Vertebrates () comprise all

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

within the

subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of "subdivision (rank), subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists ...

Vertebrata () (

chordates A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...

with backbones), including all

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 ...

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 ...

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 ( ...

amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...

s, and

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...

. Vertebrates represent the overwhelming majority of the

phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature f ...

Chordata A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...

, with currently about 69,963

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 ...

described. Vertebrates comprise such groups as the following: *

jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostomes ...

, which include

hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...

and

lampreys Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...

jawed vertebrates Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...

, which include: **

cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...

(

shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...

rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...

, and

ratfish Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At o ...

) ** bony vertebrates, which include: *** ray-fins (the majority of living

bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...

) ***

lobe-fins Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class (biology), class or subclass (biology), subclass) of the Osteichthyes, bony fishes known as the lobe-finned fishes. The group Tet ...

, which include: ****

coelacanths The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...

and

lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...

****

tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsids ...

(limbed vertebrates)

Extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

vertebrates range in size from the

frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...

species ''

Paedophryne amauensis ''Paedophryne amauensis'' is a species of microhylid frog endemic to eastern Papua New Guinea. At in snout-to-vent length, it is considered the world's smallest known vertebrate. (See also Ecological guild.) The species was listed in the ''T ...

'', at as little as , to the

blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can ...

, at up to . Vertebrates make up less than five percent of all described

animal 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 sex ...

; the rest are

invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...

, which lack

vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordata, ...

s. The vertebrates traditionally include the

, which do not have proper

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...

e due to their loss in evolution, though their closest living relatives, the

lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...

s, do. Hagfish do, however, possess a

cranium 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 ...

. For this reason, the vertebrate subphylum is sometimes referred to as "

Craniata A craniate is a member of the Craniata (sometimes called the Craniota), a proposed clade of chordate animals with a skull of hard bone or cartilage. Living representatives are the Myxini (hagfishes), Hyperoartia (including lampreys), and the muc ...

" when discussing morphology. Molecular analysis since 1992 has suggested that

are most closely related to

s, and so also are vertebrates in a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

sense. Others consider them a sister group of vertebrates in the common taxon of craniata.

Etymology

The word ''vertebrate'' derives from 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 ''vertebratus'' (

Pliny Pliny may refer to: People * Pliny the Elder (23–79 CE), ancient Roman nobleman, scientist, historian, and author of ''Naturalis Historia'' (''Pliny's Natural History'') * Pliny the Younger (died 113), ancient Roman statesman, orator, w ...

), meaning ''joint of the spine.'' ''Vertebrate'' is derived from the word ''

'', which refers to any of the bones or segments of the

spinal column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordates ...

Anatomy and morphology

All vertebrates are built along the basic chordate

body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...

: a stiff rod running through the length of the animal (vertebral column and/or

notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consis ...

), with a hollow tube of nervous tissue (the

spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spi ...

) above it and the

gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organ (biology), organs of the digestive syste ...

below. In all vertebrates, the mouth is found at, or right below, the anterior end of the animal, while the

anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...

opens to the exterior before the end of the body. The remaining part of the body continuing after the anus forms a

tail The tail is the section at the rear end of certain kinds of animals’ bodies; in general, the term refers to a distinct, flexible appendage to the torso. It is the part of the body that corresponds roughly to the sacrum and coccyx in mammals, r ...

with vertebrae and spinal cord, but no gut.Romer, A.S. (1949): ''The Vertebrate Body.'' W.B. Saunders, Philadelphia. (2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970)

Vertebral column

The defining characteristic of a vertebrate is the

, in which the

(a stiff rod of uniform composition) found in all

has been replaced by a segmented series of stiffer elements (vertebrae) separated by mobile joints (intervertebral discs, derived embryonically and evolutionarily from the notochord). However, a few vertebrates have secondarily lost this anatomy, retaining the notochord into adulthood, such as the

sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...

and

coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...

Jawed vertebrates Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...

are typified by paired appendages (fins or legs, which may be secondarily lost), but this trait is not required in order for an animal to be a vertebrate. Naturkundemuseum Berlin - Dinosaurierhalle

Naturkundemuseum Berlin - Dinosaurierhalle

Gills

All basal vertebrates breathe with

gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...

. The gills are carried right behind the head, bordering the posterior margins of a series of openings from the

pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...

to the exterior. Each

gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...

is supported by a cartilaginous or bony

gill arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arc ...

. The

have three pairs of arches,

have five to seven pairs, while the primitive

have seven. The vertebrate ancestor no doubt had more arches than this, as some of their

chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...

relatives have more than 50 pairs of gills. In

s and some primitive bony

fishes Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...

, the

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

e bear

external gills External gills are the gills of an animal, most typically an amphibian, that are exposed to the environment, rather than set inside the pharynx and covered by gill slits, as they are in most fishes. Instead, the respiratory organs are set on a frill ...

, branching off from the gill arches. These are reduced in adulthood, their function taken over by the gills proper in fishes and by

lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...

s in most amphibians. Some amphibians retain the external larval gills in adulthood, the complex internal

system as seen in fish apparently being irrevocably lost very early in the evolution of

tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...

s.Clack, J. A. (2002): Gaining ground: the origin and evolution of tetrapods. ''

Indiana University Press Indiana University Press, also known as IU Press, is an academic publisher founded in 1950 at Indiana University that specializes in the humanities and social sciences. Its headquarters are located in Bloomington, Indiana. IU Press publishes 140 ...

'', Bloomington, Indiana. 369 pp While the more derived vertebrates lack gills, the gill arches form during

fetal development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...

, and form the basis of essential structures such as

jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...

s, the

thyroid gland The thyroid, or thyroid gland, is an endocrine gland in vertebrates. In humans it is in the neck and consists of two connected lobe (anatomy), lobes. The lower two thirds of the lobes are connected by a thin band of Connective tissue, tissue cal ...

, 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 ...

, the ''columella'' (corresponding to the

stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foot ...

s) and, in mammals, the malleus and incus.

Central nervous system

The

central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all par ...

of vertebrates is based on a hollow nerve cord running along the length of the animal. Of particular importance and unique to vertebrates is the presence of

neural crest Neural crest cells are a temporary group of cells unique to vertebrates that arise from the embryonic ectoderm germ layer, and in turn give rise to a diverse cell lineage—including melanocytes, craniofacial cartilage and bone, smooth muscle, per ...

cells. These are progenitors of

stem cell In multicellular organisms, stem cells are undifferentiated or partially differentiated cells that can differentiate into various types of cells and proliferate indefinitely to produce more of the same stem cell. They are the earliest type o ...

s, and critical to coordinating the functions of cellular components.Teng, L.; Labosky, P. A. (2006)
"Neural crest stem cells"
In: Jean-Pierre Saint-Jeannet, ''Neural Crest Induction and Differentiation'', pp. 206-212, Springer Science & Business Media. . Neural crest cells migrate through the body from the nerve cord during development, and initiate the formation of neural ganglia and structures such as the jaws and skull. The vertebrates are the only

group with neural

cephalisation Cephalization is an evolutionary trend in which, over many generations, the mouth, sense organs, and nerve ganglia become concentrated at the front end of an animal, producing a head region. This is associated with movement and bilateral symmetr ...

, the concentration of

brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...

functions in the head. A slight swelling of the anterior end of the nerve cord is found in the

lancelet The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...

, a chordate, though it lacks the eyes and other complex sense

organs In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a f ...

comparable to those of vertebrates. Other chordates do not show any trends towards cephalisation. A

peripheral nervous system The peripheral nervous system (PNS) is one of two components that make up the nervous system of bilateral animals, with the other part being the central nervous system (CNS). The PNS consists of nerves and ganglia, which lie outside the brain ...

branches out from the nerve cord to innervate the various systems. The front end of the nerve tube is expanded by a thickening of the walls and expansion of the central canal of spinal cord into three primary brain vesicles: The

prosencephalon In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the Anatomical terms of location#Directional terms, rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hin ...

(forebrain),

mesencephalon The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', "m ...

(midbrain) and

rhombencephalon The hindbrain or rhombencephalon or lower brain is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes. Metencephalon ...

(hindbrain), further differentiated in the various vertebrate groups.Hildebrand, M.; Gonslow, G. (2001): Analysis of Vertebrate Structure. 5th edition. ''John Wiley & Sons, Inc''.

New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...

Two laterally placed eyes form around outgrowths from the midbrain, except in

, though this may be a secondary loss. The forebrain is well-developed and subdivided in most

s, while the midbrain dominates in many

and some

salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...

s. Vesicles of the forebrain are usually paired, giving rise to hemispheres like the

cerebral hemisphere The vertebrate cerebrum (brain) is formed by two cerebral hemispheres that are separated by a groove, the longitudinal fissure. The brain can thus be described as being divided into left and right cerebral hemispheres. Each of these hemispheres ...

s in

s. The resulting anatomy of the central nervous system, with a single hollow nerve cord topped by a series of (often paired) vesicles, is unique to vertebrates. All

with well-developed brains, such as

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...

spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...

s and

squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...

s, have a ventral rather than dorsal system of

ganglion A ganglion is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system there are both sympatheti ...

s, with a split

brain stem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is conti ...

running on each side of the mouth or gut.

Molecular signatures

In addition to the morphological characteristics used to define vertebrates (i.e. the presence of a notochord, the development of a vertebral column from the notochord, a dorsal nerve cord, pharyngeal gills, a post-anal tail, etc.), molecular markers known as

conserved signature indels Conserved signature inserts and deletions (CSIs) in protein sequences provide an important category of molecular markers for understanding phylogenetic relationships. CSIs, brought about by rare genetic changes, provide useful phylogenetic markers ...

(CSIs) in protein sequences have been identified and provide distinguishing criteria for the subphylum Vertebrata. Specifically, 5 CSIs in the following proteins: protein synthesis

elongation factor-2 Eukaryotic elongation factor 2 is a protein that in humans is encoded by the ''EEF2'' gene. It is the archaeal and eukaryotic counterpart of bacterial EF-G. This gene encodes a member of the Guanosine triphosphate, GTP-binding translation elongat ...

(EF-2),

eukaryotic translation initiation factor 3 Eukaryotic initiation factor 3 (eIF3) is a multiprotein complex that functions during the initiation phase of eukaryotic translation. It is essential for most forms of cap-dependent and cap-independent translation initiation. In humans, eIF3 co ...

(eIF3),

adenosine kinase Adenosine kinase (AdK; EC 2.7.1.20) is an enzyme that catalyzes the transfer of gamma-phosphate from Adenosine triphosphate ( ATP) to adenosine (Ado) leading to formation of Adenosine monophosphate (AMP). In addition to its well-studied role in co ...

(AdK) and a protein related to

ubiquitin carboxyl-terminal hydrolase Ubiquitin C-terminal hydrolase may refer to: * Ubiquitin carboxy-terminal hydrolase L1, an enzyme * Ubiquitinyl hydrolase 1, an enzyme {{Short pages monitor A

dagger A dagger is a fighting knife with a very sharp point and usually two sharp edges, typically designed or capable of being used as a thrusting or stabbing weapon.State v. Martin, 633 S.W.2d 80 (Mo. 1982): This is the dictionary or popular-use de ...

(†) denotes an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

, whereas all other clades have living descendants. NochelaspisMaeandrine-PaleozoologicalMuseumOfChina-May23-08

Note that, as shown in the cladogram above, the †"

Ostracodermi Ostracoderms () are the Armour (zoology), armored Agnatha, jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are cl ...

" (armoured jawless fishes) and †"

Placodermi Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally 'Plate (animal anatomy), plate-skinned') is a Class (biology), class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devoni ...

" (armoured jawed fishes) are shown to be paraphylectic groups, separated from Gnathostomes and Eugnathostomes respectively. Also note that

Teleostei Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest class (biology), infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of a ...

(

Neopterygii Neopterygii (from Greek νέος ''neos'' 'new' and πτέρυξ ''pteryx'' 'fin') is a subclass of ray-finned fish (Actinopterygii). Neopterygii includes the Holostei and the Teleostei, of which the latter comprise the vast majority of extant f ...

) and Tetrapoda (

s) each make up about 50% of today's vertebrate

diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...

, while all other groups are either

or rare. The next

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

shows the extant

clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...

of tetrapods (the four-limbed vertebrates), and a selection of

(†) groups: Note that reptile-like amphibians,

mammal-like reptile Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is no ...

s, and non-avian dinosaurs are all

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

. The placement of hagfish on the vertebrate tree of life has been controversial. Their lack of proper

vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...

(among with other characteristics found in lampreys and jawed vertebrates) led phylogenetic analyses based on

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

to place them outside Vertebrata.

Molecular data In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which succi ...

, however, indicates they are vertebrates closely related to

. A study by Miyashita ''et al''. (2019), 'reconciliated' the two types of analysis as it supports the

Cyclostomata Cyclostomi, often referred to as Cyclostomata , is a group of vertebrates that comprises the living jawless fishes: the lampreys and hagfishes. Both groups have jawless mouths with horny epidermal structures that function as teeth called ceratod ...

hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...

using only morphological data.

Number of extant species

The number of described vertebrate species are split between

s and

. The following table lists the number of described

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

species for each vertebrate

class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...

as estimated in the

IUCN Red List of Threatened Species The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biologi ...

, 2014.3.The World Conservation Union. 2014. ''

'', 2014.3. Summary Statistics for Globally Threatened Species
Table 1: Numbers of threatened species by major groups of organisms (1996–2014)
The IUCN estimates that 1,305,075 extant invertebrate species have been described, which means that less than 5% of the described animal species in the world are vertebrates.

Vertebrate species databases

The following databases maintain (more or less) up-to-date lists of vertebrate species: * Fish:

Fishbase FishBase is a global species database of fish species (specifically finfish). It is the largest and most extensively accessed online database on adult finfish on the web.

* Amphibians
Amphibiaweb
* Reptiles:

Reptile Database The Reptile Database is a scientific database that collects taxonomic information on all living reptile species (i.e. no fossil species such as dinosaurs). The database focuses on species (as opposed to higher ranks such as families) and has entrie ...

* Birds
Avibase
* Mammals
Mammal species of the World

Reproductive systems

Nearly all vertebrates undergo

sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...

. They produce

haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...

gametes by

meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...

. The smaller, motile gametes are

spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromosomes, ...

and the larger, non-motile gametes are

ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...

. These fuse by the process of

fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...

to form

diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...

zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicellula ...

s, which develop into new individuals.

Inbreeding

During sexual reproduction, mating with a close relative (

inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and o ...

) often leads to

inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...

. Inbreeding depression is considered to be largely due to expression of deleterious

recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...

mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...

s. The effects of inbreeding have been studied in many vertebrate species. In several species of fish, inbreeding was found to decrease reproductive success. Inbreeding was observed to increase juvenile mortality in 11 small animal species. A common breeding practice for pet dogs is mating between close relatives (e.g. between half- and full

sibling A sibling is a relative that shares at least one parent with the subject. A male sibling is a brother and a female sibling is a sister. A person with no siblings is an only child. While some circumstances can cause siblings to be raised separat ...

s). This practice generally has a negative effect on measures of reproductive success, including decreased litter size and puppy survival.

Incest Incest ( ) is human sexual activity between family members or close relatives. This typically includes sexual activity between people in consanguinity (blood relations), and sometimes those related by affinity (marriage or stepfamily), adoption ...

uous matings in birds result in severe fitness costs due to inbreeding depression (e.g. reduction in hatchability of eggs and reduced progeny survival).

Inbreeding avoidance

As a result of the negative fitness consequences of

, vertebrate species have evolved mechanisms to avoid inbreeding. Numerous inbreeding avoidance mechanisms operating prior to mating have been described. Toads and many other amphibians display breeding site fidelity. Individuals that return to natal ponds to breed will likely encounter

s as potential mates. Although

incest Incest ( ) is human sexual activity between family members or close relatives. This typically includes sexual activity between people in consanguinity (blood relations), and sometimes those related by affinity (marriage or stepfamily), adoption ...

is possible, ''Bufo americanus'' siblings rarely mate. These toads likely recognize and actively avoid close kin as mates.

Advertisement Advertising is the practice and techniques employed to bring attention to a product or service. Advertising aims to put a product or service in the spotlight in hopes of drawing it attention from consumers. It is typically used to promote a ...

vocalizations by males appear to serve as cues by which females recognize their kin.

Inbreeding avoidance Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. Animals only rarely exhibit inbreeding avoidance. The inbreeding avoidance ...

mechanisms can also operate subsequent to

copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetrat ...

. In guppies, a post-copulatory mechanism of inbreeding avoidance occurs based on competition between

sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...

of rival males for achieving

fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...

. In competitions between sperm from an unrelated male and from a full sibling male, a significant bias in paternity towards the unrelated male was observed. When female

sand lizard The sand lizard (''Lacerta agilis'') is a lacertid lizard distributed across most of Europe from France and across the continent to Lake Baikal in Russia. It does not occur in European Turkey. Its distribution is often patchy. In the sand lizard ...

s mate with two or more males, sperm competition within the female's reproductive tract may occur. Active selection of sperm by females appears to occur in a manner that enhances female fitness. On the basis of this selective process, the sperm of males that are more distantly related to the female are preferentially used for fertilization, rather than the

of close relatives. This preference may enhance the fitness of progeny by reducing

Outcrossing

Mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...

with unrelated or distantly related members of the same species is generally thought to provide the advantage of masking deleterious recessive mutations in progeny (see

heterosis Heterosis, hybrid vigor, or outbreeding enhancement is the improved or increased function of any biological quality in a hybrid offspring. An offspring is heterotic if its traits are enhanced as a result of mixing the genetic contributions of ...

). Vertebrates have evolved numerous diverse mechanisms for avoiding close inbreeding and promoting outcrossing (see

inbreeding avoidance Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. Animals only rarely exhibit inbreeding avoidance. The inbreeding avoidance ...

). Outcrossing as a way of avoiding inbreeding depression has been especially well studied in birds. For instance, inbreeding depression occurs in the

great tit The great tit (''Parus major'') is a passerine bird in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central Asia and east across the Palearctic to the Amur River, south to parts of North Af ...

(''Parus major'') when the offspring are produced as a result of a mating between close relatives. In natural populations of the great tit, inbreeding is avoided by dispersal of individuals from their birthplace, which reduces the chance of mating with a close relative.

Purple-crowned fairywren The purple-crowned fairywren (''Malurus coronatus'') is a species of bird in the Australasian wren family, Maluridae. It is the largest of the eleven species in the genus ''Malurus'' and is endemic to northern Australia. The species name is de ...

females paired with related males may undertake extra-pair matings that can reduce the negative effects of inbreeding, despite ecological and demographic constraints.

Southern pied babbler The southern pied babbler (''Turdoides bicolor'') is a species of bird in the family Leiothrichidae, found in dry savannah of Botswana, Namibia, South Africa, and Zimbabwe. Description The southern pied babbler is a medium-sized cooperatively b ...

s (''Turdoides bicolor'') appear to avoid inbreeding in two ways: through dispersal and by avoiding familiar group members as mates. Although both males and females disperse locally, they move outside the range where genetically related individuals are likely to be encountered. Within their group, individuals only acquire breeding positions when the opposite-sex breeder is unrelated. Cooperative breeding in birds typically occurs when

offspring In biology, offspring are the young creation of living organisms, produced either by a single organism or, in the case of sexual reproduction, two organisms. Collective offspring may be known as a brood or progeny in a more general way. This ca ...

, usually males, delay dispersal from their natal group in order to remain with the family to help rear younger kin. Female offspring rarely stay at home, dispersing over distances that allow them to breed independently or to join unrelated groups.

Parthenogenesis

Parthenogenesis is a natural form of reproduction in which growth and development of embryos occur without fertilization. Reproduction in

squamate Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest order of reptiles, comprising lizards, snakes, and amphisbaenians (worm lizards), which are collectively known as squamates or scaled reptiles. With over 10,900 species, ...

reptiles is ordinarily sexual, with males having a ZZ pair of sex determining chromosomes, and females a ZW pair. However, various species, including the Colombian Rainbow boa (''Epicrates maurus''), ''

Agkistrodon contortrix The eastern copperhead (''Agkistrodon contortrix''), also known as the copperhead, is a species of venomous snake, a pit viper, endemic to eastern North America; it is a member of the subfamily Crotalinae in the family Viperidae. The eastern ...

'' (copperhead snake) and ''

Agkistrodon piscivorus ''Agkistrodon piscivorus'' is a species of pit viper in the subfamily Crotalinae of the family Viperidae. It is one of the world's few semiaquatic vipers (along with the Florida cottonmouth), and is native to the southeastern United States. A ...

'' (cotton mouth snake) can also reproduce by facultative parthenogenesis—that is, they are capable of switching from a sexual mode of reproduction to an asexual mode—resulting in production of WW female progeny. The WW females are likely produced by terminal automixis.

Mole salamander The mole salamanders (genus ''Ambystoma'') are a group of advanced salamanders endemic to North America. The group has become famous due to the presence of the axolotl (''A. mexicanum''), widely used in research due to its paedomorphosis, and t ...

s are an ancient (2.4–3.8 million year-old) unisexual vertebrate lineage. In the

polyploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...

unisexual mole salamander females, a premeiotic endomitotic event doubles the number of chromosomes. As a result, the mature eggs produced subsequent to the two meiotic divisions have the same ploidy as the somatic cells of the female salamander.

Synapsis Synapsis is the pairing of two chromosomes that occurs during meiosis. It allows matching-up of homologous pairs prior to their segregation, and possible chromosomal crossover between them. Synapsis takes place during prophase I of meiosis. Whe ...

and recombination during meiotic prophase I in these unisexual females is thought to ordinarily occur between identical sister chromosomes and occasionally between

homologous chromosome A couple of homologous chromosomes, or homologs, are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during fertilization. Homologs have the same genes in the same loci where they provide points alon ...

s. Thus little, if any, genetic variation is produced. Recombination between homeologous chromosomes occurs only rarely, if at all. Since production of genetic variation is weak, at best, it is unlikely to provide a benefit sufficient to account for the long-term maintenance of meiosis in these organisms.

Self-fertilization

Two

killifish A killifish is any of various oviparous (egg-laying) cyprinodontiform fish (including families Aplocheilidae, Cyprinodontidae, Fundulidae, Profundulidae and Valenciidae). All together, there are 1,270 species of killifish, the biggest family ...

species, the

mangrove killifish The mangrove killifish or mangrove rivulus, ''Kryptolebias marmoratus'' (syn. ''Rivulus marmoratus''), is a species of killifish in the Family (biology), family Rivulidae. It lives in brackish and marine waters (less frequently in fresh water) a ...

(''Kryptolebias marmoratus'') and '' Kryptolebias hermaphroditus'', are the only known vertebrates to self-fertilize. They produce both eggs and sperm by meiosis and routinely reproduce by

self-fertilisation Autogamy, or self-fertilization, refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species of ...

. This capacity has apparently persisted for at least several hundred thousand years. Each individual

hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...

normally fertilizes itself through uniting inside the fish's body of an egg and a sperm that it has produced by an internal organ. In nature, this mode of reproduction can yield highly homozygous lines composed of individuals so genetically uniform as to be, in effect, identical to one another. Although inbreeding, especially in the extreme form of self-fertilization, is ordinarily regarded as detrimental because it leads to expression of deleterious recessive alleles, self-fertilization does provide the benefit of ''fertilization assurance'' ( reproductive assurance) at each generation.

Population trends

The Living Planet Index, following 16,704 populations of 4,005 species of vertebrates, shows a decline of 60% between 1970 and 2014. Since 1970, freshwater species declined 83%, and tropical populations in South and Central America declined 89%. The authors note that, "An average trend in population change is not an average of total numbers of animals lost." According to WWF, this could lead to a sixth major extinction event. The five main causes of

biodiversity loss Biodiversity loss includes the worldwide extinction of different species, as well as the local reduction or loss of species in a certain habitat, resulting in a loss of biological diversity. The latter phenomenon can be temporary or permanent, de ...

are

land-use change Land use, land-use change, and forestry (LULUCF), also referred to as Forestry and other land use (FOLU), is defined by the United Nations Climate Change Secretariat as a "greenhouse gas inventory sector that covers emissions and removals of gree ...

, overexploitation of natural resources,

climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...

pollution Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, the ...

and

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...

References

Bibliography

* *

External links

Tree of Life

* ttp://entomology.ifas.ufl.edu/fasulo/vector/chapter_07.htm Vertebrate Pestschapter in

United States Environmental Protection Agency The Environmental Protection Agency (EPA) is an independent executive agency of the United States federal government tasked with environmental protection matters. President Richard Nixon proposed the establishment of EPA on July 9, 1970; it be ...

and

University of Florida The University of Florida (Florida or UF) is a public land-grant research university in Gainesville, Florida. It is a senior member of the State University System of Florida, traces its origins to 1853, and has operated continuously on its ...

Institute of Food and Agricultural Sciences The University of Florida Institute of Food and Agricultural Sciences (UF/IFAS) is a teaching, research and Extension scientific organization focused on agriculture and natural resources. It is a partnership of federal, state, and county governmen ...

National Public Health Pesticide Applicator Training Manual
The Vertebrates

Marc W. Kirschner Marc Wallace Kirschner (born February 28, 1945) is an American cell biologist and biochemist and the founding chair of the Department of Systems Biology at Harvard Medical School. He is known for major discoveries in cell and developmental biolog ...

, ''iBioSeminars'', 2008. {{Authority control * Terreneuvian first appearances Extant Cambrian first appearances