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The loggerhead sea turtle
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Subclass: Anapsida
Order: Testudines
Family: Cheloniidae
Genus: Caretta
Rafinesque, 1814
Species: C. caretta
Binomial name
Caretta caretta
Linnaeus, 1758
Range of Caretta caretta

The loggerhead sea turtle (Caretta caretta) is a sea turtle, the only member of the genus Caretta. It is the world's most-studied and best-known sea-turtle.[1] The loggerhead sea turtle also holds the title of the world's largest hard-shelled turtle.[2] The genus name "Caretta" is a latinization of the French "caret", meaning turtle, tortoise, or sea turtle.[3] Adult loggerhead sea turtles weigh from 170 pounds (77 kg) to 350 pounds (159 kg) and measure 31 inches (79 cm) to 45 inches (114 cm).[3] Their shell is a reddish brown color, and their skin is yellow/brown. They are named for their disproportionately large head. They are the state reptile of South Carolina.[4]

Contents

Taxonomy

Two subspecies are recognized: C. caretta gigas, inhabits the Indian and Pacific Oceans, and C. caretta caretta, the Atlantic loggerhead, also found in south Italy and the Greek islands of Zakynthos, Kefalonia, Crete, and the Peloponnese and in Dalyan in southwestern Turkey. (see June Haimoff). The loggerhead is now being added to the endangered list.

C. caretta gets its name from its large head that supports the jaws necessary for crushing exoskeletons/shells of prey such as conch.[5] Linnaeus first named the loggerhead, calling it Testudo caretta. Thirty-five other names emerged over the following two centuries, but Caretta caretta has prevailed.[6]

Range and distribution

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Atlantic Ocean

The greatest concentration of loggerheads is along the southeastern coast of North America and in the Gulf of Mexico.[1] Very few loggerheads are found along the European and African coastlines.[1] Florida is the most popular nesting site with over 67,000 nests per year. Nesting extends as far north as Virginia, as far south as Brazil, and as far east as the Cape Verde Islands.[1] The Cape Verde Islands are the only significant nesting site on the eastern side of the Atlantic.

Indian Ocean

Indian Ocean loggerheads feed along the coastlines of Africa, the Arabian Peninsula, and in the Arabian Sea.[1] In Africa, the loggerhead nests from Mozambique's Bazaruto Archipelago south to South Africa's St Lucia estuary.[6] The largest population is located near the coast of western Australia.[1] The largest nesting site is in Oman, on the Arabian Peninsula, and hosts around 15,000 nests annually.[1] Western Australia and Southern Africa are other notable nesting areas.

Pacific Ocean

Loggerheads live in temperate to tropical regions of the Pacific Ocean.[6] Pacific loggerheads forage in the East China Sea, southwestern Pacific, and along the Baja Peninsula.[1] Eastern Australia and Japan are the major nesting areas.[1] Little nesting occurs on the eastern Pacific coastline however thousands of loggerheads live there during the summers.[1] There are also few breeding grounds in the northern and southern Pacific Ocean. In fact, loggerheads have been found to nest in Japan in the northern Pacific Ocean. Eastern Australia, New Caledonia, and rarely Vanuatu and Tokelau. Yakushima Island, Miyazaki, Minabe, and Atsumi beaches on the mainland provide important nesting areas. Yakushima Island is the most important of these, with three nesting grounds that 40% of all loggerheads near Japan visit. Post-nesting females often find homes in the East China Sea, while the Kuroshio Extension Bifurcation Region and the Coast of Baja California Sur, Mexico provide important juvenile foraging areas.[6] Hatchlings migrate from the east Pacific to Mexico only to return later as sub-adults.[1]

Mediterranean Sea

The Mediterranean is a nursery for juvenile loggerheads. Almost 45 percent of the juvenile population in the Mediterranean has migrated from the Atlantic.[1] Loggerheads feed in the western and eastern basins of the Mediterranean, Alboran Sea and Adriatic Sea areas respectively.[1]

Greece is the most popular nesting site with more than 3,000 nests per year.[1] The coastlines of Cyprus and Turkey are also common nesting sites.[1]

Anatomy and Morphology

Average adult loggerhead sea turtles have a weight range from 170 pounds (77 kg) to 350 pounds (159 kg).[3] On average, they measure 31 inches (79 cm) to 45 inches (114 cm).[3] The adult has a carapace (upper side of the shell) length of approximately 36 inches (91 cm).[5] The flippers and carapace range from a dull brown to a reddish-brown, and the plastron (underside) is typically pale yellow.[5] The turtle's neck and sides are brown on the tops and yellow on the sides and bottom.[5] Hatchlings vary in color from gray to brown, and lack the adults' distinct yellows and reds. They measure approximately 1.8 inches (4.6 cm) at birth, and weigh about .04 pounds (18 g).[5]

The heart-shaped carapace is thick and bony and covered by non-overlapping scutes at the seam lines. Typically, there are 11 or 12 pairs of marginal scutes, 5 pairs of both vertebral and costal scutes, and a nuchal scute that contacts the costals. The carapace is connected to the plastron by 3 pairs of inframarginal scutes that lack pores. The plastron itself features paired gular, humeral, pectoral, abdominal, femoral, and anal scutes.[6] Hatchlings' carapaces have 3 keels while the plastrons have 2.[6]

Ecology

Habitat

During its lifetime, the loggerhead sea turtle inhabits terrestrial, oceanic, and neritic ecosystems.[5] Upon birth, hatchlings enter the ocean and swim away from land for several days. The "post-hatchlings" then return to find a home in a local downwelling.[5] Downwellings are created when surface waters converge and feature floating material such as seaweed. They may stay there for months, feeding on floating material and occasionally swimming to keep from getting too cold. Loggerheads are endangered if water temperatures dip below 50 °F (10 °C).[7] Eventually, they enter ocean currents farther from shore and move into the oceanic zone[5] where they remain until they become juveniles (between ages 7 and 12). They then move into the neritic zone and remain there until reaching adulthood.[5] The neritic zone is also an important area for adult foraging and inter-nesting habitats.[5]

Feeding

The loggerhead mainly feeds on bottom dwelling invertebrates, including horseshoe crabs, clams, mussels, molluscs, crustaceans, fish, jellyfish, crabs, shrimp, Portuguese Man o' War and other small to medium-sized marine animals, which they crush with their large and powerful jaws. Their powerful jaw muscles allow them to easily crush shellfish.[5] During migration through the open sea, loggerheads eat jellyfishes, floating mollusks, floating egg clusters, squids, and flying fishes.[citation needed]

Life history

Breeding

The female first creates a nest and then enters a mating period that can last up to six weeks.[8] Loggerhead sea turtles court their mates, but these behaviors have not been thoroughly examined. Nuzzling, biting, head movements, and flipper movements may be forms of male courtship behavior. Females may produce cloacal pheromones to suggest reproductive ability. Males approach females and attempt to mount them, while females resist. The male and female may circle each other. If the male has competitors, the female may leave the males to struggle with each other. The winner then mounts the female. Other courting males bite the male during mating, damaging his flippers and tail, sometimes to the point of exposing bones. Such damage can cause the male to dismount, requiring weeks to heal. The male's curved claws damage the shoulders of the female's shell when he mounts her. He may injure her by biting her neck during mating.[8]

 Photo of a baby loggerhead crawling across the sand.
Baby Loggerhead Sea Turtle.

Over several months, females produces several egg clutches and then become quiescent, producing no eggs for an average two to three years. Unlike other sea turtles, courtship and mating usually do not take place near the nesting beach but rather along migration routes between feeding and breeding grounds. In the Mediterranean, Loggerheads mate from late March to early June. The nesting season peaks in June and July, but varies by nesting beach.

All sea turtles have similar basic nesting behaviors. Females return to lay their eggs on or near the beach where they hatched. They haul out of the water, climb the beach, excavate a body pit, lay eggs, fill the egg chamber, fill the body pit, and finally return to sea. The nesting area must be selected carefully because it will have an affect on characteristics of the turtle such as fitness, emergence ratio, and vulnerability to nest predators.[8] Clutch size ranges from 70mm to 150mm. Each egg is roughly the size and shape of a ping-pong ball.

 A female loggerhead sea turtle from the back, laying eggs into the hole it has dug.
Loggerhead Sea Turtle laying eggs.

Loggerhead turtles are the most common sea turtle to nest in the United States. Loggerheads nest from Texas to North Carolina, requiring soft sandy beaches where there is little or light pollution. The largest concentration is in south Florida. Florida statistics since 1998 describe its lowest nesting levels in 17 years, declining from 85,988 nests in 1998 to approximately 45,084 in 2007.[citation needed]

Early life

The loggerheads sex is determined during incubation. Temperatures range from 26-32 degrees Celsius. Lower temperatures produce males. [2] After incubating for approximately 80 days, hatchlings emerge, usually at night when predation is reduced. Hatchlings average about 1.8 inches (4.6 cm) in length and weigh around 20 grams (0.71 oz) at birth.[5]

Maturation

Once in the ocean, Atlantic turtles use ocean currents to travel to the Sargasso Sea using the Sargassum as protection until they mature.[9]

An alternative to migration is hibernation to varying degrees as the water cools. By February they submerge for up to seven hours at a time, emerging for only seven minutes to breathe. Although outdone by freshwater turtles, these are the longest recorded dives for any air-breathing marine vertebrate.[10]

Loggerheads mature sexually at around age 35.[5] Nesting loggerheads have a straight caraprace length of 70–109 centimetres (28–43 in). Seventy is the minimum size for breeding. However, not all loggerheads begin nesting after reaching minimum size. Therefore carapace length is not a good indicator of sexual maturity.[8] Most loggerheads that reach adulthood live more than 30 years, and often live past 50 years.

Like all marine turtles, the loggerhead prepares for reproduction in its foraging area. This takes place several years before the loggerhead migrates to a mating area.[8]

Importance to humans

Loggerhead Sea Turtles were once intensively hunted for their meat and eggs. Turtle meat and eggs in general are still widely used today for food sources despite international regulations. In Mexico, eating turtle eggs is a very common tradition and the eggs can be easily bought in the markets. The locals claim that the turtle egg acts as an aphrodisiac.[11] However, eating loggerhead meat or eggs can lead to serious illness. This can be attributed to high levels of toxic metals that build up in the turtles through bioaccumulation and harmful bacteria that can be found in the loggerheads such as Pseudomonas aeruginosa and Serratia marcescens.[12][13] Their fat is used in cosmetics and medication and the shells are used to make items such as combs. As a result, C. caretta is now internationally protected.

Conservation

Threats

 An orange diamond sign with the words "Loggerhead Turtle Nesting Area" is blocking off a roped-off area on the beach where a loggerhead has laid eggs.
Loggerhead Sea Turtle nest roped off as part of the Sea Turtle Protection Project on Hilton Head Island.

Fishing gear is currently the biggest threat to loggerheads. They are most commonly caught in longlines and gillnets but have also been accidentally captured in traps, pots, trawls, and dredges.[5] Caught in this gear, loggerheads risk serious injury or drowning. Turtle excluder devices for nets and other gear help protect them. Due to the fishing traps, the loggerheads are now on the endangered list.

Because hatchlings usually follow the brightest light to the ocean's edge, artificial lights can lead them astray.

Since loggerheads spend a significant portion of their lives in open ocean environments, floating debris such as plastic pellets and ghost fishing gear can be ingested, causing harm. [14]

Fibropapillomatosis disease has been a problem for loggerheads. It causes internal and external tumors that disrupt essential behaviors and, if on the eyes, permanent blindness.[14]

Human activities are often threatening to many marine animals. Loggerhead turtles are no exception. The creation of docks and marinas can destroy near shore habitats and risk harming the loggerheads even more through gasoline and oil discharge at the marina fueling stations. Boat traffic and dredging can degrade the loggerheads' habitats and can also potentially injure or kill loggerheads by hitting them. [14]

 A Loggerhead sea turtle escapes a circular fisherman's net via a TED.
Loggerhead Sea Turtle escapes from fishing net through a Turtle Excluder Device (TED).

Conservation Efforts

Since the loggerhead occupies such a broad range, successful conservation requires efforts from multiple countries.[5]

Loggerhead turtles are classified as endangered by the International Union for the Conservation of Nature and are listed under Appendix I of the Convention on International Trade in Endangered Species, making international trade of the loggerhead sea turtle illegal.[5] In the United States, the U.S. Fish and Wildlife Service and National Marine Fisheries Service classify them as a threatened species under the Endangered Species Act.[15] Loggerheads are listed as Endangered under both Australia's Environment Protection and Biodiversity Conservation Act 1999 and Queensland's Nature Conservation Act 1992. Annex 2 of the Specially Protected Areas and Wildlife (SPAW) Protocol of the Cartagena Convention also protects the loggerheads. The National Marine Fisheries Service and the U.S. Fish and Wildlife services hold joint jurisdiction over the loggerhead sea turtles in the United States[5]

The turtles inhabit the Italian islands of Lampedusa and Linosa, off the coast of Sicily, and in Calabria, where they are particularly endangered. They also nest on the beaches of Cyprus, especially Akamas and Alagadi Beach.[16]

In many places during the nesting season, workers search the coastline for nests. They uncover the nest and count its eggs. If necessary they relocate it to protect it from humans and other threats. Plastic fencing can protect eggs from large predators such as raccoons or even dogs. The fence's gauge must be large enough to allow the hatchlings to pass through without difficulty. The nests are checked daily for disturbances. Several days after eggs hatch, the workers examine the nest, tallying hatched eggs, undeveloped eggs, and dead hatchlings. Any remaining live hatchlings are either raised and released or taken to research facilities. Those that appear strong and healthy may be immediately released to the ocean. Typically, those that lacked the strength to hatch and climb to the surface by that point would have died otherwise.

Hatchlings use the journey from nest to ocean to build strength for the coming swim. Therefore, helping them to reach the ocean lowers their chances of survival.[citation needed] The Fripp Island, South Carolina Turtle Patrol each year arranges driftwood to guide the hatchlings towards the closest shoreline.

Gallery

Footnotes

  1. ^ a b c d e f g h i j k l m n o p Spotila, James R. (2004). Sea Turtles: A Complete Guide to their Biology, Behavior, and Conservation. Baltimore, Maryland: The Johns Hopkins University Press and Oakwood Arts. pp. 163–179. ISBN 0-8018-8007-6. 
  2. ^ a b c MarineBio (March). "Caretta caretta Loggerhead Sea Turtle". http://marinebio.org/species.asp?id=163. Retrieved March 2, 2010. 
  3. ^ a b c d e "Loggerhead Sea Turtle (Caretta caretta)" (in English). Virginia: Virginia Herpetological Society. http://fwie.fw.vt.edu/VHS/reptiles/turtles/loggerhead-sea-turtle/loggerhead_sea_turtle.htm. Retrieved 2/15/10. 
  4. ^ [1]
  5. ^ a b c d e f g h i j k l m n o p q Bolten, A.B. (2003). "Loggerhead Turtle (Caretta caretta)". NOAA Fisheries. NOAA Fisheries. http://www.nmfs.noaa.gov/pr/species/turtles/loggerhead.htm. Retrieved January 31, 2010. 
  6. ^ a b c d e f g Terese, Conant; Peter H. Dutton, Tomoharu Eguchi, Sheryan P. Epperly, Christina C. Fahy, Matthew H. Godfrey, Sandra L. MacPherson, Earl E. Possardt, Barbara A. Schroeder, Jeffrey A. Seminoff, Melissa L. Snover, Carrie M. Upite, and Blair E. Witherington (August 2009). [http://www.seaturtle.org/PDF/ConantTA_2009_NMFSTechReport.pdf LOGGERHEAD SEA TURTLE (CARETTA CARETTA) 2009 STATUS REVIEW UNDER THE U.S. ENDANGERED SPECIES ACT]. Loggerhead Biological Review Team. pp. 222. http://www.seaturtle.org/PDF/ConantTA_2009_NMFSTechReport.pdf. 
  7. ^ a b Basu, Rebecca (1 March 2010). "Winter is culprit in manatee death toll". Melbourne, Florida: Florida Today. pp. 3A. http://www.floridatoday.com/article/20100301/NEWS01/3010320/Winter-is-culprit-in-manatee-death-toll. 
  8. ^ a b c d e f Miller, Jeffrey D.; Limpus, Colin J.; Godfrey, Matthew H. (in press). [http://ww2.coastal.edu/msci302/ST-repro2.pdf "Nest Site Selection, Oviposition, Eggs, Development, Hatching and Emergence of Loggerhead Turtles"]. in Bolten, Alan; Witherington, Blair. Biology and Conservation of Loggerhead Sea Turtle. University of Florida Press. http://ww2.coastal.edu/msci302/ST-repro2.pdf. Retrieved February 2, 2010. 
  9. ^ BBC NEWS | Science/Nature | Turtles return home after UK stay
  10. ^ Hochscheid, S.; F. Bentivegna, G.C. Hay (2005). "First records of dive durations for a hibernating sea turtle". Biol. Lett. 1 (1): 82–6. doi:10.1098/rsbl.2004.0250. PMID 17148134. PMC 1629053. http://rsbl.royalsocietypublishing.org/content/1/1/82.long. Retrieved 2 December 2009. "These represent the longest dives ever reported for a diving marine vertebrate.". 
  11. ^ a b Dellios, Hugh (September 18). "Mexico Cracks Open Myth of Sea Turtle Eggs". http://www.redorbit.com/news/science/243267/mexico_cracks_open_myth_of_sea_turtle_eggs/index.html. Retrieved March 2, 2010. 
  12. ^ a b Aguirre, A.; Gardner, S. (July 12). "Hazards Associated with the Consumption of Sea Turtle Meat and Eggs: A Review for Health Care Workers and the General Public". http://www.springerlink.com/content/l641g4k4870q234v/. Retrieved March 2, 2010. 
  13. ^ a b Craven, Kathryn; Taylor, Judy (September 11). "Identification of Bacterial Isolates from Unhatched Loggerhead (Caretta caretta) Sea Turtle Eggs in Georgia, USA". http://www.seaturtle.org/mtn/archives/mtn115/mtn115p9.shtml. Retrieved March 16, 2010. 
  14. ^ a b c d NOAA Fisheries (2010). "Threats to Marine Turtles". Endangered marine animal preservation. NOAA Fisheries Office of Protected Resources. http://www.nmfs.noaa.gov/pr/species/turtles/threats.htm. Retrieved February 7, 2010. 
  15. ^ [2]
  16. ^ "Conservation of the Akamas Peninsula (Cyprus)" (PDF). CONVENTION ON THE CONSERVATION OF EUROPEAN WILDLIFE AND NATURAL HABITATS. Strasbourg: Council of Europe. 29 November - 3 December-2004. http://www.coe.int/t/dg4/cultureheritage/conventions/Bern/T-PVS/sc24_files15_en.pdf. Retrieved 2009-01-02. 
  17. ^ My Scribe (May 8, 2009). "Loggerhead Facts". Loggerhead preservation. KiawahTurtles. http://www.kiawahturtles.com/TurtleFAQ.htm. Retrieved February 4, 2010. 

References

  • Loggerhead Sea Turtles. Washington, D.C.: Smithsonian Books. 2003. 
  • Lutz, Peter L.; Musick, John A.; Wyneken, Jeanette (1997). The Biology of Sea Turtles. 1. Boca Raton, Florida: CRC Press. 
  • Lutz, Peter L.; Musick, John A.; Wyneken, Jeanette (2003). The Biology of Sea Turtles. 2. Boca Raton, Florida: CRC Press. 
  • Gulko, D.; Eckert, K.L. (2004). Sea Turtles: An Ecological Guide. Honolulu, Hawai’i: Mutual Publishing. 

External links


Loggerhead sea turtle
Fossil range: 110–0 Ma
Cretaceous - Recent[1]
File:Loggerhead sea
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Subclass: Anapsida
Order: Testudines
Family: Cheloniidae
Genus: Caretta
Rafinesque, 1814
Species: C. caretta
Binomial name
Caretta caretta
Leonhard Stejneger, 1902

The loggerhead sea turtle (Caretta caretta), or loggerhead, is an oceanic turtle distributed throughout the world. It is a marine reptile, belonging to the family Cheloniidae. The loggerhead is the world's largest hard-shelled turtle, measuring up to 213 centimeters (84 in) long when fully grown. The adult loggerhead sea turtle weighs approximately 135 kilograms (300 lb). The skin ranges from yellow to brown in color, and the shell is typically reddish-brown. There are no external differences in gender until the turtle becomes an adult. The most obvious difference being, adult males have thicker tails and shorter plastrons than the females.

The loggerhead sea turtle is found in the Atlantic, Pacific, and Indian oceans as well as the Mediterranean Sea. It spends most of its life in saltwater and estuarine habitats, with females briefly coming ashore to lay eggs. The loggerhead sea turtle has a low reproductive rate; females lay an average of four egg clutches and then become quiescent, producing no eggs for two to three years. The loggerhead reaches sexual maturity within 17–33 years and has a lifespan of 47–67 years.

The loggerhead sea turtle is omnivorous, feeding mainly on bottom dwelling invertebrates. Its large and powerful jaws serve as an effective tool in dismantling its prey. Young loggerheads are exploited by numerous predators; the eggs are especially vulnerable to terrestrial organisms. Once the turtles reach adulthood, their formidable size limits predation to large marine organisms such as sharks.

Loggerheads are considered an endangered species and are protected by the International Union for the Conservation of Nature. Untended fishing gear is responsible for many loggerhead deaths. Turtles may also suffocate if they are trapped in fishing trawls. Turtle excluder devices (TEDs) have been implemented in efforts to reduce mortality by providing the turtle an escape route. Loss of suitable nesting beaches and the introduction of exotic predators has also taken a toll on loggerhead populations. Efforts to restore their numbers will require international cooperation since the turtles roam vast areas of ocean and critical nesting beaches are scattered among several countries.

Contents

Taxonomy

Carolus Linnaeus gave the loggerhead its first binomial name, Testudo caretta, in 1758.[3][5] Thirty-five other names emerged over the following two centuries, with the combination Caretta caretta first introduced in 1902 by Leonhard Stejneger.[4] The English common name "loggerhead" refers to the animal's large head.[6][7] The Loggerhead sea turtle belongs to the family Cheloniidae, which includes all sea turtles except the leatherback sea turtle.[8] The subspecific classification of the loggerhead sea turtle is debated, but most authors consider the loggerhead a single polymorphic species.[9] Molecular genetics has confirmed hybridization of the Loggerhead sea turtle with the Kemp's Ridley, Hawksbill turtle, and Green turtle. The extent of natural hybridization is not yet determined; however, second generation hybrids have been reported, suggesting that some hybrids are fertile.[10]

Habitat

Loggerhead sea turtles spend most of their life in the open ocean and in shallow coastal waters. They rarely come ashore, with the exception of the females' brief visits to construct nests and deposit eggs. Hatchling loggerhead turtles live in floating mats of Sargassum algae.[11] Adults and juveniles live along the continental shelf as well as in shallow coastal estuaries.[12] In the northwestern Atlantic Ocean, age plays a factor in habitat preference. Juveniles are more frequently found in shallow estuarine habitats with limited ocean access compared to non-nesting adults.[13] Loggerheads occupy waters with surface temperature ranging from

  1. REDIRECT Template:Convert/°C during non-nesting season. Temperatures from
  2. REDIRECT Template:Convert/°C are most suitable for nesting females.[14]

Juvenile loggerheads share the Sargassum habitat with a variety of other organisms. The mats of Sargassum contain as many as 100 different species of animals that the juveniles feed on. Some of the prey, such as ants, flies, aphids, leafhoppers, and beetles are carried by the wind to these areas. Endemic prey of the Sargassum mats include barnacles, small crab larvae, fish eggs, and hydrozoan colonies.[11] Marine mammals and commercial fish such as tuna, dolphin fish, and amberjacks also inhabit the Sargassum mats.[15]

Distribution

File:Loggerhead
Range of the loggerhead sea turtle

The loggerhead sea turtle has a cosmopolitan distribution, nesting over the broadest geographical range of any sea turtle. The loggerhead inhabits the Atlantic, Indian, and Pacific Oceans and the Mediterranean Sea.[16]

In the Atlantic Ocean, the greatest concentration of loggerheads is along the southeastern coast of North America and in the Gulf of Mexico.[16] Very few loggerheads are found along the European and African coastlines.[17] Florida is the most popular nesting site with over 67,000 nests built per year.[17] Nesting extends as far north as Virginia, as far south as Brazil, and as far east as the Cape Verde Islands.[17] The Cape Verde Islands are the only significant nesting site on the eastern side of the Atlantic. Loggerheads found in the Atlantic Ocean feed from Canada to Brazil.[16]

In the Indian Ocean, loggerheads feed along the coastlines of Africa, the Arabian Peninsula, and in the Arabian Sea.[18] Along the African coastline, loggerheads nest from Mozambique's Bazaruto Archipelago to South Africa's St Lucia estuary.[19] The largest Indian Ocean nesting site is Oman, on the Arabian Peninsula, which hosts around 15,000 nests, giving it the second largest nesting population of loggerheads in the world. Western Australia is another notable nesting area, with 1,000-2,000 nests per year.[18]

Pacific loggerheads live in temperate to tropical regions.[19] They forage in the East China Sea, the southwestern Pacific, and along the Baja California peninsula. Eastern Australia and Japan are the major nesting areas. Pacific loggerheads occasionally nest in Vanuatu and Tokelau. Yakushima Island is the most important site with three nesting grounds that 40% of all nearby loggerheads visit.[18] Post-nesting females often find homes in the East China Sea, while the Kuroshio Extension Bifurcation region provides important juvenile foraging areas.[19] Eastern Pacific populations are concentrated off the coast of Baja California, where upwelling provides rich feeding grounds for juvenile turtles and subadults (individuals past the juvenile stage that lack adult characteristics). Nesting sites along the eastern Pacific Basin are rare. mtDNA sequence polymorphism analysis and tracking studies suggest that 95% of the population along the coast of the Americas hatch on the Japanese Islands in the western Pacific.[20] The turtles are transported by the prevailing currents across the full length of the northern Pacific, a migration route that is one of the longest of any marine animal.[20] The return journey to the natal beaches in Japan has been long suspected, although the trip would cross unproductive clear water with few feeding opportunities.[21] Evidence of a return journey came from an adult female loggerhead named Adelita, who in 1996, equipped with a satellite tracking device made the 9,000-mile trip from Mexico across the Pacific. Adelita was the first animal of any kind ever tracked across an ocean basin.[22]

The Mediterranean is a nursery for juveniles as well as a common place for adults in the spring and summer months.[17][23] Almost 45% of the Mediterranean juvenile population has migrated from the Atlantic.[17] Loggerheads feed in the Alboran Sea and the Adriatic Sea.[17] Greece is the most popular nesting site along the Mediterranean with more than 3,000 nests per year.[18] Because of this, Greek authorities do not allow planes to take off or land at night in Zakynthos due to the nesting turtles.[24] In addition to the Greek coast, the coastlines of Cyprus and Turkey are also common nesting sites.[18]

Evolutionary history

All modern sea turtles likely descended from a single common ancestor during the Cretaceous Period; however evidence of a specific ancestor is lacking.[25] Loggerheads, like all other sea turtles excluding the leatherback sea turtle, are members of the ancient family Cheloniidae. About 40 million years ago, a branch of the Cheloniidae gave rise to loggerhead sea turtle.[26] Of the six species of living Cheloniidae, loggerheads are more closely related to the Kemp's Ridley sea turtle, olive Ridley, and the hawksbill turtle than they are to the flatback turtle and the green turtle.

Around three million years ago, during the Pliocene epoch, Central America emerged from the sea, effectively cutting off currents between the Atlantic and Indo-Pacific Oceans. The rerouting of ocean currents lead to climatic changes as the Earth entered a glacial cycle. Cold water upwelling around the Cape of Good Hope and reduction in water temperature at Cape Horn formed cold water barriers to migrating turtles. The result was a complete isolation of the Atlantic and Pacific populations of loggerheads.[27] During the most recent ice age, the beaches of southeastern North America were too cold for sea turtle eggs. As the Earth began to warm, loggerheads moved farther north, colonizing the northern beaches. Because of this, turtles nesting between North Carolina and northern Florida represent a different genetic population than those in southern Florida.[27]

The distinct populations of loggerheads have unique characteristics and genetic differences. For example, Mediterranean loggerheads are smaller, on average, than Atlantic Ocean loggerheads.[18] North Atlantic and Mediterranean loggerhead sea turtles are descendants of colonizing loggerheads from Tongaland, South Africa. South African loggerhead genes are still present in these populations today.[27]

Anatomy and morphology

File:Loggerhead
The carapace of this loggerhead is reddish-brown; five vertebral scutes run down the turtle's midline bordered by five pairs of costal scutes.

The loggerhead sea turtle is the world's largest hard-shelled turtle.[28] Adult loggerheads have an average weight range of 80 to 200 kilograms (180 to 440 lb) and a length range of 70 to 95 centimeters (28 to 37 in).[28] The maximum reported weight is 545 kilograms (1,200 lb) and the maximum carapace length is 213 centimeters (84 in).[28] The head and carapace (upper shell) range from a yellow-orange to a reddish-brown, while the plastron (underside) is typically pale yellow.[29] The turtle's neck and sides are brown on the tops and yellow on the sides and bottom.[6]

The turtle's shell is divided into two sections: carapace and plastron. The carapace is further divided into large plates, or scutes.[29] Typically, there are 11 or 12 pairs of marginal scutes which rim the carapace.[5] Five vertebral scutes run down the carapace's midline, while five pairs of costal scutes border them.[30] The nuchal scute is located at the base of the head.[30] The carapace connects to the plastron by 3 pairs of inframarginal scutes forming the bridge of the shell.[30] The plastron features paired gular, humeral, pectoral, abdominal, femoral, and anal scutes.[5] The shell serves as external armor, although loggerhead sea turtles cannot retract their head or flippers into their shells.[31]

Sexual dimorphism of the loggerhead sea turtle is only apparent in adults. Adult males have longer tails and claws than females. The males' plastrons are shorter than the females', presumably to accommodate the males' larger tails. The carapace of males are wider and less domed than the females, and males typically have wider heads than females.[32] The sex of juveniles and subadults cannot be determined through external anatomy, but can be observed through dissection, laparoscopy (an operation performed on the abdomen), histological examination (cell anatomy), and radioimmunological assays (immune study dealing with radiolabeling).[32]

Lachrymal glands located behind each eye allow the loggerhead to maintain osmotic balance by eliminating the excess salt obtained from ingesting ocean water. On land, the excretion of excess salt gives the false impression that the turtle is crying.[33]

Ecology and behavior

File:Loggerhead Sea Turtle (Caretta caretta)
A resting loggerhead sea turtle

Loggerhead sea turtles observed in captivity and in the wild are most active during the day. In captivity, the loggerheads' daily activities are divided between swimming and resting on the bottom. While resting, loggerheads spread their forelimbs to about mid-stroke swimming position. They remain motionless with eyes open or half-shut and are easily alerted during this state. At night, captive loggerheads sleep in the same position with their eyes tightly shut and are slow to react.[14] Loggerheads spend up to 85% of their day submerged, with males being the more active divers than females. The average duration of dives is 15-30 minutes, but they can stay submerged for up to four hours.[34] Juvenile loggerheads and adults differ in their swimming methods. Juveniles keep their forelimbs pressed to the side of their carapace and propel themselves by kicking with their hind limbs. As the juveniles mature, their swimming method is progressively replaced with the adults' alternating-limb method. The loggerheads depend entirely on this method of swimming by the time they are one year old.[35]

Water temperature affects the sea turtle's metabolic rate.[14] Lethargy is induced at temperatures between

  1. REDIRECT Template:Convert/°C. The loggerhead takes on a floating, cold-stunned posture when temperatures drop to approximately
  2. REDIRECT Template:Convert/°C.[14] However, younger loggerheads are more resistant to cold and do not become stunned until temperatures drop below
  3. REDIRECT Template:Convert/°C. The loggerheads' migration help prevent instances of cold-stunning.[36] Higher water temperatures cause an increase in metabolism and heart rate. A loggerhead's body temperature increases in warmer waters more quickly than it decreases in colder water. The loggerhead sea turtle's critical thermal maximum is currently unknown.[36]

Female-female aggression, being uncommon (especially in marine vertebrates), is common among loggerheads. Ritualized aggression escalates from passive threat displays to combat. This conflict primarily occurs over access to feeding grounds. Escalation typically follows four steps.[37] First, initial contact is stimulated by visual or tactile cues. Second, confrontation occurs, beginning with passive confrontations characterized by wide head-tail circling. They begin aggressive confrontation when one turtle ceases to circle and directly faces the other. Third, sparring occurs with turtles snapping at each other’s jaws. The final stage, separation, is either mutual, with both turtles swimming away in opposite directions, or involves chasing one out of the immediate vicinity.[37] Escalation is determined by several factors including: hormone levels, energy expenditure, expected outcome, and importance of location. At all stages an upright tail shows willingness to escalate, while a curled tail shows willingness to submit. Because higher aggression is metabolically costly and potentially debilitating, contact is much more likely to escalate when the conflict is over access to good foraging grounds.[37] Further aggression has also been reported in captive loggerheads. The turtles are seemingly territorial and will fight with other loggerheads and sea turtles of different species.[34]

Predators

File:Fox up a
The red fox is a predator of loggerhead nests in Australia.

Loggerheads have numerous predators. Egg and nestling predators include oligochaete worms, beetles, fly larvae, ants, parasitoid wasp larvae, flesh flies, crabs, snakes, gulls, crows, opposums, bears, rats, armadillos, honey badgers, skunks, canids, raccoons, cats, hogs, and humans. During their migration from their nest to the sea, hatchlings are predated by dipteran larvae, crabs, toads, lizards, snakes, birds, and mammals. In the ocean, predators of the loggerhead juveniles and adults include sharks, other fish such as parrotfish, moray eels, portunid crabs, gulls, monk seals, and killer whales. Nesting females are attacked by flesh flies, feral dogs, and humans. Salt marsh mosquitoes can also pester nesting females.[38][39]

In Australia, the introduction of the red fox (Vulpes vulpes) by English settlers in the 19th century lead to significant reductions in loggerhead sea turtle populations. In one coastal section in eastern Australia during the 1970s, predation of turtle eggs destroyed up to 95% of all clutches laid.[40] Aggressive efforts to destroy foxes in the 1980s and 90's has reduced this impact; however, it is estimated that it will be the year 2020 before populations will experience complete recovery from such dramatic losses.[41]

Along the southeastern coast of the United States, the raccoon (Procyon lotor) is the most destructive predator of nesting sites. Mortality rates of nearly 100% of all clutches laid in a season have been recorded on some Florida beaches.[40] This is attributed to an increase in raccoon populations, which have flourished in urban environments. Aggressive efforts to protect nesting sites by covering them with wire mesh has significantly reduced the impact of raccoon predation on loggerhead sea turtle eggs.[41]

Up to 40% of nesting females around the world have wounds believed to come from shark attacks.[39] Infectious bacteria such as Pseudomonas and Salmonella attack loggerhead hatchlings and eggs. Fungi such as Penicillium infect loggerhead sea turtle nests and cloaca.[39]

Disease and parasites

Fibropapillomatosis disease threatens loggerheads in the form of the herpes-type virus which causes internal and external tumors. These tumors disrupt essential behaviors and, if on the eyes, cause permanent blindness.[42] Trematodes of the family Spirorchiidae inhabit tissues throughout the body of the loggerhead including vital organs such as the heart and the brain.[43] Trematode infection can be highly debilitating. For example, inflammatory trematode lesions can cause endocarditis and neurological disease.[43] A nematode, Angiostoma carettae, also infects loggerheads.[44] The nematodes cause histologic lesions in the respiratory tract.[44]

More than 100 species of animals from 13 phyla as well as 37 kinds of algae live on loggerheads' backs.[45] These parasitic organisms, which increase drag, offer no known benefit to the turtle, although it has been suggested that the dulling effect of organisms on shell color improves camouflaging ability.[45]

Feeding

File:Moon jelly - adult (rev2).jpg
An adult Aurelia jellyfish which loggerheads eat during migration through the open sea

The loggerhead sea turtle is omnivorous, feeding mainly on bottom dwelling invertebrates, such as gastropods, bivalves, and decapods. The loggerhead has a greater list of known prey than any other sea turtle. Other food items include sponges, corals, sea pens, polychaete worms, sea anemones, cephalopods, barnacles, brachiopods, isopods, insects, bryozoans, sea urchins, sand dollars, sea cucumbers, starfish, fish (eggs, juveniles, and adults), wrasses, hatchling turtles (including members of its own species), algae, and vascular plants.[46] During migration through the open sea, loggerheads eat jellyfish, floating mollusks, floating egg clusters, squid, and flying fish.[6]

Loggerheads crush prey with their large and powerful jaws.[6][38] Projecting scale points on the anterior margin of the forelimbs allow manipulation of the food. These points can be used as "pseudo-claws" to tear large pieces of food in the loggerhead's mouth. The loggerhead will turn its neck sideways to consume the torn food on the scale points.[38] Inward pointing, mucosal (mucus-covered) papillae found in the foreregion of the loggerhead's esophagus filter out foreign bodies, such as fish hooks. The next region of the esophagus is nonpapillated with numerous mucosal folds. The rate of digestion in loggerheads is temperature dependent; digestion rate increases as temperature increases.[38]

Life history

Early life

[[File:|thumb|upright|right|alt= Loggerhead hatchling crawling through the sand.|A hatchling loggerhead sea turtle on its journey to the ocean]]

Hatchlings range in color from light brown to almost black, lacking the adult's distinct yellows and reds.[29] Upon hatching, they measure approximately 4.6 centimeters (1.8 in) and weigh about 20 grams (0.71 oz).[6] The eggs are typically laid on the beach in an area above the high-tide line. The eggs are laid near the water so that hatchlings can return to the sea.[47] The loggerhead's sex is dictated by the temperature of the underground nest. Incubation temperatures generally range from

  1. REDIRECT Template:Convert/°C. Studies show that loggerhead sea turtle eggs that are kept at a constant incubating temperature of
  2. REDIRECT Template:Convert/°C become females. Eggs incubating at
  3. REDIRECT Template:Convert/°C become males. An incubation temperature of
  4. REDIRECT Template:Convert/°C results in an equal ratio of male to female hatchlings.[48] Hatchlings from eggs in the middle of the clutch tend to be the largest, grow the fastest, and be the most active during the first few days of sea life.[40]

After incubating for approximately 80 days, hatchlings dig through the sand to the surface. This usually occurs at night when darkness increases the chance of escaping predation and damage from extreme sand surface temperatures is reduced.[47] Hatchlings enter the ocean by navigating toward the brighter horizon created by the reflection of the moon and star light off the water's surface.[49]

Hatchlings can lose up to 20% of their body mass due to evaporation of water as they journey from nest to ocean.[50] Hatchlings initially use the undertow to push them five to ten yards away from the shore.[50] Once in the ocean, hatchling loggerheads swim for about 20 hours, bringing them far offshore.[29] The loggerhead sea turtles have an iron compound, magnetite, in their brains that allows the turtles to perceive the Earth's magnetic field.[51] They use this magnetite for navigation. Many hatchlings use Sargassum in the open ocean as protection until they reach 45 centimeters (18 in).[29] Hatchling loggerheads live in this pelagic environment until they reach juvenile age, and then they migrate to nearshore waters.[29]

Maturation

File:Caretta caretta
A mature loggerhead sea turtle

When ocean waters cool, loggerheads must migrate to warmer areas or hibernate to some degree. In the coldest months, loggerheads submerge for up to seven hours at a time, emerging for only seven minutes to breathe. Although outdone by freshwater turtles, these are among the longest recorded dives for any air-breathing marine vertebrate.[52] During their seasonal migration, juvenile loggerheads have the ability to use both magnetic and visual cues.[53] When both aids are available, they are used in conjunction; if one aid is not available, the other suffices.[53]

Like all marine turtles, the loggerhead prepares for reproduction in its foraging area. This takes place several years before the loggerhead migrates to a mating area.[54] Female loggerheads first reproduce at ages 28–33 in Southeastern United States and Australia, and at ages 17–30 in South Africa. Age at first reproduction in the Mediterranean, Oman, Japan, and Brazil are unknown.[55] Nesting loggerheads have a straight carapace length of 70–109 centimeters (28–43 in). Because of the large range, carapace length is not a reliable indicator of sexual maturity.[54] Loggerheads have an estimated maximum lifespan of 47–67 years in the wild.[46]

Reproduction

File:Logger eggs
A loggerhead sea turtle laying eggs

Female loggerheads first reproduce between the ages of 17–33.[55] The loggerhead mating period lasts up to six weeks.[56] Loggerhead sea turtles court their mates, however these behaviors have not been thoroughly examined.[57] Male forms of courtship behavior include nuzzling, biting, and head and flipper movements.[57] Studies suggest that females produce cloacal pheromones to indicate reproductive ability.[57] Before mating, the male approaches a female and attempt to mount her, while she resists. Next, the male and female begin to circle each other. If the male has competitors, the female may let the males struggle with each other. The winner then mounts the female; the male's curved claws usually damage the shoulders of the female's shell during this process. Other courting males bite the male while he is attempting to copulate, damaging his flippers and tail, possibly exposing bones. Such damage can cause the male to dismount and require weeks to heal.[57] While nesting, females produce an average of 3.9 egg clutches and then become quiescent, producing no eggs for an average of two to three years.[56][58] Unlike other sea turtles, courtship and mating usually do not take place near the nesting beach but rather along migration routes between feeding and breeding grounds.[57] Recent evidence indicates that ovulation in loggerheads is mating-induced.[59] Through the act of mating, the female ovulates eggs which are fertilized by the male. This is unique as mating-induced ovulation is rare outside of mammals.[59] In the Mediterranean, loggerheads mate from late March to early June. The nesting season peaks in June and July, but varies by nesting beach.[60]

Loggerheads may display multiple paternity.[61] Multiple paternity is possible due to sperm storage. The female can store sperm from multiple males in her oviducts until ovulation.[62] A single clutch may have as many as five fathers, each contributing sperm to a portion of the clutch.[61] Multiple paternity and female size are positively correlated.[61] Two hypotheses explain this correlation. One states that males favor large females due to perceived higher fecundity (ability to reproduce).[61] The other states that because larger females are able to swim more quickly to mating grounds, they have a longer mating period.[61]

All sea turtles have similar basic nesting behaviors. Females return to lay eggs at intervals of 12–17 days during the nesting season, on or near the beach where they hatched.[58][63] They exit the water, climb the beach, and scrape away the surface sand to form a body pit. With their hind limbs, they excavate an egg chamber in which the eggs are deposited. The females then cover the egg chamber and body pit with sand, and finally return to the sea.[64] This process takes 1–2 hours and occurs in open sand areas or on top of sand dunes.[58] The nesting area must be selected carefully because it affects characteristics such as fitness, emergence ratio, and vulnerability to nest predators.[47] Loggerhead sea turtles have an average clutch size of 110 eggs.[65]

Conservation

Loggerhead sea turtles were once intensively hunted for their meat and eggs, however consumption has decreased due to worldwide illegality. Despite this, turtle meat and eggs are still consumed in countries where regulations are not strictly enforced.[66] In Mexico, turtle eggs are a common meal; locals claim that the egg is an aphrodisiac.[67] Eating turtle eggs or meat can cause serious illness due to harmful bacteria such as Pseudomonas aeruginosa and Serratia marcescens, and high levels of toxic metals that build up through bioaccumulation[66][68]

Threats

File:Loggerhead nesting
Loggerhead sea turtle nest roped off as part of the Sea Turtle Protection Project on Hilton Head Island

Many human activities have negative effects on loggerhead sea turtle populations. The prolonged time required for loggerheads to reach sexual maturity and the high mortality rates of eggs and young turtles from natural phenomena compound the problems of population reduction as a consequence of human activities.[69]

Fishing gear is the biggest threat to loggerheads in the open ocean. They often become entangled in longlines or gillnets. They also become stuck in traps, pots, trawls, and dredges.[6] Caught in this unattended equipment, loggerheads risk serious injury or drowning. Turtle excluder devices for nets and other traps reduce the number being accidentally caught.

Nearly 24,000 metric tons of plastic is dumped into the ocean each year. Turtles ingest a wide array of this floating debris, including plastic bags, plastic sheets, plastic pellets, balloons and abandoned fishing line.[70] Loggerheads may mistake the floating plastic for jellyfish, a common food item. The ingested plastic causes numerous health concerns including: intestinal blockage, reduced nutrient absorption, suffocation, ulcerations, malnutrition or starvation. Ingested plastics release toxic compounds, including polychlorinated biphenyls, which may accumulate in internal tissues. Such toxins may lead to a thinning of eggshells, tissue damage or deviation from natural behaviors.[71]

Artificial lighting discourages nesting and interferes with the hatchlings ability to navigate to the water's edge. Females prefer nesting on beaches free of artificial lighting. On developed beaches, nests are often clustered around tall buildings, perhaps because they block-out the man-made light sources.[54] Loggerhead hatchlings are drawn toward the brighter area over the water which is the consequence of the reflection of moon and star light. Loggerheads, confused by the brighter artificial light, navigate inland, away from the protective waters. This exposes them to dehydration and predation as the sun rises.[49] Artificial lighting causes tens of thousands of hatchling deaths per year.[72]

Destruction and encroachment of habitat by humans is another threat to loggerhead sea turtles. Optimum nesting beaches are open sand beaches above the high tide line. However, beach development deprives them of suitable nesting areas, forcing them to nest closer to the surf.[58] Urbanization often leads to the siltation of sandy beaches, decreasing their viability.[58] Construction of docks and marinas can destroy near shore habitats. Boat traffic and dredging degrades habitat and can also injure or kill turtles when boats collide with turtles at or near the surface.[42]

Annual variations in climatic temperatures can effect sex ratios, since loggerheads have temperature-dependent sex determination. High sand temperatures may skew gender ratios in favor of females. In one study (Mrosovsky, N. & Provancha, J. 1992), nesting sites exposed to unseasonably warm temperatures over a three year period produced 87-99% females. This raises concerns over the connection between rapid global temperature changes and the possibility of population extinction.[73] A more localized effect on gender skewing comes from the construction of tall buildings which reduce sun exposure, lowering the average sand temperature. This results in a shift in gender ratios that favor the emergence of male turtles.[40] Global warming affects the sex of the turtles.

Conservation efforts

File:Logger ted
Loggerhead sea turtle escapes from fishing net through a turtle excluder device (TED)

Since the loggerhead occupies such a broad range, successful conservation requires efforts from multiple countries.[6]

Loggerhead sea turtles are classified as endangered by the International Union for the Conservation of Nature and are listed under Appendix I of the Convention on International Trade in Endangered Species, making international trade illegal.[6] In the United States, the U.S. Fish and Wildlife Service (FWS) and National Marine Fisheries Service (NMFS) classify them as a threatened species under the Endangered Species Act.[6] Loggerheads are listed as endangered under both Australia's Environment Protection and Biodiversity Conservation Act 1999 and Queensland's Nature Conservation Act 1992. The Convention on Migratory Species works for the conservation of loggerhead sea turtles on the Atlantic coast of Africa as well as in the Indian Ocean and South-East.[74][75] Throughout Japan, the Sea Turtle Association of Japan aids in the conservation of loggerhead sea turtles.[76] Greece's ARCHELON works for the conservation of loggerheads.[77] The Marine Research Foundation works for conservation of loggerhead sea turtles in Oman.[78] Annex 2 of the Specially Protected Areas and Wildlife (SPAW) Protocol of the Cartagena Convention, which deals with pollution that could harm marine ecosystems, also protects them.[6][79] Conservation organizations worldwide have worked with the shrimp trawling industry to develop TEDs that can exclude even the largest turtles. These devices are mandatory for all shrimp trawlers.[6]

File:Loggerhead
Loggerhead sea turtle, Evelyn, is equipped with a satellite tag and a SeaTag-GEO solar powered, geomagnetic tag

In many places during the nesting season, workers and volunteers search the coastline for nests.[80] During the nesting season, researchers may also go out during the evening to look for nesting females for tagging studies, gather barnacles, and tissues samples. Volunteers may, if necessary, relocate the nests for protection from threats such as high spring tides and predators, and monitor the nests daily for disturbances. After the eggs hatch, volunteers uncover and tally hatched eggs, undeveloped eggs, and dead hatchlings. Any remaining live hatchlings are released or taken to research facilities. Typically, those that lack the vitality to hatch and climb to the surface die.[81] Hatchlings use the journey from nest to ocean to build strength for the coming swim. Helping them to reach the ocean bypasses this strength building exercise and lowers their chances of survival.[81]

References

Footnotes
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Bibliography

Further reading

  • Bolten, Alan B.; Witherington, Blair E. (2003). Loggerhead Sea Turtles. Washington, D.C.: Smithsonian Books. ISBN 1588341364. 
  • Lutz, Peter L.; Musick, John A.; Wyneken, Jeanette (1997). The Biology of Sea Turtles. 1. Boca Raton, Florida: CRC Press. ISBN 0849384222. 
  • Lutz, Peter L.; Musick, John A.; Wyneken, Jeanette (2003). The Biology of Sea Turtles. 2. Boca Raton, Florida: CRC Press. ISBN 0849384222. 
  • Gulko, D.; Eckert, K.L. (2004). Sea Turtles: An Ecological Guide. Honolulu, Hawai’i: Mutual Publishing. ISBN 1566476518. 

External links



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