Turtles are an
order
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
of
reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s known as Testudines, characterized by a special
shell
Shell may refer to:
Architecture and design
* Shell (structure), a thin structure
** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses
** Thin-shell structure
Science Biology
* Seashell, a hard ou ...
developed mainly from their ribs. Modern turtles are divided into two major groups, the
Pleurodira
The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differen ...
(side necked turtles) and
Cryptodira
The Cryptodira ('' el, hidden neck'') are a suborder of Testudines that includes most living tortoises and turtles. Cryptodira differ from Pleurodira (side-necked turtles) in that they lower their necks and pull the heads straight back into the ...
(hidden necked turtles), which differ in the way the head retracts. There are 360 living and recently extinct
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of turtles, including land-dwelling
tortoise
Tortoises () are reptiles of the family Testudinidae of the order Testudines (Latin: ''tortoise''). Like other turtles, tortoises have a turtle shell, shell to protect from predation and other threats. The shell in tortoises is generally hard, ...
s and freshwater
terrapin
Terrapins are one of several small species of turtle (order Testudines) living in fresh or brackish water. Terrapins do not form a taxonomic unit and may not be closely related. Many belong to the families Geoemydidae and Emydidae.
The name "t ...
s. They are found on most continents, some islands and, in the case of
sea turtle
Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhead, ...
s, much of the ocean. Like other
amniotes
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
(reptiles,
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s, and
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s) they breathe air and do not lay eggs underwater, although many species live in or around water.
Turtle shells are made mostly of
bone
A bone is a Stiffness, rigid Organ (biology), organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red blood cell, red and white blood cells, store minerals, provid ...
; the upper part is the domed
carapace
A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
, while the underside is the flatter
plastron or belly-plate. Its outer surface is covered in
scales
Scale or scales may refer to:
Mathematics
* Scale (descriptive set theory), an object defined on a set of points
* Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original
* Scale factor, a number w ...
made of
keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
, the material of hair, horns, and claws. The carapace bones develop from ribs that grow sideways and develop into broad flat plates that join up to cover the body. Turtles are
ectotherm
An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
s or "cold-blooded", meaning that their internal temperature varies with their direct environment. They are generally opportunistic
omnivore
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s and mainly feed on plants and animals with limited movements. Many turtles
migrate
Migration, migratory, or migrate may refer to: Human migration
* Human migration, physical movement by humans from one region to another
** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
short distances seasonally. Sea turtles are the only reptiles that migrate long distances to lay
their eggs on a favored beach.
Turtles have
appeared in myths and folktales around the world. Some terrestrial and freshwater species are widely kept as pets. Turtles have been
hunted for their meat, for use in traditional medicine, and for their shells. Sea turtles are often killed accidentally as
bycatch
Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
in fishing nets. Turtle habitats around the world are being destroyed. As a result of these pressures, many species are extinct or threatened with extinction.
Naming and etymology
The word ''turtle'' is derived from the
French
French (french: français(e), link=no) may refer to:
* Something of, from, or related to France
** French language, which originated in France, and its various dialects and accents
** French people, a nation and ethnic group identified with Franc ...
or ('turtle,
tortoise
Tortoises () are reptiles of the family Testudinidae of the order Testudines (Latin: ''tortoise''). Like other turtles, tortoises have a turtle shell, shell to protect from predation and other threats. The shell in tortoises is generally hard, ...
'). It is a
common name
In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
and may be used without knowledge of taxonomic distinctions. In North America, it may denote the order as a whole. In Britain, the name is used for
sea turtle
Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhead, ...
s as opposed to freshwater
terrapin
Terrapins are one of several small species of turtle (order Testudines) living in fresh or brackish water. Terrapins do not form a taxonomic unit and may not be closely related. Many belong to the families Geoemydidae and Emydidae.
The name "t ...
s and land-dwelling tortoises. In Australia, which lacks true tortoises (family Testudinidae), non-marine turtles were traditionally called tortoises, but more recently turtle has been used for the entire group.
The name of the order, ''Testudines'' ( ), is based on the
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word for tortoise, ; and was coined by German naturalist
August Batsch
August Johann Georg Karl Batsch (28 October 1761 – 29 September 1802) was a German naturalist. He was a recognised authority on mushrooms, and also described new species of ferns, bryophytes, and seed plants.
Life and career
Batsch was born ...
in 1788.
The order has also been historically known as ''Chelonii'' (
Latreille 1800) and ''Chelonia'' (Ross and
Macartney 1802),
which are based on the
Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
word for tortoise: (). Testudines is the official order name due to the
principle of priority
270px, '' valid name.
Priority is a fundamental principle of modern botanical nomenclature and zoological nomenclature. Essentially, it is the principle of recognising the first valid application of a name to a plant or animal. There are two asp ...
.
The term chelonian is used as a formal name for members of the group.
Anatomy and physiology
Size
The largest
living
Living or The Living may refer to:
Common meanings
*Life, a condition that distinguishes organisms from inorganic objects and dead organisms
** Living species, one that is not extinct
*Personal life, the course of an individual human's life
* Hu ...
species of turtle (and fourth-largest
reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
) is the
leatherback turtle
The leatherback sea turtle (''Dermochelys coriacea''), sometimes called the lute turtle or leathery turtle or simply the luth, is the largest of all living turtles and the heaviest non-crocodilian reptile, reaching lengths of up to and weights ...
, which can reach over in length and weigh over . The largest known turtle was ''
Archelon ischyros
''Archelon'' is an extinct marine turtle from the Late Cretaceous, and is the largest turtle ever to have been documented, with the biggest specimen measuring from head to tail and in body mass. It is known only from the Dakota Pierre Shal ...
'', a
Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
sea turtle up to long, wide between the tips of the front flippers, and estimated to have weighed over . The smallest living turtle is ''
Chersobius signatus
''Chersobius signatus'' is the world's smallest species of tortoise (family Testudinidae). The species is commonly known as the speckled tortoise and also known locally as the speckled padloper and internationally as the speckled Cape tortoise. ...
'' of South Africa, measuring no more than in length and weighing .
Shell
![Turtle skeleton cross-section, labelled as infographic](https://upload.wikimedia.org/wikipedia/commons/1/19/Turtle_skeleton_cross-section%2C_labelled_as_infographic.svg)
The shell of a turtle is unique among
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s and serves to protect the animal and provide shelter from the elements.
It is primarily made of 50–60 bones and consists of two parts: the domed, dorsal (back)
carapace
A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
and the flatter, ventral (belly)
plastron. They are connected by lateral (side) extensions of the plastron.
[
The carapace is fused with the vertebrae and ribs while the plastron is formed from bones of the ]shoulder girdle
The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of t ...
, sternum
The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sh ...
, and gastralia
Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these ...
(abdominal ribs).[ During development, the ribs grow sideways into a carapacial ridge, unique to turtles, entering the ]dermis
The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
(inner skin) of the back to support the carapace. The development is signaled locally by proteins known as fibroblast growth factor
Fibroblast growth factors (FGF) are a family of cell signalling proteins produced by macrophages; they are involved in a wide variety of processes, most notably as crucial elements for normal development in animal cells. Any irregularities in the ...
s that include FGF10
Fibroblast growth factor 10 is a protein that in humans is encoded by the ''FGF10'' gene.
Function
The protein encoded by this gene is a member of the fibroblast growth factor (FGF) family. FGF family members possess broad mitogenic and cell s ...
. The shoulder girdle in turtles is made up of two bones, the scapula and the coracoid
A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
. Both the shoulder and pelvic girdles of turtles are located within the shell and hence are effectively within the rib cage. The trunk ribs grow over the shoulder girdle during development.
The shell is covered in epidermal
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
(outer skin) scales known as scute
A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
s that are made of keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
, the same substance that makes up hair and fingernails. Typically, a turtle has 38 scutes on the carapace and 16 on the plastron, giving them 54 in total. Carapace scutes are divided into "marginals" around the margin and "vertebrals" over the vertebral column, though the scute that overlays the neck is called the "cervical". "Pleurals" are present between the marginals and vertebrals. Plastron scutes include gulars (throat), humerals, pectorals, abdominals, and anals. Side-necked turtles
The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differen ...
additionally have "intergular" scutes between the gulars. Turtle scutes are usually structured like mosaic
A mosaic is a pattern or image made of small regular or irregular pieces of colored stone, glass or ceramic, held in place by plaster/mortar, and covering a surface. Mosaics are often used as floor and wall decoration, and were particularly pop ...
tiles, but some species, like the hawksbill sea turtle
The hawksbill sea turtle (''Eretmochelys imbricata'') is a critically endangered sea turtle belonging to the family Cheloniidae. It is the only extant species in the genus ''Eretmochelys''. The species has a global distribution, that is largel ...
, have overlapping scutes on the carapace.
The shapes of turtle shells vary with the adaptations of the individual species, and sometimes with sex. Land-dwelling turtles are more dome-shaped, which appears to make them more resistant to being crushed by large animals. Aquatic turtles have flatter, smoother shells that allow them to cut through the water. Sea turtles in particular have streamlined shells that reduce drag and increase stability in the open ocean. Some turtle species have pointy or spiked shells that provide extra protection from predators and camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
against the leafy ground. The lumps of a tortoise shell can tilt its body when it gets flipped over, allowing it to flip back. In male tortoises, the tip of the plastron is thickened and used for butting and ramming during combat.
Shells vary in flexibility. Some species, such as box turtle
Box turtle is the common name for several species of turtle. It may refer to those of the genus '' Cuora'' or ''Pyxidea'', which are the Asian box turtles, or more commonly to species of the genus '' Terrapene'', the North American box turtles. T ...
s, lack the lateral extensions and instead have the carapace bones fully fused or ankylosed
Ankylosis is a stiffness of a joint due to abnormal adhesion and rigidity of the bones of the joint, which may be the result of injury or disease. The rigidity may be complete or partial and may be due to inflammation of the tendinous or muscula ...
together. Several species have hinges on their shells, usually on the plastron, which allow them to expand and contract. Softshell turtle
The Trionychidae are a taxonomic family of a number of turtle genera, commonly known as softshell turtles. The family was erected by Leopold Fitzinger in 1826. Softshells include some of the world's largest freshwater turtles, though many can ad ...
s have rubbery edges, due to the loss of bones. The leatherback turtle has hardly any bones in its shell, but has thick connective tissue
Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
and an outer layer of leathery skin.
Head and neck
The turtle's skull is unique among living amniote
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s (which includes reptiles, birds and mammals), it is solid and rigid with no openings for muscle attachment (temporal fenestrae
The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
). Muscles instead attach to recesses in the back of the skull. Turtle skulls vary in shape, from the long and narrow skulls of softshells to the broad and flattened skull of the mata mata
Matamata () is a town in Waikato, New Zealand. It is located near the base of the Kaimai Ranges, and is a thriving farming area known for Thoroughbred horse breeding and training pursuits. It is part of the Matamata-Piako District, which takes ...
. Some turtle species have developed large and thick heads, allowing for greater muscle mass and stronger bites.
Turtles that are carnivorous or durophagous
Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs. It is mostly used to describe fish, but is also used when describing reptiles, including fossil tu ...
(eating hard-shelled animals) have the most powerful bites. For example, the durophagous ''Mesoclemmys nasuta
''Mesoclemmys nasuta'' is a species of turtle that lives in Bolivia, Colombia, Peru, Brazil and the Guyanas (north-eastern South America).
A study found that this durophagous
Durophagy is the eating behavior of animals that consume hard-shelle ...
'' has a bite force of . Species that are insectivorous
A robber fly eating a hoverfly
An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects.
The first vertebrate insectivores were ...
, piscivorous
A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evoluti ...
(fish-eating), or omnivorous
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutri ...
have lower bite forces. Living turtles lack teeth but have beaks made of keratin sheaths along the edges of the jaws.[ These sheaths may have sharp edges for cutting meat, serrations for clipping plants, or broad plates for breaking ]mollusks
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is esti ...
.
The necks of turtles are highly flexible, possibly to compensate for their rigid shells. Some species, like sea turtles, have short necks while others, such as snake-necked turtles, have long ones. Despite this, all turtle species have eight neck vertebrae, a consistency not found in other reptiles but similar to mammals. Some snake-necked turtles have both long necks and large heads, limiting their ability to lift them when not in water. Some turtles have folded structures in the larynx
The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
or glottis
The glottis is the opening between the vocal folds (the rima glottidis). The glottis is crucial in producing vowels and voiced consonants.
Etymology
From Ancient Greek ''γλωττίς'' (glōttís), derived from ''γλῶττα'' (glôtta), va ...
that vibrate to produce sound. Other species have elastin
Elastin is a protein that in humans is encoded by the ''ELN'' gene. Elastin is a key component of the extracellular matrix in gnathostomes (jawed vertebrates). It is highly elastic and present in connective tissue allowing many tissues in the bod ...
-rich vocal cord
In humans, vocal cords, also known as vocal folds or voice reeds, are folds of throat tissues that are key in creating sounds through vocalization. The size of vocal cords affects the pitch of voice. Open when breathing and vibrating for speech ...
s.
Limbs and locomotion
Due to their heavy shells, turtles are slow-moving on land. A desert tortoise
The desert tortoise (''Gopherus agassizii''), is a species of tortoise in the Family (biology), family Testudinidae. The species is native to the Mojave Desert, Mojave and Sonoran Deserts of the southwestern United States and northwestern Mexic ...
moves at only . By contrast, sea turtles can swim at .[ The limbs of turtles are adapted for various means of locomotion and habits and most have five toes. Tortoises are specialized for terrestrial environments and have column-like legs with elephant-like feet and short toes. The ]gopher tortoise
The gopher tortoise (''Gopherus polyphemus'') is a species of tortoise in the family Testudinidae. The species is native to the southeastern United States. The gopher tortoise is seen as a keystone species because it digs burrows that provide ...
has flattened front limbs for digging in the substrate. Freshwater turtles have more flexible legs and longer toes with webbing
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, giving them thrust in the water. Some of these species, such as snapping turtles
The Chelydridae is a family of turtles that has seven extinct and two extant genera. The extant genera are the snapping turtles, ''Chelydra'' and '' Macrochelys''. Both are endemic to the Western Hemisphere. The extinct genera are '' Acherontemy ...
and mud turtle
''Kinosternon'' is a genus of small aquatic turtles from the Americas known commonly as mud turtles.
Geographic range
They are found in the United States, Mexico, Central America, and South America. The greatest species richness is in Mexico, a ...
s, mainly walk along the water bottom, as they would on land. Others, such as terrapins, swim by paddling with all four limbs, switching between the opposing front and hind limbs, which keeps their direction stable.[
]
Sea turtles and the pig-nosed turtle
The pig-nosed turtle (''Carettochelys insculpta''), also known as the pitted-shelled turtle or Fly River turtle, is a species of turtle native to northern Australia and southern New Guinea. It is the only living member of the family Carettoche ...
are the most specialized for swimming. Their front limbs have evolved into flippers while the shorter hind limbs are shaped more like rudders. The front limbs provide most of the thrust for swimming, while the hind limbs serve as stabilizers.[ Sea turtles such as the ]green sea turtle
The green sea turtle (''Chelonia mydas''), also known as the green turtle, black (sea) turtle or Pacific green turtle, is a species of large sea turtle of the family Cheloniidae. It is the only species in the genus ''Chelonia''. Its range exten ...
rotate the front limb flippers like a bird's wings to generate a propulsive force on both the upstroke and on the downstroke. This is in contrast to similar-sized freshwater turtles (measurements having been made on young animals in each case) such as the Caspian turtle
The Caspian turtle or striped-neck terrapin (''Mauremys caspica'') is a species of turtle in the family Geoemydidae (=Bataguridae), living in the eastern Mediterranean region from southwestern former USSR and central Iran to Saudi Arabia, Bahrai ...
, which uses the front limbs like the oars of a rowing boat, creating substantial negative thrust on the recovery stroke in each cycle. In addition, the streamlining of the marine turtles reduces drag. As a result, marine turtles produce a propulsive force twice as large, and swim six times as fast, as freshwater turtles. The swimming efficiency of young marine turtles is similar to that of fast-swimming fish of open water, like mackerel
Mackerel is a common name applied to a number of different species of pelagic fish, mostly from the family Scombridae. They are found in both temperate and tropical seas, mostly living along the coast or offshore in the oceanic environment.
...
.
Compared to other reptiles, turtles tend to have reduced tails, but these vary in both length and thickness among species and between sexes. Snapping turtles and the big-headed turtle
The big-headed turtle (''Platysternon megacephalum'') is a species of turtle in the family Platysternidae from Southeast Asia and southern China.
Background
Previously considered a distinct family placed on occasion in " Kinosternoidea", it w ...
have longer tails; the latter uses it for balance while climbing. The cloaca
In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
is found underneath and at the base, and the tail itself houses the reproductive organs. Hence, males have longer tails to contain the penis. In sea turtles, the tail is longer and more prehensile
Prehensility is the quality of an appendage or organ (anatomy), organ that has Adaptation (biology), adapted for grasping or holding. The word is derived from the Latin term ''prehendere'', meaning "to grasp". The ability to grasp is likely der ...
in males, who use it to grasp mates. Several turtle species have spines on their tails.
Senses
Turtles make use of vision
Vision, Visions, or The Vision may refer to:
Perception Optical perception
* Visual perception, the sense of sight
* Visual system, the physical mechanism of eyesight
* Computer vision, a field dealing with how computers can be made to gain un ...
to find food and mates, avoid predators, and orient themselves. The retina
The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
's light-sensitive cells include both rods for vision in low light, and cones
A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex.
A cone is formed by a set of line segments, half-lines, or lines conn ...
with three different photopigment
Photopigments are unstable pigments that undergo a chemical change when they absorb light. The term is generally applied to the non-protein chromophore moiety of photosensitive chromoproteins, such as the pigments involved in photosynthesis and p ...
s for bright light, where they have full-color vision. There is possibly a fourth type of cone that detects ultraviolet
Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
, as hatchling sea turtles respond experimentally to ultraviolet light, but it is unknown if they can distinguish this from longer wavelengths. A freshwater turtle, the red-eared slider
The red-eared slider or red-eared terrapin (''Trachemys scripta elegans'') is a subspecies of the pond slider (''Trachemys scripta''), a semiaquatic turtle belonging to the family Emydidae. It is the most popular pet turtle in the United States, ...
, has an exceptional seven types of cone cell.
Sea turtles orient themselves on land by night, using visual features detected in dim light. They can use their eyes in clear surface water, muddy coasts, the darkness of the deep ocean, and also above water. Unlike in terrestrial turtles, the cornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical power ...
, the curved surface that lets light into the eye, does not help to focus light on the retina, so focusing underwater is handled entirely by the lens, behind the cornea. The cone cells contain oil droplets placed to shift perception toward the red part of the spectrum, improving color discrimination. Visual acuity, studied in hatchlings, is highest in a horizontal band with retinal cells packed about twice as densely as elsewhere. This gives the best vision along the visual horizon. Sea turtles do not appear to use polarized light
Polarization (also polarisation) is a property applying to transverse waves that specifies the geometrical orientation of the oscillations. In a transverse wave, the direction of the oscillation is perpendicular to the direction of motion of the ...
for orientation as many other animals do. The deep-diving leatherback turtle lacks specific adaptations to low light, such as large eyes, large lenses, or a reflective tapetum. It may rely on seeing the bioluminescence
Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some b ...
of prey when hunting in deep water.
Turtles have no ear openings; the eardrum
In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear
The outer ear, external ear, or auris externa is the extern ...
is covered with scales and encircled by a bony otic capsule
The bony labyrinth (also osseous labyrinth or otic capsule) is the rigid, bony outer wall of the inner ear in the temporal bone. It consists of three parts: the vestibule, semicircular canals, and cochlea. These are cavities hollowed out of the s ...
, which is absent in other reptiles. Their hearing thresholds are high in comparison to other reptiles, reaching up to 500 Hz in air, but underwater they are more attuned to lower frequencies. The loggerhead sea turtle
The loggerhead sea turtle (''Caretta caretta'') is a species of oceanic turtle distributed throughout the world. It is a marine reptile, belonging to the family Cheloniidae. The average loggerhead measures around in carapace length when fully ...
has been shown experimentally to respond to low sounds, with maximal sensitivity between 100 and 400 Hz.
Turtles have olfactory
The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste.
In humans, it ...
(smell) and vomeronasal
The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. T ...
receptors along the nasal cavity, the latter of which are used to detect chemical signals. Experiments on green sea turtles showed they could learn to respond to a selection of different odorant chemicals such as triethylamine
Triethylamine is the chemical compound with the formula N(CH2CH3)3, commonly abbreviated Et3N. It is also abbreviated TEA, yet this abbreviation must be used carefully to avoid confusion with triethanolamine or tetraethylammonium, for which TEA ...
and cinnamaldehyde
Cinnamaldehyde is an organic compound with the formula(C9H8O) C6H5CH=CHCHO. Occurring naturally as predominantly the ''trans'' (''E'') isomer, it gives cinnamon its flavor and odor. It is a phenylpropanoid that is naturally synthesized by the shik ...
, which were detected by olfaction in the nose. Such signals could be used in navigation.
Breathing
The rigid shell of turtles is not capable of expanding and making room for the lungs, as in other amniotes, so they have had to evolve special adaptations for respiration. The lungs of turtles are attached directly to the carapace above while below, connective tissue attaches them to the organs. They have multiple lateral (side) and medial (middle) chambers (the numbers of which vary between species) and one terminal (end) chamber.
The lungs are ventilated using specific groups of abdominal muscles attached to the organs that pull and push on them. Specifically, it is the turtle's large liver that compresses the lungs. Underneath the lungs, in the coelomic cavity
The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...
, the liver is connected to the right lung by the root
In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...
, and the stomach is directly attached to the left lung, and to the liver by a mesentery
The mesentery is an organ that attaches the intestines to the posterior abdominal wall in humans and is formed by the double fold of peritoneum. It helps in storing fat and allowing blood vessels, lymphatics, and nerves to supply the intestines ...
. When the liver is pulled down, inhalation begins. Supporting the lungs is a wall or septum
In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate.
Examples
Human anatomy
* Interatri ...
, which is thought to prevent them from collapsing. During exhalation, the contraction of the transversus abdominis muscle
The transverse abdominal muscle (TVA), also known as the transverse abdominis, transversalis muscle and transversus abdominis muscle, is a muscle layer of the anterior and lateral (front and side) abdominal wall which is deep to (layered below) th ...
propels the organs into the lungs and expels air. Conversely, during inhalation, the relaxing and flattening of the oblique abdominis muscle pulls the transversus back down, allowing air back into the lungs.
Although many turtles spend large amounts of their lives underwater, all turtles breathe air and must surface at regular intervals to refill their lungs. Depending on the species, immersion periods vary between a minute and an hour. Some species can respire through the cloaca, which contains large sacs that are lined with many finger-like projections that take up dissolved oxygen
Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
from the water.
Circulation
Turtles share the linked circulatory
The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
and pulmonary (lung) systems of vertebrates, where the three-chambered heart pumps deoxygenated blood through the lungs and then pumps the returned oxygenated blood
Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the ci ...
through the body's tissues. The cardiopulmonary system has both structural and physiological adaptations that distinguish it from other vertebrates. Turtles have a large lung volume and can move blood through non-pulmonary blood vessels, including some within the heart, to avoid the lungs while they are not breathing. They can hold their breath for much longer periods than other reptiles and they can tolerate the resulting low oxygen levels. They can moderate the increase in acidity during anaerobic respiration, anaerobic (non-oxygen-based) respiration by buffer solution, chemical buffering and they can lie dormant for months, in aestivation or brumation.
The heart has two Atrium (heart), atria but only one Ventricle (heart), ventricle. The ventricle is subdivided into three chambers. A muscular ridge enables a complex pattern of blood flow so that the blood can be directed either to the lungs via the pulmonary artery, or to the body via the aorta. The ability to separate the two outflows varies between species. The leatherback has a powerful muscular ridge enabling almost complete separation of the outflows, supporting its actively swimming lifestyle. The ridge is less well developed in freshwater turtles like the sliders (''Trachemys'').
Turtles are capable of enduring periods of anaerobic respiration longer than many other vertebrates. This process breaks down sugars incompletely to lactic acid, rather than all the way to carbon dioxide and water as in aerobic respiration, aerobic (oxygen-based) respiration. They make use of the shell as a source of additional buffering agents for combating increased acidity, and as a sink for lactic acid.
Osmoregulation
In sea turtles, the bladder is one unit and in most freshwater turtles, it is double-lobed. Sea turtle bladders are connected to two small accessory bladders, located at the sides to the neck of the urinary bladder and above the Pubis (bone), pubis. Arid-living tortoises have bladders that serve as reserves of water, storing up to 20% of their body weight in fluids. The fluids are normally low in solutes, but higher during droughts when the reptile gains potassium salts from its plant diet. The bladder stores these salts until the tortoise finds fresh drinking water. To regulate the amount of salt in their bodies, sea turtles and diamondback terrapins secrete excess salt in a thick sticky substance from their lacrimal gland, tear glands. Because of this, sea turtles may appear to be "crying" when on land.
Thermoregulation
Turtles, like other reptiles, have a limited ability to Homoiotherm, regulate their body temperature. This ability varies between species, and with body size. Small pond turtles regulate their temperature by crawling out of the water and basking in the sun, while small terrestrial turtles move between sunny and shady places to adjust their temperature. Large species, both terrestrial and marine, have sufficient mass to give them substantial thermal inertia, meaning that they heat up or cool down over many hours. The Aldabra giant tortoise weighs up to some and is able to allow its temperature to rise to some on a hot day, and to fall naturally to around by night. Some giant tortoises seek out shade to avoid overheating on sunny days. On Grand Terre Island, food is scarce inland, shade is scarce near the coast, and the tortoises compete for space under the few trees on hot days. Large males may push smaller females out of the shade, and some then overheat and die.
Adult sea turtles, too, have large enough bodies that they can to some extent control their temperature. The largest turtle, the leatherback, can swim in the waters off Nova Scotia, which may be as cold as , while their body temperature has been measured at up to warmer than the surrounding water. To help keep their temperature up, they have a system of countercurrent heat exchange in the blood vessels between their body core and the skin of their flippers. The vessels supplying the head are insulated by fat around the neck.
Behavior
Diet and feeding
Most turtle species are opportunistic omnivores; land-dwelling species are more Herbivore, herbivorous and aquatic ones more Carnivore, carnivorous. Generally lacking speed and agility, most turtles feed either on plant material or on animals with limited movements like mollusks, worms, and insect larvae.[ Some species, such as the African helmeted turtle and snapping turtles, eat fish, amphibians, reptiles (including other turtles), birds, and mammals. They may take them by ambush predator, ambush but also scavenge. The alligator snapping turtle has a worm-like appendage on its tongue that it Aggressive mimicry, uses to lure fish into its mouth. Tortoises are the most herbivorous group, consuming grasses, leaves, and fruits. Many turtle species, including tortoises, supplement their diet with eggshells, animal bones, hair, and droppings for extra nutrients.
Turtles generally eat their food in a straightforward way, though some species have special feeding techniques.][ The yellow-spotted river turtle and the painted turtle may filter feed by skimming the water surface with their mouth and throat open to collect particles of food. When the mouth closes, the throat constricts and water is pushed out through the nostrils and the gap in between the jaws. Some species employ a "gape-and-suck method" where the turtle opens its jaws and expands its throat widely, sucking the prey in.]
The diet of an individual within a species may change with age, sex, and season, and may also differ between populations. In many species, juveniles are generally carnivorous but become more herbivorous as adults.[ With Barbour's map turtle, the larger female mainly eats mollusks while the male usually eats arthropods.][ Blanding's turtle may feed mainly on snails or crayfish depending on the population. The European pond turtle has been recorded as being mostly carnivorous much of the year but switching to Nymphaea alba, water lilies during the summer. Some species have developed Generalist and specialist species, specialized diets such as the hawksbill, which eats sponges, the leatherback, which feeds on jellyfish, and the Mekong snail-eating turtle.][
]
Communication and intelligence
While popularly thought of as mute, turtles make various sounds to communicate. Tortoises may bellow when courting and mating. Various species of both freshwater and sea turtles emit short, low-frequency calls from the time they are in the egg to when they are adults. These vocalizations may serve to create group cohesion when Animal migration, migrating. The oblong turtle has a particularly large vocal range; producing sounds described as clacks, clicks, squawks, hoots, various kinds of chirps, wails, ', grunts, growls, blow bursts, howls, and drum rolls.
Play behavior has been documented in some turtle species. In the laboratory, Florida red-bellied cooters can learn novel tasks and have demonstrated a long-term memory of at least 7.5 months. Similarly, giant tortoises can learn and remember tasks, and master lessons much faster when trained in groups. Tortoises appear to be able to retain operant conditioning nine years after their initial training. Studies have shown that turtles can navigate the environment using landmarks and a map-like system resulting in accurate direct routes towards a goal. Navigation in turtles have been correlated to high cognition function in the medial cortex region of the brain.
Defense
When sensing danger, a turtle may flee, freeze or withdraw into its shell. Freshwater turtles flee into the water, though the Sonora mud turtle may take refuge on land as the shallow temporary ponds they inhabit make them vulnerable. When startled, a softshell turtle may dive underwater and bury itself under the sea floor. If a predator persists, the turtle may bite or discharge from its cloaca. Several species produce foul-smelling chemicals from musk glands. Other tactics include threat displays and Bell's hinge-back tortoise can Apparent death, play dead. When attacked, big-headed turtle hatchlings squeal, possibly startling the predator.
Migration
Turtles are the only reptiles that migrate long distances, more specifically the marine species that can travel up to thousands of kilometers. Some non-marine turtles, such as the species of ''Geochelone'' (terrestrial), ''Chelydra'' (freshwater), and ''Malaclemys'' (estuarine), migrate seasonally over much shorter distances, up to around , to lay eggs. Such short migrations are comparable to those of some lizards, snakes, and crocodilians. Sea turtles nest in a specific area, such as a beach, leaving the eggs to hatch unattended. The young turtles leave that area, migrating long distances in the years or decades in which they grow to maturity, and then return seemingly to the same area every few years to mate and lay eggs, though the precision varies between species and populations. This "natal homing" has appeared remarkable to biologists, though there is now plentiful evidence for it, including from genetics.
How sea turtles navigate to their breeding beaches remains unknown. One possibility is Imprinting (psychology), imprinting as in salmon, where the young learn the chemical signature, effectively the scent, of their home waters before leaving, and remember that when the time comes for them to return as adults. Another possible cue is the orientation of the earth's magnetic field at the natal beach. There is experimental evidence that turtles have an effective magnetic sense, and that they use this in Animal navigation, navigation. Proof that homing occurs is derived from genetic analysis of populations of loggerheads, hawksbills, leatherbacks, and Olive ridley sea turtle, olive ridleys by nesting place. For each of these species, the populations in different places have their own mitochondrial DNA genetic signatures that persist over the years. This shows that the populations are distinct and that homing must be occurring reliably.
Reproduction and lifecycle
Turtles have a wide variety of mating behaviors but do not form Pair bond, pair-bonds or social groups. In green sea turtles, females generally outnumber males. In terrestrial species, males are often larger than females and fighting between males establishes a dominance hierarchy for access to mates. For most semi-aquatic and bottom-walking aquatic species, combat occurs less often. Males of these species instead may use their size advantage to Sexual coercion among animals, mate forcibly. In fully aquatic species, males are often smaller than females and rely on courtship displays to gain mating access to females.
Courtship and mounting
Courtship varies between species, and with habitat. It is often complex in aquatic species, both marine and freshwater, but simpler in the semi-aquatic mud turtles and snapping turtles. A male tortoise bobs his head, then subdues the female by biting and butting her before mounting.[ The male scorpion mud turtle approaches the female from the rear, and often resorts to aggressive methods such as biting the female's tail or hind limbs, followed by a mounting.]
Female choice is important in some species, and female green sea turtles are not always receptive. As such, they have evolved behaviors to avoid the male's attempts at copulation, such as swimming away, confronting the male followed by biting or taking up a refusal position with her body vertical, her limbs widely outspread, and her plastron facing the male. If the water is too shallow for the refusal position, the females resort to beaching themselves, as the males do not follow them ashore.
All turtles fertilize internally; mounting and copulation can be difficult. In many species, males have a concave plastron that interlocks with the female's carapace. In species like the Russian tortoise, the male has a lighter shell and longer legs. The high, rounded shape of box turtles are particular obstacles for mounting. The male eastern box turtle leans backward and hooks onto the back of the female's plastron. Aquatic turtles mount in water, and female sea turtles support the mounting male while swimming and diving. During copulation, the male turtle aligns his tail with the female's so he can insert his penis into her cloaca. Some female turtles can Female sperm storage, store sperm from multiple males and their Clutch (eggs), egg clutches can have multiple sires.
Eggs and hatchlings
Turtles, including sea turtles, lay their eggs on land, although some lay eggs close near water that rises and falls in level, submerging the eggs. While most species build nests and lay eggs where they forage, some travel miles. The common snapping turtle walks on land, while sea turtles travel even further; the leatherback swims some to its nesting beaches.[ Most turtles create a nest for their eggs. Females usually dig a flask-like chamber in the substrate. Other species lay their eggs in vegetation or crevices. Females choose nesting locations based on environmental factors such as temperature and humidity, which are important for developing embryos. Depending on the species, the number of eggs laid varies from one to over 100. Larger females can lay eggs that are greater in number or bigger in size. Compared to freshwater turtles, tortoises deposit fewer but larger eggs. Females can lay multiple clutches throughout a season, particularly in species that experience unpredictable monsoons.
]
Most mother turtles do no more in the way of parental care than covering their eggs and immediately leaving, though some species guard their nests for days or weeks. Eggs vary between rounded, oval, elongated, and between hard- and soft-shelled. Most species have their temperature-dependent sex determination, sex determined by temperature. In some species, higher temperatures produce females and lower ones produce males, while in others, milder temperatures produce males and both hot and cold extremes produce females.[ There is experimental evidence that the embryos of ''Mauremys reevesii'' can move around inside their eggs to select the best temperature for development, thus influencing their sexual destiny.] In other species, sex is Sex-determination system#Chromosomal systems, determined genetically. The length of incubation for turtle eggs varies from two to three months for temperate species, and four months to over a year for tropical species.[ Species that live in warm temperate climates can embryonic diapause, delay their development.
Hatching young turtles break out of the shell using an egg tooth, a sharp projection that exists temporarily on their upper beak.][ Hatchlings dig themselves out of the nest and find safety in vegetation or water. Some species stay in the nest for longer, be it for overwintering or to wait for the rain to loosen the soil for them to dig out.][ Young turtles are highly vulnerable to predators, both in the egg and as hatchlings. Mortality is high during this period but significantly decreases when they reach adulthood. Most species grow quickly during their early years and slow down when they are mature.
]
Lifespan
Turtles can live long lives. The oldest living turtle and land animal is said to be a Seychelles giant tortoise named Jonathan (tortoise), Jonathan, who turned 187 in 2019. A Galápagos tortoise named Harriet (tortoise), Harriet was collected by Charles Darwin in 1835; it died in 2006, having lived for at least 176 years. Most wild turtles do not reach that age. Turtles keep growing new scutes under the previous scutes every year, allowing researchers to estimate how long they have lived. They also negligible senescence, age slowly. The survival rate for adult turtles can reach 99% per year.[
]
Systematics and evolution
Fossil history
Zoologists have sought to explain the evolutionary origin of the turtles, and in particular of their unique shells. In 1914, Jan Versluys proposed that bony plates in the dermis, called osteoderms, fused to the ribs beneath them, later called the "Polka Dot Ancestor" by Olivier Rieppel. The theory accounted for the evolution of fossil pareiasaurs from ''Bradysaurus'' to ''Anthodon (reptile), Anthodon'', but not for how the ribs could have become attached to the bony dermal plates.
More recent discoveries have painted a different scenario for the evolution of the turtle's shell. The Crown group#Stem groups, stem-turtles ''Eunotosaurus'' of the Middle Permian, ''Pappochelys'' of the Middle Triassic, and ''Eorhynchochelys'' of the Late Triassic lacked carapaces and plastrons but had shortened torsos, expanded ribs, and lengthened dorsal vertebrae. Also in the Late Triassic, ''Odontochelys'' had a partial shell consisting of a complete bony plastron and an incomplete carapace. The development of a shell reached completion with the Late Triassic ''Proganochelys'', with its fully developed carapace and plastron. Adaptations that lead to the evolution of the shell may have originally been for digging and a fossorial lifestyle.
The oldest known members of the Pleurodira lineage are the Platychelyidae, from the Late Jurassic. The oldest known unambiguous cryptodire is ''Sinaspideretes,'' a close relative of softshell turtles, from the Late Jurassic of China. During the Late Cretaceous and Cenozoic, members of the pleurodire families Bothremydidae and Podocnemididae became widely distributed in the Northern Hemisphere due to their coastal habits. The oldest known soft-shelled turtles and sea turtles appeared during the Early Cretaceous. Tortoises originated in Asia during the Eocene. A late surviving group of stem-turtles, the Meiolaniidae, survived in Australasia into the Pleistocene and Holocene.
External relationships
The turtles' exact ancestry has been disputed. It was believed they were the only surviving branch of the ancient evolutionary grade Anapsida, which includes groups such as Procolophonidae, procolophonids and pareiasaurs. All anapsid skulls lack a temporal opening while all other living amniotes have temporal openings. It was later suggested that the anapsid-like turtle skulls may be due to backward evolution rather than to anapsid descent. Fossil evidence has shown that early stem-turtles possessed small temporal openings.
Some early morphological phylogenetics, phylogenetic studies have placed turtles closer to Lepidosauria (tuataras, lizards, and snakes) than to Archosauria (crocodilians and birds). By contrast, several Molecular phylogenetics, molecular studies place turtles either within Archosauria, or, more commonly, as a sister group to extant archosaurs, though an analysis conducted by Tyler Lyson and colleagues (2012) recovered turtles as the sister group of lepidosaurs instead. Ylenia Chiari and colleagues (2012) analyzed 248 nuclear genes from 16 vertebrates and suggested that turtles share a most recent common ancestor, more recent common ancestor with birds and crocodilians. The date of separation of turtles and birds and crocodilians was estimated to be during the Permian. Through genomic-scale phylogenetic study of ultra-conserved elements (UCEs) to clarify the placement of turtles within reptiles, Nicholas Crawford and colleagues (2012) similarly found that turtles are closer to birds and crocodilians.
Using the draft (unfinished) genome sequences of the green sea turtle and the Chinese softshell turtle, Zhuo Wang and colleagues (2013) concluded that turtles are likely a sister group of crocodilians and birds. The external phylogeny of the turtles is shown in the cladogram below.
Internal relationships
Modern turtles and their extinct relatives with a complete shell are classified within the clade Testudinata. The most recent common ancestor of living turtles, corresponding to the split between Pleurodira (side-necked species) and Cryptodira
The Cryptodira ('' el, hidden neck'') are a suborder of Testudines that includes most living tortoises and turtles. Cryptodira differ from Pleurodira (side-necked turtles) in that they lower their necks and pull the heads straight back into the ...
(hidden necked species), is estimated to have occurred around during the Late Triassic. Robert Thompson and colleagues (2021) comment that living turtles have low diversity, relative to how long they existed. Diversity has been stable, according to their analysis, except for a single rapid increase around the Eocene-Oligocene boundary some 30 million years ago, and a large regional extinction at roughly the same time. They suggest that global climate change caused both events, as the cooling and drying caused the land to become arid and turtles to become extinct there, while new continental margins opened up by the climate change provided habitats for other species to evolve.
The cladogram, from Nicholas Crawford and colleagues 2015, shows the internal phylogeny of the Testudines down to the level of Family (biology), families. The analysis by Thompson and colleagues in 2021 supports the same structure down to the family level.
Differences between the two suborders
Turtles are divided into two living suborders: Cryptodira and Pleurodira. The two groups differ in the way the neck is retracted for protection. Pleurodirans retract their neck to the side and in front of the shoulder girdles, whereas cryptodirans retract their neck backward into their shell. These motions are enabled by the morphology and arrangement of neck vertebrae. Sea turtles (which belong to Cryptodira) have mostly lost the ability to retract their heads.
The adductor muscles in the lower jaw create a pulley-like system in both subgroups. However, the bones that the muscles articulate with differ. In Pleurodira, the pulley is formed with the pterygoid bones of the palate, but in Cryptodira the pulley is formed with the otic capsule. Both systems help to vertically redirect the Anatomical terms of motion#Abduction and adduction, adductor muscles and maintain a powerful bite.
A further difference between the suborders is the attachment of the pelvis. In Cryptodira, the pelvis is free, linked to the shell only by ligaments. In Pleurodira, the pelvis is Suture (anatomy), sutured, joined with bony connections, to the carapace and to the plastron, creating a pair of large columns of bone at the back end of the turtle, linking the two parts of the shell.
Distribution and habitat
Turtles are widely distributed across the world's continents, oceans, and islands with terrestrial, fully aquatic, and semi-aquatic species. Sea turtles are mainly tropical and subtropical, but leatherbacks can be found in colder areas of the Atlantic and Pacific. Living Pleurodira all live in freshwater and are found only in the Southern Hemisphere. The Cryptodira include terrestrial, freshwater, and marine species, and these range more widely. The world regions richest in non-marine turtle species are the Amazon basin, the Gulf of Mexico Drainage basin, drainages of the United States, and parts of South and Southeast Asia.
For turtles in colder climates, their distribution is limited by constraints on reproduction, which is reduced by long hibernations. North American species barely range above the southern Canadian border. Some turtles are found at high altitudes, for example, the species ''Terrapene ornata'' occurs up to in New Mexico. Conversely, the leatherback sea turtle can dive over . Species of the genus ''Gopherus'' can tolerate both below freezing and over in body temperature, though they are most active at .
Conservation
Among vertebrate orders, turtles are second only to primates in the percentage of threatened species. 360 modern species have existed since 1500 AD. Of these, 51–56% are considered threatened and 60% considered threatened or extinct. Turtles face many threats, including habitat destruction, harvesting for consumption, the pet trade, light pollution, and climate change. Asian species have a particularly high extinction risk, primarily due to their long-term unsustainable exploitation for food and medicine, and about 83% of Asia's non-marine turtle species are considered threatened. As of 2021, turtle extinction is progressing much faster than during the Cretaceous-Tertiary extinction. At this rate, all turtles could be extinct in a few centuries.
Turtle hatchery, hatcheries can be set up when protection against flooding, erosion, predation, or heavy poaching is required. Chinese markets have sought to satisfy an increasing demand for turtle meat with farmed turtles. In 2007 it was estimated that over a thousand turtle farms operated in China. All the same, wild turtles continue to be caught and sent to market in large numbers, resulting in what conservationists have called "the Asian turtle crisis".[ In the words of the biologist George Amato, the hunting of turtles "vacuumed up entire species from areas in Southeast Asia", even as biologists still did not know how many species lived in the region. In 2000, all the Asian box turtles were placed on the CITES list of endangered species.][
Harvesting wild turtles is legal in some American states, and there has been a growing demand for American turtles in China.] The Florida Fish and Wildlife Conservation Commission estimated in 2008 that around 3,000 pounds of softshell turtles were exported weekly via Tampa International Airport. However, the great majority of turtles exported from the US between 2002 and 2005 were farmed.
Large numbers of sea turtles are accidentally killed in Longline fishing, longlines, gillnetting, gillnets, and trawling nets as bycatch
Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
. A 2010 study suggested that over 8 million had been killed between 1990 and 2008; the Eastern Pacific and the Mediterranean were identified as among the areas worst affected. Since the 1980s, the United States has required all Shrimp trawling, shrimp trawlers to fit their nets with turtle excluder devices that prevent turtles from being entangled in the net and drowning.
More locally, other human activities are affecting marine turtles. In Australia, Queensland's shark culling program, which uses shark nets and drum line (shark control), drum lines, has killed over 5,000 turtles as bycatch between 1962 and 2015; including 719 loggerhead turtles and 33 hawksbill sea turtles, which are listed as critically endangered.
Native turtle populations can also be threatened by Invasive species, invasive ones. The central North American red-eared slider turtle has been listed among the "100 of the World's Worst Invasive Alien Species, world's worst invasive species", pet turtle having been released globally. They appear to compete with native turtle species in eastern and western North America, Europe, and Japan.
Human uses
In culture
File:Kurmavatara (tortoise incarnation of Vishnu), from Garhwa, Allahabad District.jpg, alt=Photograph of temple sculpture in India, 4th-century sculpture of Kurma, turtle avatar of Vishnu. Garhwa, Garhwa, India
File:PSM V10 D562 The hindoo earth.jpg, alt=Lithograph drawing of world resting on 4 elephants standing on a giant turtle, World resting on four elephants on the back of the World Turtle. Western depiction of "The Hindu Earth", 1877
File:Kangxi-Lugou-rebuilding-stele-3581.jpg, alt=Chinese funeral stone held up by a stone tortoise, Bixi (tortoise), Bixi supporting Kangxi Emperor's stele, Beijing, 1698
File:Alice par John Tenniel 34.png, alt=Children's book illustration with turtle figure standing on hind legs, The Mock Turtle in Lewis Carroll's 1865 ''Alice's Adventures in Wonderland''
File:The Turtle 1898 Manhattan Theatre poster.jpg, alt=Painting of a turtle standing on hind legs, with top hat and cane, on theatre poster, Poster for 1898 production of ''The Turtle'' at the Manhattan Theatre, Broadway
File:Terrapin Shell Leg Rattles Worn by Lead Woman Dancer, Oconaluftee Village, NC. - NARA - 281630.jpg, alt=Photograph of cloth with four terrapin shell rattles to be tied around a dancer's leg, Terrapin shell leg rattles worn by lead Cherokee woman dancer, 20th century
Turtles have featured in human cultures across the world since ancient times. They are generally viewed positively despite not being "cuddly" or flashy; their association with the ancient times and old age have contributed to their endearing image.
In Hindu mythology, the World Turtle, named Kurma, Kurma or Kacchapa, supports four elephants on his back; they, in turn, carry the weight of the whole world on their backs. The turtle is one of the ten avatars or incarnations of the god Vishnu. The yoga pose Kurmasana is named for the avatar.[ revised from American Academy of Religions conference, San Francisco, 19 November 2011.] World Turtles are found in Native American cultures including the Algonquian peoples, Algonquian, Iroquois, and Lenape. They tell many versions of the creation story of Turtle Island (Native American folklore), Turtle Island. One version has Muskrat pile up earth on Turtle's back, creating the continent of North America. An Iroquois version has the pregnant Sky Woman fall through a hole in the sky between a tree's roots, where she is caught by birds who land her safely on Turtle's back; the Earth grows around her. The turtle here is altruistic, but the world is a heavy burden, and the turtle sometimes shakes itself to relieve the load, causing earthquakes.
A turtle was the symbol of the Ancient Mesopotamian god Enki from the 3rd millennium BCE onward. An ancient Greek origin myth told that only the tortoise refused the invitation of the gods Zeus and Hera to their wedding, as it preferred to stay at home. Zeus then ordered it to carry its house with it, ever after. Another of their gods, Hermes, invented a seven-stringed chelys, lyre made with the shell of a tortoise. In the Shang dynasty China, Chinese practice of plastromancy, dating back to 1200 BCE, oracles were obtained by inscribing questions on turtle plastrons using the Oracle bone script, oldest known form of Chinese characters, burning the plastron, and interpreting the resulting cracks. Later, the turtle was one of the Four Symbols, four sacred animals in Confucianism, while in the Han period, steles were mounted on top of stone turtles, later linked with Bixi, the turtle-shelled son of the Dragon King. Marine turtles feature significantly in Australian Aboriginal art. The army of Ancient Rome used the Testudo formation, ("tortoise") formation where soldiers would form a shield wall for protection.
In Aesop's Fables, "The Tortoise and the Hare" tells how an unequal race may be won by the slower partner. Lewis Carroll's 1865 ''Alice's Adventures in Wonderland'' features a Mock Turtle, named for Mock turtle soup, a soup meant to imitate the expensive soup made from real turtle meat. In 1896, the French playwright Léon Gandillot wrote a comedy in three acts named that was "a Parisian sensation" in its run in France, and came to the Manhattan Theatre, Broadway, New York, in 1898 as ''The Turtle''. A "cosmic turtle" and the island motif reappear in Gary Snyder's 1974 novel ''Turtle Island (book), Turtle Island'', and again in Terry Pratchett's ''Discworld'' series as Great A'Tuin, starting with the 1983 novel ''The Colour of Magic''. It is supposedly of the species ''Chelys galactica'', the galactic turtle, complete with four elephants on its back to support Discworld. Turtles have been featured in comic books and animations such as the 1984 Teenage Mutant Ninja Turtles.
As pets
Some turtles, particularly small terrestrial and freshwater species, are kept as pets. The demand for pet turtles increased in the 1950s, with the US being the main supplier, particularly of farm-bred red-eared sliders. The popularity for exotic pets has led to an increase in illegal Wildlife trade, wildlife trafficking. Around 21% of the value of live animal trade is in reptiles, and turtles are among the more popularly traded species. Poor husbandry of tortoises can cause chronic rhinitis (nasal swelling), overgrown beaks, hyperparathyroidism (which softens their skeleton), constipation, various reproductive problems, and injuries from dogs. In the early 20th century, people in the United States have organized and gambled on Turtle racing, turtle races.
As food and other uses
The flesh of captured wild turtles continues to be eaten in Asian cultures, while turtle soup was once a popular dish in English cuisine. Gopher tortoise stew has been popular with some groups in Florida. The supposed aphrodisiac or medicinal properties of turtle eggs created a large trade for them in Southeast Asia. Hard-shell turtle plastrons and soft-shell carapaces are widely used in traditional Chinese medicine; Taiwan imported nearly 200 metric tons of hard-shells from its neighbors yearly from 1999 to 2008. A popular medicinal preparation based on herbs and turtle shells is ''guilinggao'' jelly. The substance tortoiseshell, usually from the hawksbill turtle, has been used for centuries to make jewelry, tools, and ornaments around the Western Pacific. Hawksbills have accordingly been hunted for their shells. The trading of tortoiseshell was internationally banned in 1977 by CITES. Some cultures have used turtle shells to make music: Native American shamans made them into ceremonial rattles, while Aztecs, Maya civilization, Mayas, and Mixtecs made ' drums.
File:StateLibQld 2 395489 Catching turtles, wood engraving, 1875.jpg, Catching turtles in Australia, 1875, alt=Historic engraving of men catching turtles on a beach
File:TurtleSeafood.jpg, Turtles on sale as food in Canada, 2007, alt=photo of turtles on sale as food in a shop
File:Turtle plastrons as TCM in Xi'an market.jpg, Turtle plastrons for traditional Chinese medicine, alt=Photograph of a box of turtle plastrons in a market
File:Peigne en écaille.jpg, A tortoiseshell comb; the material was expensive and decorative, and widely used for small items., alt=Photograph of a decoratively ridged comb made of tortoiseshell
File:Sheldonbasking.JPG, A pet red-eared slider basking on a floating platform under a sun lamp, alt=Photograph of a pet turtle in a terrarium
See also
* World Turtle Day
References
Citations
Cited sources
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External links
Turtle Survival Alliance
Turtle Conservancy
Symposium on Turtle Evolution
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Turtles, Turtles
Articles containing video clips
Kimmeridgian first appearances
Extant Late Jurassic first appearances
Turtle taxonomy