwest Atlantic, Northeast Atlantic, South Atlantic, and Mediterranean Sea (Bowen et al., 1994; Encalada et al., 1998; Pearce, 2001; Carerras et al., 2007; C. Monzon-Arguello, Instituto Canario de Ciencias Marinas—Spain, personal communication, 2008; Monzon-Arguello et al., 2009). There is no evidence or reason to believe that female loggerheads from the Northeast Atlantic, Mediterranean Sea, or South Atlantic nesting beaches would repopulate the Northwest Atlantic nesting beaches should these populations be lost (Bowen et al., 1994; Bowen, 2003). Data from satellite telemetry studies and flipper tag returns, as described in detail in the Discreteness Determination section above, have shown that the vast majority of nesting females from the Northwest Atlantic return to the same nesting area; they reveal no evidence of movement of adults south of the equator or east of 40° W longitude

se populations be lost (Bowen et al., 1994; Bowen, 2003). Data from satellite telemetry studies and flipper tag returns, as described in detail in the Discreteness Determination section above, have shown that the vast majority of nesting females from the Northwest Atlantic return to the same nesting area; they reveal no evidence of movement of adults south of the equator or east of 40° W longitude. This discrete population segment is genetically unique ( see Discreteness Determination section above) and the BRT indicated that these unique haplotypes could represent adaptive differences; thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the Northwest Atlantic Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

The BRT considered 60° N latitude and the equator as the north and south boundaries, respectively, and 40° W longitude as the west boundary of the Northeast Atlantic Ocean population segment. The BRT considered the boundary between the Northeast Atlantic Ocean and Mediterranean Sea population segments as 5°36′ W longitude (Strait of Gibraltar). These boundaries are based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the Northeast Atlantic Ocean discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The Northeast Atlantic Ocean population segment encompasses half of an hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon

cant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The Northeast Atlantic Ocean population segment encompasses half of an hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon. Genetic studies have shown that adult populations are highly structured with no overlap in distribution among adult loggerheads from the Northwest Atlantic, Northeast Atlantic, South Atlantic, and Mediterranean Sea (Bowen et al., 1994; Encalada et al., 1998; Pearce, 2001; Carerras et al., 2007; C. Monzon-Arguello, Instituto Canario de Ciencias Marinas—Spain, personal communication, 2008; Monzon-Arguello et al., 2009). There is no evidence or reason to believe that female loggerheads from the Northwest Atlantic, Mediterranean Sea, or South Atlantic nesting beaches would repopulate the Northeast Atlantic nesting beaches should these populations be lost (Bowen et al., 1994; Bowen, 2003). There is also no evidence see Discreteness Determination section above) and the BRT indicated that these unique haplotypes could represent adaptive differences; thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the Northeast Atlantic Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

it satisfies the significance element of the DPS policy.

The BRT considered the Mediterranean Sea west to 5°36′ W longitude (Strait of Gibraltar) as the boundary of the Mediterranean Sea population segment based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the Mediterranean Sea discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The Mediterranean Sea population segment encompasses the entire Mediterranean Sea basin, and its loss would result in a significant gap in the range of the taxon. Genetic studies have shown that adult populations are highly structured with no overlap in distribution among adult loggerheads from the Northwest Atlantic, Northeast Atlantic, South Atlantic, and Mediterranean Sea (Bowen et al., 1994; Encalada et al., 1998; Pearce, 2001; Carerras et al., 2007; C. Monzon-Arguello, Instituto Canario de Ciencias Marinas—Spain, personal communication, 2008; Monzon-Arguello et al., 2009). There is no evidence or reason to believe that female loggerheads from the Northwest Atlantic, Northeast Atlantic, or South Atlantic nesting beaches would repopulate the Mediterranean Sea nesting beaches should these populations be lost (Bowen et al., 1994; Bowen, 2003). As previously described, adults from the Mediterranean Sea population segment appear to forage closer to the nesting beaches in the eastern basin, and most flipper tag recoveries from females nesting in Greece have occurred in the Adriatic Sea and off Tunisia (Margaritoulis et al., 2003; Lazar et al., 2004). There is no evidence of movement of adult Mediterranean Sea loggerheads west of the Strait of Gibraltar (5°36' W longitude)

rom the Mediterranean Sea population segment appear to forage closer to the nesting beaches in the eastern basin, and most flipper tag recoveries from females nesting in Greece have occurred in the Adriatic Sea and off Tunisia (Margaritoulis et al., 2003; Lazar et al., 2004). There is no evidence of movement of adult Mediterranean Sea loggerheads west of the Strait of Gibraltar (5°36' W longitude). This discrete population segment is genetically unique ( see Discreteness Determination section above) and the BRT indicated that these unique haplotypes could represent adaptive differences; thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the Mediterranean Sea population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

The BRT considered the equator and 60° S latitude as the north and south boundaries, respectively, and 20° E longitude at Cape Agulhas on the southern tip of Africa and 67° W longitude as the east and west boundaries, respectively, of the South Atlantic Ocean population segment based on oceanographic features, loggerhead sightings, thermal tolerance, fishery bycatch data, and information on loggerhead distribution from satellite telemetry and flipper tagging studies. The BRT determined that the South Atlantic Ocean discrete population segment is biologically and ecologically significant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The South Atlantic Ocean population segment encompasses an entire hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon

gnificant because the loss of this population segment would result in a significant gap in the range of the taxon, and the population segment differs markedly from other population segments of the species in its genetic characteristics. The South Atlantic Ocean population segment encompasses an entire hemispheric ocean basin, and its loss would result in a significant gap in the range of the taxon. Genetic studies have shown that adult populations are highly structured with no overlap in distribution among adult loggerheads from the Northwest Atlantic, Northeast Atlantic, South Atlantic, and Mediterranean Sea (Bowen et al., 1994; Encalada et al., 1998; Pearce, 2001; Carerras et al., 2007; C. Monzon-Arguello, Instituto Canario de Ciencias Marinas-Spain, personal communication, 2008; Monzon-Arguello et al., 2009). There is no evidence or reason to believe that female loggerheads from the Northwest Atlantic, Northeast Atlantic, or Mediterranean Sea nesting beaches would repopulate the South Atlantic nesting beaches should these populations be lost (Bowen et al., 1994; Bowen, 2003). This discrete population segment is genetically unique ( see Discreteness Determination section above) and the BRT indicated that these unique haplotypes could represent adaptive differences; thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the South Atlantic Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy

unique haplotypes could represent adaptive differences; thus, the loss of this discrete population segment would represent a significant loss of genetic diversity. Based on this information, the BRT concluded, and we concur, that the South Atlantic Ocean population segment is significant to the taxon to which it belongs, and, therefore, that it satisfies the significance element of the DPS policy.

In summary, based on the information provided in the Discreteness Determination and Significance Determination sections above, the BRT identified nine loggerhead DPSs distributed globally: (1) North Pacific Ocean DPS, (2) South Pacific Ocean DPS, (3) North Indian Ocean DPS, (4) Southeast Indo-Pacific Ocean DPS, (5) Southwest Indian Ocean DPS, (6) Northwest Atlantic Ocean DPS, (7) Northeast Atlantic Ocean DPS, (8) Mediterranean Sea DPS, and (9) South Atlantic Ocean DPS. We concur with the findings and application of the DPS policy described by the BRT and conclude that the nine DPSs identified by the BRT warrant delineation as DPSs ( i.e., they are discrete and significant).

Significant Portion of the Range

We have determined that the range of each DPS contributes meaningfully to the conservation of the DPS and that populations that may contribute more or less to the conservation of each DPS throughout a portion of its range cannot be identified due to the highly migratory nature of the listed entity.

The loggerhead sea turtle is highly migratory and crosses multiple domestic and international geopolitical boundaries. Depending on the life stage, they may occur in oceanic waters or along the continental shelf of landmasses, or transit back and forth between oceanic and neritic habitats. Protection and management of both the terrestrial and marine environments is essential to recovering the listed entity. Management measures implemented by any State, foreign nation, or political subdivision likely would only affect individual sea turtles during certain stages and seasons of the life cycle

tinental shelf of landmasses, or transit back and forth between oceanic and neritic habitats. Protection and management of both the terrestrial and marine environments is essential to recovering the listed entity. Management measures implemented by any State, foreign nation, or political subdivision likely would only affect individual sea turtles during certain stages and seasons of the life cycle. Management measures implemented by any State, foreign nation, or political subdivision may also affect individuals from multiple DPSs because juvenile turtles from disparate DPSs can overlap on foraging grounds or migratory corridors ( e.g., Northwest Atlantic, Northeast Atlantic, and Mediterranean Sea DPSs). The “significant” term in “significant portion of the range” refers to the contribution of the population(s)

Status of the Nine Loggerhead DPSs

Abundance estimates across all life stages do not exist for the nine DPSs. Within the global range of the species, and within each DPS, the primary data available are collected on nesting beaches, either as counts of nests or counts of nesting females, or a combination of both (either direct or extrapolated). Information on abundance and trends away from the nesting beaches is limited or non-existent, primarily because these data are, relative to nesting beach studies, logistically difficult and expensive to obtain. Therefore, the primary information source for directly evaluating status and trends of the nine DPSs is nesting beach data.

North Pacific Ocean DPS

In the North Pacific, loggerhead nesting is essentially restricted to Japan where monitoring of loggerhead nesting began in the 1950s on some beaches, and expanded to include most known nesting beaches since approximately 1990. Kamezaki et al. (2003) reviewed census data collected from most of the Japanese nesting beaches. Although most surveys were initiated in the 1980s and 1990s, some data collection efforts were initiated in the 1950s

s essentially restricted to Japan where monitoring of loggerhead nesting began in the 1950s on some beaches, and expanded to include most known nesting beaches since approximately 1990. Kamezaki et al. (2003) reviewed census data collected from most of the Japanese nesting beaches. Although most surveys were initiated in the 1980s and 1990s, some data collection efforts were initiated in the 1950s. Along the Japanese coast, nine major nesting beaches (greater than 100 nests per season) and six “submajor” beaches (10-100 nests per season) were identified. Census data from 12 of these 15 beaches provide composite information on longer-term trends in the Japanese nesting assemblage. Using information collected on these beaches, Kamezaki et al. (2003) concluded a substantial decline (50-90 percent) in the size of the annual loggerhead nesting population in Japan in recent decades. Snover (2008) combined nesting data from the Sea Turtle Association of Japan and data from Kamezaki et al. (2002) to provide a recent 18-year time series of nesting data from 1990-2007. Nesting declined from an initial peak of approximately 6,638 nests in 1990-1991, followed by a steep decline to a low of 2,064 nests in 1997. During the past decade, nesting increased gradually to 5,167 nests in 2005, declined and then rose again to a high of just under 11,000 nests in 2008. Estimated nest numbers for 2009 are on the order of 7,000-8,000 nests. While nesting numbers have gradually increased in recent years and the number for 2009 is similar to the start of the time series in 1990, historical evidence indicates that there has been a substantial decline over the last half of the 20th century.

South Pacific Ocean DPS

the time series in 1990, historical evidence indicates that there has been a substantial decline over the last half of the 20th century.

South Pacific Ocean DPS

In the South Pacific, loggerhead nesting is almost entirely restricted to eastern Australia (primarily Queensland) and New Caledonia, with the majority of nesting occurring in eastern Australia, a population that has been well studied. The size of the annual breeding population (females only) has been monitored at numerous rookeries in Australia since 1968 (Limpus and Limpus, 2003), and these data constitute the primary measure of the current status of the DPS. The total nesting population for Queensland was approximately 3,500 females in the 1976-1977 nesting season (Limpus, 1985; Limpus and Reimer, 1994). Little more than two decades later, Limpus and Limpus (2003) estimated this nesting population at less than 500 females in the 1999-2000 nesting season. There has been a marked decline in the number of females breeding annually since the mid-1970s, with an estimated 50 to 80 percent decline in the number of breeding females at various Australian rookeries up to 1990 (Limpus and Reimer, 1994) and a decline of approximately 86 percent by 1999 (Limpus and Limpus, 2003). Comparable nesting surveys have not been conducted in New Caledonia however. Information from pilot surveys conducted in 2005, combined with oral history information collected, suggest that there has been a decline in loggerhead nesting (Limpus et al., 2006). Based on data from the pilot study, only 60 to 70 loggerheads nested on the four surveyed New Caledonia beaches during the 2004-2005 nesting season (Limpus et al., 2006).

004-2005 nesting season (Limpus et al., 2006).

Studies of eastern Australia loggerheads at their foraging areas provide some information on the status of non-breeding loggerheads of the South Pacific Ocean DPS. Chaloupka and Limpus (2001) determined that the resident loggerhead population on coral reefs of the southern Great Barrier Reef declined at 3 percent per year from 1985 to the late 1990s. The observed decline was hypothesized as a result of recruitment failure, given few anthropogenic impacts and constant high annual survivorship measured at this foraging habitat (Chaloupka and Limpus, 2001). Concurrently, a decline in new recruits was measured in these foraging areas (Limpus and Limpus, 2003).

North Indian Ocean DPS

The North Indian Ocean hosts the largest nesting assemblage of loggerheads in the eastern hemisphere; the vast majority of these loggerheads nest in Oman (Baldwin et al., 2003). Nesting occurs in greatest density on Masirah Island; the number of emergences ranges from 27-102 per km nightly (Ross, 1998). Nesting densities have complicated the implementation of standardized nesting beach surveys, and more precise nesting data have only been collected since 2008. Extrapolations resulting from partial surveys and tagging in 1977-1978 provided broad estimates of 19,000-60,000 females nesting annually at Masirah Island, while a more recent partial survey in 1991 provides an estimate of 23,000 nesting females at Masirah Island (Baldwin, 1992; Ross, 1979, 1998; Ross and Barwani 1982). A reinterpretation of these estimates, assuming 50 percent nesting success (as compared to 100 percent in the original estimates), resulted in an estimate of 20,000 to 40,000 females nesting annually (Baldwin et al., 2003). Reliable trends in nesting cannot be determined due to the lack of standardized surveys at Masirah Island prior to 2008. In 2008, about 50,000 nests were estimated based on daily surveys of the highest density nesting beaches and weekly surveys on all remaining island nesting beaches

e original estimates), resulted in an estimate of 20,000 to 40,000 females nesting annually (Baldwin et al., 2003). Reliable trends in nesting cannot be determined due to the lack of standardized surveys at Masirah Island prior to 2008. In 2008, about 50,000 nests were estimated based on daily surveys of the highest density nesting beaches and weekly surveys on all remaining island nesting beaches. Even using the low end of the 1977-1978 estimates of 20,000 nesting females at Masirah, this suggests a significant decline in the size of the nesting population and is consistent with observations by local rangers that the population has declined dramatically in the last three decades (E. Possardt, FWS, personal communication, 2008). If the higher estimates are accurate then the decline would be greater than 70 percent.

In addition to the nesting beaches on Masirah Island, over 3,000 nests per year have been recorded in Oman on the Al-Halaniyat Islands and, along the Oman mainland of the Arabian Sea, approximately 2,000 nests are deposited annually (Salm, 1991; Salm et al., 1993). In Yemen, on Socotra Island, 50-100 loggerheads were estimated to have nested in 1999 (Pilcher and Saad, 2000). A time series of nesting data based on standardized surveys is not available to determine trends for these nesting sites.

Loggerhead nesting is rare elsewhere in the northern Indian Ocean and in some cases is complicated by inaccurate species identification (Shanker, 2004; Tripathy, 2005). A small number of nesting females use the beaches of Sri Lanka every year; however, there are no records that Sri Lanka has ever been a major nesting area for loggerheads (Kapurusinghe, 2006). Loggerheads have been reported nesting in low numbers in Myanmar; however, these data may not be reliable because of misidentification of species (Thorbjarnarson et al., 2000)

Shanker, 2004; Tripathy, 2005). A small number of nesting females use the beaches of Sri Lanka every year; however, there are no records that Sri Lanka has ever been a major nesting area for loggerheads (Kapurusinghe, 2006). Loggerheads have been reported nesting in low numbers in Myanmar; however, these data may not be reliable because of misidentification of species (Thorbjarnarson et al., 2000).

Southeast-Indo Pacific Ocean DPS

In the eastern Indian Ocean, loggerhead nesting is restricted to western Australia (Dodd, 1988), and this nesting population is the largest in Australia (Wirsing et al., unpublished data, cited in Natural Heritage Trust, 2005). Dirk Hartog Island hosts about 70-75 percent of nesting individuals in the eastern Indian Ocean (Baldwin et al., 2003). Surveys have been conducted on the island for the duration of six nesting seasons between 1993/1994 and 1999/2000 (Baldwin et al., 2003). An estimated 800-1,500 loggerheads nest annually on Dirk Hartog Island beaches (Baldwin et al., 2003).

Fewer loggerheads (approximately 150-350 per season) are reported nesting on the Muiron Islands; however, more nesting loggerheads are reported here than on North West Cape (approximately 50-150 per season) (Baldwin et al., 2003). Although data are insufficient to determine trends, evidence suggests the nesting population in the Muiron Islands and North West Cape region was depleted before recent beach monitoring programs began (Nishemura and Nakahigashi, 1990; Poiner et al., 1990; Poiner and Harris, 1996).

Southwest Indian Ocean DPS

and North West Cape region was depleted before recent beach monitoring programs began (Nishemura and Nakahigashi, 1990; Poiner et al., 1990; Poiner and Harris, 1996).

Southwest Indian Ocean DPS

In the Southwest Indian Ocean, the highest concentration of nesting occurs on the coast of Tongaland, South Africa, where surveys and management practices were instituted in 1963 (Baldwin et al., 2003). A trend analysis of index nesting beach data from this region from 1965 to 2008 indicates an increasing nesting population between the first decade of surveys, which documented 500-800 nests annually, and the last 8 years, which documented 1,100-1,500 nests annually (Nel, 2008). These data represent approximately 50 percent of all nesting within South Africa and are believed to be representative of trends in the region. Loggerhead nesting occurs elsewhere in South Africa, but sampling is not consistent and no trend data are available. The total number of females nesting annually in South Africa is estimated between 500-2,000 (Baldwin et al., 2003). In Mozambique, surveys have been instituted much more recently; likely less than 100 females nest annually and no trend data are available (Baldwin et al., 2003). Similarly, in Madagascar, loggerheads have been documented nesting in low numbers, but no trend data are available (Rakotonirina, 2001).

Northwest Atlantic Ocean DPS

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 (Sternberg, 1981; Ehrhart, 1989; Ehrhart et al., 2003; NMFS and FWS, 2008). Collectively, the Northwest 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. NMFS and FWS (2008), Witherington et al

is concentrated in the southeastern U.S. and on the Yucatan Peninsula in Mexico (Sternberg, 1981; Ehrhart, 1989; Ehrhart et al., 2003; NMFS and FWS, 2008). Collectively, the Northwest 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. NMFS and FWS (2008), Witherington et al. (2009), and TEWG (2009) provide comprehensive analyses of the status of the nesting assemblages within the Northwest Atlantic Ocean DPS using standardized data collected over survey periods ranging from 10 to 23 years. The results of these analyses, using different analytical approaches, were consistent in their findings—there has been a significant, overall nesting decline within this DPS.

NMFS and FWS (2008) identified five recovery units (nesting subpopulations) in the Northwest Atlantic Ocean: the Northern U.S. (Florida/Georgia border to southern Virginia); Peninsular Florida (Florida/Georgia border south through Pinellas County, 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 Greater Caribbean (Mexico through French Guiana, The Bahamas, Lesser and Greater Antilles). Declining trends in the annual number of nests were documented for all recovery units for which there were adequate data. The most significant declining trend has been documented for the Peninsular Florida Recovery Unit, where nesting declined 26 percent over the 20-year period from 1989-2008, and declined 41 percent over the period 1998-2008 (NMFS and FWS, 2008; Witherington et al., 2009). The most standardized nest count from this recovery unit in 2009 recorded the fourth lowest loggerhead nesting in the 21-year monitoring period, reinforcing the assessment of nesting decline (B. Witherington, FWC, personal communication, 2010)

clined 26 percent over the 20-year period from 1989-2008, and declined 41 percent over the period 1998-2008 (NMFS and FWS, 2008; Witherington et al., 2009). The most standardized nest count from this recovery unit in 2009 recorded the fourth lowest loggerhead nesting in the 21-year monitoring period, reinforcing the assessment of nesting decline (B. Witherington, FWC, personal communication, 2010). The Peninsular Florida Recovery Unit represents approximately 87 percent of all nesting effort in the Northwest Atlantic Ocean DPS (Ehrhart et al., 2003). The Northern U.S. Recovery Unit is the second largest recovery unit within the DPS and is declining significantly at 1.3 percent annually since 1983 (NMFS and FWS, 2008). The Greater Caribbean Recovery Unit is the third largest recovery unit within the Northwest Atlantic Ocean DPS, with the majority of nesting at Quintana Roo, Mexico. TEWG (2009) reported a greater than 5 percent annual decline in loggerhead nesting from 1995-2006 at Quintana Roo.

In an effort to evaluate loggerhead population status and trends beyond the nesting beach, NMFS and FWS (2008) and TEWG (2009) reviewed data from in-water studies within the range of the Northwest Atlantic Ocean DPS. NMFS and FWS (2008), in the Recovery Plan for the Northwest Atlantic Population of the Loggerhead Sea Turtle, summarized population trend data reported from nine in-water study sites, located between Long Island Sound, New York, and Florida Bay, Florida, where loggerheads were regularly captured and where efforts were made to provide local indices of abundance. The study periods for these nine sites varied. The earliest began in 1987, and the most recent were initiated in 2000. None included annual sampling. Results reported from four of the studies indicated no discernible trend, two studies reported declining trends, and two studies reported increasing trends

ads were regularly captured and where efforts were made to provide local indices of abundance. The study periods for these nine sites varied. The earliest began in 1987, and the most recent were initiated in 2000. None included annual sampling. Results reported from four of the studies indicated no discernible trend, two studies reported declining trends, and two studies reported increasing trends. Trends at one study site, Mosquito Lagoon, Florida, indicated either no trend (all data) or a declining trend (more recent data), depending on whether all sample years were used or only the more recent, and likely more comparable sample years, were used. TEWG (2009) used raw data from six of the aforementioned nine in-water study sites to conduct trend analyses. Results from three of the four sites located in the southeast U.S. showed an increasing trend in the abundance of loggerheads, one showed no discernible trend, and the two sites located in the northeast U.S. showed a decreasing trend in abundance of loggerheads. Both NMFS and FWS (2008) and TEWG (2009) stress that population trend results currently available from in-water studies must be viewed with caution given the limited number of sampling sites, size of sampling areas, biases in sampling, and caveats associated with the analyses.

Northeast Atlantic Ocean DPS

In the northeastern Atlantic, the Cape Verde Islands support the only large nesting population of loggerheads in the region (Fretey, 2001). Nesting occurs at some level on most of the islands in the archipelago with the largest nesting numbers reported from the island of Boa Vista where studies have been ongoing since 1998 (Lazar and Holcer, 1998; Lopez-Jurado et al. , 2000; Fretey, 2001; Varo Cruz et al. , 2007; Loureiro, 2008; M. Tiwari, NMFS, personal communication, 2008). On Boa Vista Island, 833 and 1,917 nests were reported in 2001 and 2002 respectively from 3.1 km of beach (Varo Cruz et al. , 2007) and between 1998 and 2002 the local project had tagged 2,856 females (Varo Cruz et al

ve been ongoing since 1998 (Lazar and Holcer, 1998; Lopez-Jurado et al. , 2000; Fretey, 2001; Varo Cruz et al. , 2007; Loureiro, 2008; M. Tiwari, NMFS, personal communication, 2008). On Boa Vista Island, 833 and 1,917 nests were reported in 2001 and 2002 respectively from 3.1 km of beach (Varo Cruz et al. , 2007) and between 1998 and 2002 the local project had tagged 2,856 females (Varo Cruz et al. , 2007). More recently, in 2005, 5,396 nests and 3,121 females were reported from 9 km of beach on Boa Vista Island (Lopez-Jurado et al. , 2007). From Santiago Island, 66 nests were reported from four beaches in 2007 and 53 nests from five beaches in 2008 ( http://tartarugascaboverde.wordpress.com/santiago ). Due to limited data available, a population trend cannot currently be determined for the Cape Verde population; however, available information on the directed killing of nesting females suggests that this nesting population is under severe pressure and likely significantly reduced from historic levels. Loureiro (2008) reported a reduction in nesting from historic levels at Santiago Island, based on interviews with elders. Elsewhere in the northeastern Atlantic, loggerhead nesting is non-existent or occurs at very low levels. In Morocco, anecdotal reports indicated high numbers of nesting turtles in southern Morocco (Pasteur and Bons, 1960), but a few recent surveys of the Atlantic coastline have suggested a dramatic decline (Tiwari et al. , 2001, 2006). A few nests have been reported from Mauritania (Arvy et al. , 2000) and Sierra Leone (E. Aruna, Conservation Society of Sierra Leone, personal communication, 2008). Some loggerhead nesting in Senegal and elsewhere along the coast of West Africa has been reported; however, a more recent and reliable confirmation is needed (Fretey, 2001).

Mediterranean Sea DPS

Nesting occurs throughout the central and eastern Mediterranean in Italy, Greece, Cyprus, Turkey, Syria, Lebanon, Israel, the Sinai, Egypt, Libya, and Tunisia (Sternberg, 1981; Margaritoulis et al

8). Some loggerhead nesting in Senegal and elsewhere along the coast of West Africa has been reported; however, a more recent and reliable confirmation is needed (Fretey, 2001).

Mediterranean Sea DPS

Nesting occurs throughout the central and eastern Mediterranean in Italy, Greece, Cyprus, Turkey, Syria, Lebanon, Israel, the Sinai, Egypt, Libya, and Tunisia (Sternberg, 1981; Margaritoulis et al. , 2003; SWOT, 2007). In addition, sporadic nesting has been reported from the western Mediterranean, but the vast majority of nesting (greater than 80 percent) occurs in Greece and Turkey (Margaritoulis et al. , 2003). The documented annual nesting of loggerheads in the Mediterranean averages about 5,000 nests (Margaritoulis et al. , 2003). There is no discernible trend in nesting at the two longest monitoring projects in Greece, Laganas Bay (Margaritoulis, 2005) and southern Kyparissia Bay (Margaritoulis and Rees, 2001). However, the nesting trend at Rethymno Beach, which hosts approximately 7 percent of all documented loggerhead nesting in the Mediterranean, shows a highly significant declining trend (1990-2004) (Margaritoulis et al. , 2009). In Turkey, intermittent nesting surveys have been conducted since the 1970s with more consistent surveys conducted on some beaches only since the 1990s, making it difficult to assess trends in nesting. Ilgaz et al. (2007) reported a declining trend at Fethiye Beach from 1993-2004, this beach represents approximately 10 percent of loggerhead nesting in Turkey (Margaritoulis et al. , 2003).

South Atlantic Ocean DPS

outh Atlantic Ocean DPS

In the South Atlantic nesting occurs primarily along the mainland coast of Brazil from Sergipe south to Rio de Janeiro, with peak concentrations in northern Bahia, Espírito Santo, and northern Rio de Janeiro with peak nesting along the coast of Bahia (Marcovaldi and Chaloupka, 2007). Prior to 1980, loggerhead nesting populations in Brazil were considered severely depleted. Recently, Marcovaldi and Chaloupka (2007) reported a long-term, sustained increasing trend in nesting abundance over a 16-year period from 1988 through 2003 on 22 surveyed beaches containing more than 75 percent of all loggerhead nesting in Brazil. A total of 4,837 nests were reported from these survey beaches for the 2003-2004 nesting season (Marcovaldi and Chaloupka, 2007).

Summary of Factors Affecting the Nine Loggerhead DPSs

Section 4 of the ESA (16 U.S.C. 1533) and implementing regulations at 50 CFR part 424 set forth procedures for adding species to the Federal List of Endangered and Threatened Species. Under section 4(a) of the Act, we must determine if a species is threatened or endangered because of any of the following five factors: (A) The present or threatened destruction, modification, or curtailment of its habitat or range; (B) overutilization for commercial, recreational, scientific, or educational purposes; (C) disease or predation; (D) the inadequacy of existing regulatory mechanisms; or (E) other natural or manmade factors affecting its continued existence.

We have described the effects of various factors leading to the decline of the loggerhead sea turtle in the original listing determination (43 FR 32800; July 28, 1978) and other documents (NMFS and USFWS, 1998, 2007, 2008). In making this finding, information regarding the status of each of the nine loggerhead DPSs is considered in relation to the five factors provided in section 4(a)(1) of the ESA

e have described the effects of various factors leading to the decline of the loggerhead sea turtle in the original listing determination (43 FR 32800; July 28, 1978) and other documents (NMFS and USFWS, 1998, 2007, 2008). In making this finding, information regarding the status of each of the nine loggerhead DPSs is considered in relation to the five factors provided in section 4(a)(1) of the ESA. The reader is directed to section 5 of the Status Review for a more detailed discussion of the factors affecting the nine identified loggerhead DPSs. In section 5.1., a general description of the threats that occur for all DPSs is presented under the relevant section 4(a)(1) factor. In section 5.2, threats that are specific to a particular DPS are presented by DPS under each section 4(a)(1) factor. That information is incorporated here by reference; the following is a summary of that information by DPS.

North Pacific Ocean DPS

A. The Present or Threatened Destruction, Modification, or Curtailment of its Habitat or Range

Terrestrial Zone

Destruction and modification of loggerhead nesting habitat in the North Pacific result from coastal development and construction, placement of erosion control structures and other barriers to nesting, beachfront lighting, vehicular and pedestrian traffic, sand extraction, beach erosion, beach sand placement, beach pollution, removal of native vegetation, and planting of non-native vegetation (NMFS and USFWS, 1998). Beaches in Japan where loggerheads nest are extensively eroded due to dredging and dams constructed upstream, and are obstructed by seawalls as well. Unfortunately, no quantitative studies have been conducted to determine the impact to the loggerhead nesting populations (Kamezaki et al. , 2003). However, it is clear that loggerhead nesting habitat has been impacted by erosion and extensive beach use by tourists, both of which have contributed to unusually high mortality of eggs and pre-emergent hatchlings at many Japanese rookeries (Matsuzawa, 2006)

tunately, no quantitative studies have been conducted to determine the impact to the loggerhead nesting

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