Six of the seven sea turtle species, all but the flatback, are present in U.S. waters, and are listed as endangered and/or threatened under the Endangered Species Act.[6] All but the flatback turtle are listed as threatened with extinction globally on the IUCN Red List of Threatened Species. The flatback turtle is found only in the waters of Australia, Papua New Guinea, and Indonesia.[7][6]
Sea turtles can be categorized as hard-shelled (cheloniid) or leathery-shelled (dermochelyid).[8] The only dermochelyid species of sea turtle is the leatherback.[8]
Description
For each of the seven species of sea turtles, females and males are the same size. As adults, it is possible to tell male turtles from female turtles by their long tails with a cloacal opening near the tip. Adult female sea turtles have shorter tails, with a cloacal opening near the base. Hatchling and sub-adult turtles do not exhibit sexual dimorphism; it is not possible to determine their sex by looking at them.[9]
In general, sea turtles have a more fusiform body plan than their terrestrial or freshwater counterparts. This tapering at both ends reduces volume and means that sea turtles cannot retract their head and limbs into their shells for protection, unlike many other turtles and tortoises.[10] However, the streamlined body plan reduces friction and drag in the water and allows sea turtles to swim more easily and swiftly.
The leatherback sea turtle is the largest sea turtle, reaching 1.4 to more than 1.8 m (4.6 to 5.9 ft) in length and weighing between 300 and 640 kg (661 to 1,411 lbs).[11] Other sea turtle species are smaller, ranging from as little as 60 cm (2 ft) long in the case of the Kemp's ridley, which is the smallest sea turtle species, to 120 cm (3.9 ft) long in the case of the green turtle, the second largest.[5][12]
The skulls of sea turtles have cheek regions that are enclosed in bone.[13][14] Although this condition appears to resemble that found in the earliest known fossil reptiles (anapsids), it is possible it is a more recently evolved trait in sea turtles, placing them outside the anapsids.[15][13]
Taxonomy and evolution
Sea turtles, along with other turtles and tortoises, are part of the order Testudines. All species except the leatherback sea turtle are in the family Cheloniidae. The superfamily name Chelonioidea and family name Cheloniidae are based on the Ancient Greek word for tortoise: χελώνη (khelone).[16] The leatherback sea turtle is the only extant member of the family Dermochelyidae.
Fossil evidence of marine turtles goes back to the
bothremydids, also survived well into the Cenozoic. Other pleurodires are also thought to have lived at sea, such as Araripemys[18] and extinct pelomedusids.[19] Modern sea turtles are not descended from more than one of the groups of sea-going turtles that have existed in the past; they instead constitute a single radiation that became distinct from all other turtles at least 110 million years ago.[20][21][22] Their closest extant relatives are in fact the snapping turtles (Chelydridae), musk turtles (Kinosternidae), and hickatee (Dermatemyidae) of the Americas, which alongside the sea turtles constitute the clade Americhelydia.[23]
The oldest possible representative of the lineage (
Desmatochelys padillaifrom the Early Cretaceous. Desmatochelys was a protostegid, a lineage that would later give rise to some very large species but went extinct at the end of the Cretaceous. Presently thought to be outside the crown group that contains modern sea turtles (Chelonioidea), the exact relationships of protostegids to modern sea turtles are still debated due to their primitive morphology; they may be the sister group to the Chelonoidea, or an unrelated turtle lineage that convergently evolved similar adaptations.[24][25] The earliest "true" sea turtle that is known from fossils is Nichollsemys from the Early Cretaceous (Albian) of Canada. In 2022, the giant fossil species Leviathanochelys was described from Spain. This species inhabited the oceans covering Europe in the Late Cretaceous and rivaled the concurrent giant protostegids such as Archelon and Protostega as one of the largest turtles to ever exist. Unlike the protostegids, which have an uncertain relationship to modern sea turtles, Leviathanochelys is thought to be a true sea turtle of the superfamily Chelonioidea.[26]
Sea turtles' limbs and brains have evolved to adapt to their diets. Their limbs originally evolved for locomotion, but more recently evolved to aid them in feeding. They use their limbs to hold, swipe, and forage their food. This helps them eat more efficiently.[27][28]
Cladogram
Below is a cladogram showing the phylogenetic relationships of living and extinct sea turtles in the Chelonioidea based on Evers et al. (2019):[29]
Phylogenetic relations of living and extinct chelonioid species
Sea turtles are generally found in the waters over continental shelves. During the first three to five years of life, sea turtles spend most of their time in the pelagic zone floating in seaweed mats. Green sea turtles in particular are often found in Sargassum mats, in which they find food, shelter and water.[31] Once the sea turtle has reached adulthood it moves closer to the shore.[32] Females will come ashore to lay their eggs on sandy beaches during the nesting season.[33]
Sea turtles migrate to reach their spawning beaches, which are limited in numbers. Living in the ocean therefore means they usually migrate over large distances. All sea turtles have large body sizes, which is helpful for moving large distances. Large body sizes also offer good protection against the large predators (notably sharks) found in the ocean.[34]
In 2020, diminished human activity resulting from the COVID-19 virus caused an increase in sea turtle nesting. Some areas in Thailand saw an abnormally high number of nests, and Florida experienced a similar phenomenon. Less plastic and light pollution could explain these observations.[35]
Life cycle
Sea turtles are thought to reach sexual maturity from about 10−20 years old depending on species and methodology. However, reliable estimates are difficult to ascertain.[36][37][38][39] Mature sea turtles may migrate thousands of miles to reach breeding sites. After mating at sea, adult female sea turtles return to land to lay their eggs. Different species of sea turtles exhibit various levels of philopatry. In the extreme case, females return to the same beach where they hatched. This can take place every two to four years in maturity.
The mature nesting female hauls herself onto the beach, nearly always at night, and finds suitable sand in which to create a nest. Using her hind flippers, she digs a circular hole 40 to 50 centimetres (16 to 20 in) deep. After the hole is dug, the female then starts filling the nest with her clutch of soft-shelled eggs. Depending on the species, a typical clutch may contain 50–350 eggs. After laying, she re-fills the nest with sand, re-sculpting and smoothing the surface, and then camouflaging the nest with vegetation until it is relatively undetectable visually.[31] She may also dig decoy nests.[40] The whole process takes 30 to 60 minutes. She then returns to the ocean, leaving the eggs untended.[41]
Females may lay 1–8 clutches in a single season. Female sea turtles alternate between mating in the water and laying their eggs on land. Most sea turtle species nest individually. But ridley sea turtles come ashore en masse, known as an arribada (arrival). With the Kemp's ridley sea turtle this occurs during the day.
Sea turtles have temperature-dependent sex determination, meaning the developing baby sea turtle's sex depends on the temperature it is exposed to.[42][43][44][45][46] Warmer temperatures produce female hatchlings, while cooler temperatures produce male hatchlings.[42][43][44][45][46][47] The eggs will incubate for 50–60 days. The eggs in one nest hatch together over a short period of time. The baby sea turtles break free of the egg shell, dig through the sand, and crawl into the sea. Most species of sea turtles hatch at night. However, the Kemp's ridley sea turtle commonly hatches during the day. Sea turtle nests that hatch during the day are more vulnerable to predators, and may encounter more human activity on the beach.
Larger hatchlings have a higher probability of survival than smaller individuals, which can be explained by the fact that larger offspring are faster and thus less exposed to predation. Predators can only functionally intake so much; larger individuals are not targeted as often. A study conducted on this topic shows that body size is positively correlated with speed, so larger baby sea turtles are exposed to predators for a shorter amount of time.
chelonians
has led to the evolutionary development of large body sizes.
In 1987, Carr discovered that the young of green and loggerhead sea turtles spent a great deal of their
pelagic waters. In the open ocean, pre-juveniles of this particular species were found to feed on zooplankton and smaller nekton before they are recruited into inshore seagrass meadows as obligate herbivores.[32][49]
Physiology
Osmoregulation
Sea turtles maintain an internal environment that is
lachrymal gland in the orbital cavity, capable of producing tears with a higher salt concentration than sea water.[52]
Leatherback sea turtles face an increased osmotic challenge compared to other species of sea turtle, since their primary prey are jellyfish and other gelatinous plankton, whose fluids have the same concentration of salts as sea water. The much larger lachrymal gland found in leatherback sea turtles may have evolved to cope with the higher intake of salts from their prey. A constant output of concentrated salty tears may be required to balance the input of salts from regular feeding, even considering leatherback sea turtle tears can have a salt ion concentration almost twice that of other species of sea turtle.[53]
Hatchlings depend on drinking sea water immediately upon entering the ocean to replenish water lost during the hatching process. Salt gland functioning begins quickly after hatching, so that the young sea turtles can establish ion and water balance soon after entering the ocean. Survival and physiological performance hinge on immediate and efficient hydration following emergence from the nest.[51]
Thermoregulation
All sea turtles are poikilotherms.[54] However, leatherback sea turtles (family Dermochelyidae) are able to maintain a body temperature 8 °C (14 °F) warmer than the ambient water by thermoregulation through the trait of gigantothermy.[54][55]
Green sea turtles in the relatively cooler Pacific are known to haul themselves out of the water on remote islands to bask in the sun.
Sea turtles are air-breathing reptiles that have lungs, so they regularly surface to breathe. Sea turtles spend a majority of their time underwater, so they must be able to hold their breath for long periods.[57] Dive duration largely depends on activity. A foraging sea turtle may typically spend 5–40 minutes underwater[57] while a sleeping sea turtle can remain underwater for 4–7 hours.[58][59] Remarkably, sea turtle respiration remains aerobic for the vast majority of voluntary dive time.[57][59] When a sea turtle is forcibly submerged (e.g. entangled in a trawl net) its diving endurance is substantially reduced, so it is more susceptible to drowning.[57]
When surfacing to breathe, a sea turtle can quickly refill its lungs with a single explosive exhalation and rapid inhalation. Their large lungs permit rapid exchange of oxygen and avoid trapping gases during deep dives.
Cold-stunning is a phenomenon that occurs when sea turtles enter cold ocean water (7–10 °C (45–50 °F)), which causes the turtles to float to the surface and therefore makes it impossible for them to swim.[60]
LED light and with a camera equipped with an orange optical filter to capture only the fluorescence light.[63][64]
Sensory modalities
Navigation
Below the surface, the sensory cues available for navigation change dramatically.[65] Light availability decreases quickly with depth, and is refracted by the movement of water when present, celestial cues are often obscured, and ocean currents cause continuous drift.[65] Most sea turtle species migrate over significant distances to nesting or foraging grounds, some even crossing entire ocean basins.[66] Passive drifting within major current systems, such as those in the North Atlantic Gyre, can result in ejection well outside of the temperature tolerance range of a given species, causing heat stress, hypothermia, or death.[66] In order to reliably navigate within strong gyre currents in the open ocean, migrating sea turtles possess both a bicoordinate magnetic map and magnetic compass sense, using a form of navigation termed Magnetoreception.[66][65][67] Specific migratory routes have been shown to vary between individuals, making the possession of both a magnetic map and compass sense advantageous for sea turtles.[66]
A bicoordinate magnetic map gives sea turtles the ability to determine their position relative to a goal with both latitudinal and longitudinal information, and requires the detection and interpretation of more than one magnetic parameter going in opposite directions to generate, such as
Inclination angle.[67][68] A magnetic compass sense allows sea turtles to determine and maintain a specific magnetic heading or orientation.[68] These magnetic senses are thought to be inherited, as hatchling sea turtles swim in directions that would keep them on course when exposed to the magnetic field signatures of various locations along their species' migratory routes.[68][69]
Natal homing behavior is well described in sea turtles, and genetic testing of turtle populations at different nesting sites has shown that magnetic field is a more reliable indicator of genetic similarity than physical distance between sites.[70] Additionally, nesting sites have been recorded to "drift" along with isoline shifts in the magnetic field.[71] Magnetoreception is thought to be the primary navigation tool used by nesting sea turtles in returning to natal beaches.[70][71] There are three major theories explaining natal site learning: inherited magnetic information, socially facilitated migration, and geomagnetic imprinting.[67] Some support has been found for geomagnetic imprinting, including successful experiments transplanting populations of sea turtles by relocating them prior to hatching, but the exact mechanism is still not known.[67]
Ecology
Diet
The loggerhead, Kemp's ridley, olive ridley, and hawksbill sea turtles are omnivorous their entire life. Omnivorous turtles may eat a wide variety of plant and animal life including decapods, seagrasses, seaweed, sponges, mollusks, cnidarians, Echinoderms, worms and fish.[72][73][74][75] However, some species specialize on certain prey.
The diet of green sea turtles changes with age.[76] Juveniles are omnivorous, but as they mature they become exclusively herbivorous.[73][76] This diet shift has an effect on the green sea turtle's morphology.[77][78] Green sea turtles have a serrated jaw that is used to eat sea grass and algae.[79]
Leatherback sea turtles feed almost exclusively on jellyfish and help control jellyfish populations.[80][81]
Hawksbill sea turtles principally eat sponges, which constitute 70–95% of their diets in the Caribbean.[82]
Larynx mechanisms
There was little information regarding the sea turtle's larynx. Sea turtles, like other turtle species, lack an epiglottis to cover the larynx entrance. Key findings from an experiment reveal the following in regards to the larynx morphology: a close apposition between the linguolaryngeal cleft's smooth mucosal walls and the laryngeal folds, a dorsal part of the glottis, the glottal mucosa attached to the arytenoid cartilage, and the way the hyoid sling is arranged and the relationship between the compressor laryngis muscle and cricoid cartilage. The glottal opening and closing mechanisms have been examined. During the opening stage, two abductor artytenoideae muscles swing arytenoid cartilages and the glottis walls. As a result, the glottis profile is transformed from a slit to a triangle. In the closing stage, the tongue is drawn posteriorly due to the close apposition of the glottis walls and linguolaryngeal cleft walls and hyoglossal sling contractions.[83]
Relationship with humans
Sea turtles are caught worldwide, although it is illegal to hunt most species in many countries.
Ancient Chinese texts dating to the 5th century B.C.E. describe sea turtles as exotic delicacies.[87] Many coastal communities around the world depend on sea turtles as a source of protein, often harvesting several sea turtles at once and keeping them alive on their backs until needed. Coastal peoples gather sea turtle eggs for consumption.[88]
To a much lesser extent, some species are targeted for their shells.
ancient Romans processed sea turtle scutes (primarily from the hawksbill sea turtle) for various articles and ornaments used by their elites, such as combs and brushes.[91] The skin of the flippers is prized for use as shoes and assorted leather goods.[92] In various West African countries, sea turtles are harvested for traditional medicinal use.[citation needed
]
The
Moche people of ancient Peru worshipped the sea and its animals. They often depicted sea turtles in their art.[93]J. R. R. Tolkien's poem "Fastitocalon" echoes a second-century Latin tale in the Physiologus of the Aspidochelone ("round-shielded turtle"); it is so large that sailors mistakenly land and light a fire on its back, and are drowned when it dives.[94][95]
Beach towns, such as Tortuguero, Costa Rica, have transitioned from a tourism industry that made profits from selling sea turtle meat and shells to an ecotourism-based economy. Tortuguero is considered to be the founding location of sea turtle conservation. In the 1960s the cultural demand for sea turtle meat, shells, and eggs was quickly killing the once-abundant sea turtle populations that nested on the beach. The Caribbean Conservation Corporation began working with villagers to promote ecotourism as a permanent substitute to sea turtle hunting. Sea turtle nesting grounds became sustainable. Tourists love to come and visit the nesting grounds, although it causes a lot of stress to the sea turtles because all of the eggs can get damaged or harmed.[96] Since the creation of a sea turtle ecotourism-based economy, Tortugero annually houses thousands of tourists who visit the protected 35-kilometre (22 mi) beach that hosts sea turtle walks and nesting grounds.[97][98] Walks to observe the nesting sea turtles require a certified guide and this controls and minimises disturbance of the beaches. It also gives the locals a financial interest in conservation and the guides now defend the sea turtles from threats such as poaching; efforts in Costa Rica's Pacific Coast are facilitated by a nonprofit organization, Sea Turtles Forever.[99] Thousands of people are involved in sea turtle walks, and substantial revenues accrue from the fees paid for the privilege.[100]
In other parts of the world where sea turtle breeding sites are threatened by human activity, volunteers often patrol beaches as a part of conservation activities, which may include relocating sea turtle eggs to hatcheries, or assisting hatching sea turtles in reaching the ocean.[101] Locations in which such efforts exist include the east coast of India,[102]São Tomé and Príncipe,[103]Sham Wan in Hong Kong,[104] and the coast of Florida.[105]
Importance to ecosystems
Sea turtles play key roles in two habitat types: oceans and beaches/dunes.
In the oceans, sea turtles, especially green sea turtles, are among the very few creatures (manatees are another) that eat
sea grass. Sea grass needs to be constantly cut short to help it grow across the sea floor. Sea turtle grazing helps maintain the health of the sea grass beds. Sea grass beds provide breeding and developmental grounds for numerous marine animals. Without them, many marine species humans harvest would be lost, as would the lower levels of the food chain. The reactions could result in many more marine species eventually becoming endangered or extinct.[106]
IUCN Red List classifies three species of sea turtle as either "endangered" or "critically endangered".[109] An additional three species are classified as "vulnerable".[109] The flatback sea turtle is considered as "data deficient", meaning that its conservation status is unclear due to lack of data.[109] All species of sea turtle are listed in CITES Appendix I, restricting international trade of sea turtles and sea turtle products.[4][110] However, the usefulness of global assessments for sea turtles has been questioned,[111] particularly due to the presence of distinct genetic stocks and spatially separated regional management units (RMUs).[112] Each RMU is subject to a unique set of threats that generally cross jurisdictional boundaries, resulting in some sub-populations of the same species' showing recovery while others continue to decline. This has triggered the IUCN to conduct threat assessments at the sub-population level for some species recently. These new assessments have highlighted an unexpected mismatch between where conservation relevant science has been conducted on sea turtles, and where there is the greatest need for conservation.[113] For example, as at August 2017, about 69% of studies using stable isotope analysis to understand the foraging distribution of sea turtles have been conducted in RMUs listed as "least concern" by the IUCN.[113]
Additionally, all populations of sea turtles that occur in United States waters are listed as threatened or endangered by the
US Endangered Species Act (ESA).[114] The US listing status of the loggerhead sea turtle is under review as of 2012.[114]
*The ESA manages sea turtles by population and not by species.
Management
In the Caribbean, researchers are having some success in assisting a comeback.[128] In September 2007, Corpus Christi, Texas, wildlife officials found 128 Kemp's ridley sea turtle nests on Texas beaches, a record number, including 81 on North Padre Island (Padre Island National Seashore) and four on Mustang Island. Wildlife officials released 10,594 Kemp's ridley sea turtle hatchlings along the Texas coast in recent years.
The
Pawikans) illegal. However, the law seems to have had little effect as sea turtle eggs are still in demand in Batangan markets. In September 2007, several Chinese poachers were apprehended off the Turtle Islands in the country's southernmost province of Tawi-Tawi. The poachers had collected more than a hundred sea turtles, along with 10,000 sea turtle eggs.[129]
Evaluating the progress of conservation programs is difficult, because many sea turtle populations have not been assessed adequately.[130] Most information on sea turtle populations comes from counting nests on beaches, but this does not provide an accurate picture of the whole sea turtle population.[131] A 2010 United States National Research Council report concluded that more detailed information on sea turtles' life cycles, such as birth rates and mortality, is needed.[132]
Nest relocation may not be a useful conservation technique for sea turtles. In one study on the freshwater Arrau turtle (
Podocnemis expansa) researchers examined the effects of nest relocation.[133] They discovered that clutches of this freshwater turtle that were transplanted to a new location had higher mortality rates and more morphological abnormalities compared to non-transplanted clutches.[133] However, in a study of loggerhead sea turtles (Caretta caretta), Dellert et al. found that relocating nests at risk of inundation increased the success of eggs and hatchlings and decreased the risk of inundation.[134]
Predators and disease
Most sea turtle mortality happens early in life. Sea turtles usually lay around 100 eggs at a time, but on average only one of the eggs from the nest will survive to adulthood.[135] Raccoons, foxes, and seabirds may raid nests or hatchlings may be eaten within minutes of hatching as they make their initial run for the ocean.[136] Once in the water, they are susceptible to seabirds, large fish and even other sea turtles.
Adult sea turtles have few predators. Large aquatic carnivores such as sharks and crocodiles are their biggest threats; however, reports of terrestrial predators attacking nesting females are not uncommon. Jaguars have been reported to smash into sea turtle shells with their paws, and scoop out the flesh.[137]
While many of the things that endanger sea turtles are natural predators,[136] increasingly many threats to the sea turtle species have arrived with the ever-growing presence of humans.[138]