Vertebrates () are

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

s with a

vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...

(backbone or spine), and a

cranium The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...

, or skull. The vertebral column surrounds and protects the

spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...

, while the cranium protects the

brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...

. The vertebrates make up the

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

Vertebrata with some 65,000 species, by far the largest ranked grouping in the

phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...

Chordata A chordate ( ) is a bilaterian animal belonging to the phylum Chordata ( ). All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (Apomorphy and synapomorphy, synapomorphies) th ...

. The vertebrates include

mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...

s, and various classes of

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

and

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

s. The fish include the jawless

Agnatha Agnatha (; ) or jawless fish is a paraphyletic infraphylum of animals in the subphylum Vertebrata of the phylum Chordata, characterized by the lack of jaws. The group consists of both extant taxon, living (Cyclostomi, cyclostomes such as hagfish ...

, and the jawed

Gnathostomata Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant vertebrates, including all living bony fishes (both ray-finned ...

. The jawed fish include both the

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

and the

bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...

. Bony fish include the

lobe-finned fish Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...

, which gave rise to the

tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...

s, the animals with four limbs. Despite their success, vertebrates still only make up less than five percent of all described

animal species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of classification and ...

. The first vertebrates appeared in the Cambrian explosion some 518 million years ago. Jawed vertebrates evolved in the

Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...

, followed by bony fishes in the

Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...

. The first amphibians appeared on land in the

Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...

. During the

Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...

s and

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

s appeared, the latter giving rise to

s in the

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...

. Extant species are roughly equally divided between fishes of all kinds, and tetrapods. Populations of many species have been in steep decline since 1970 because of

land-use change Land use is an umbrella term to describe what happens on a parcel of land. It concerns the benefits derived from using the land, and also the land management actions that humans carry out there. The following categories are used for land use: for ...

overexploitation Overexploitation, also called overharvesting or ecological overshoot, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to ...

natural resource Natural resources are resources that are drawn from nature and used with few modifications. This includes the sources of valued characteristics such as commercial and industrial use, aesthetic value, scientific interest, and cultural value. ...

climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...

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

and the impact of

invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...

Characteristics

Unique features

Vertebrates belong to

, a

characterised by five

synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...

(unique characteristics): namely a

notochord The notochord is an elastic, rod-like structure found in chordates. In vertebrates the notochord is an embryonic structure that disintegrates, as the vertebrae develop, to become the nucleus pulposus in the intervertebral discs of the verteb ...

, a hollow nerve cord along the back, an

endostyle The endostyle is an organ found in invertebrate chordate species of tunicates, lancelets, and in the larval stage of vertebrate lampreys. It assists in filter-feeding. It has evolved into the thyroid in vertebrate chordates. Since the endo ...

(often as a

thyroid The thyroid, or thyroid gland, is an endocrine gland in vertebrates. In humans, it is a butterfly-shaped gland located in the neck below the Adam's apple. It consists of two connected lobes. The lower two thirds of the lobes are connected by ...

gland), and pharyngeal

gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...

s arranged in pairs. Vertebrates share these characteristics with other chordates. Vertebrates are distinguished from all other animals, including other chordates, by multiple synapomorphies: namely the vertebral column, skull of bone or cartilage, large brain divided into 3 or more sections, a muscular heart with multiple chambers; an inner ear with

semicircular canals The semicircular canals are three semicircular interconnected tubes located in the innermost part of each ear, the inner ear. The three canals are the lateral, anterior and posterior semicircular canals. They are the part of the bony labyrinth, ...

; sense organs including eyes, ears, and nose; and digestive organs including intestine, liver, pancreas, and stomach.

Physical

Vertebrates (and other chordates) belong to the

Bilateria Bilateria () is a large clade of animals characterised by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front (or "head") and a rear (or "tail") end, as well as a left� ...

, a group of animals with mirror symmetrical bodies. They move, typically by swimming, using

muscle Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...

s along the back, supported by a strong but flexible

skeletal A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fram ...

structure, the spine or

. The name 'vertebrate' derives from the

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

, 'jointed', from ''

vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...

'', 'joint', in turn from Latin , 'to turn'. As embryos, vertebrates still have a notochord; as adults, all but the jawless fishes have a vertebral column, made of

bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...

cartilage Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...

, instead. Vertebrate embryos have

pharyngeal arch The pharyngeal arches, also known as visceral arches'','' are transient structures seen in the Animal embryonic development, embryonic development of humans and other vertebrates, that are recognisable precursors for many structures. In fish, t ...

es; in adult

, these support the

s, while in adult

s they develop into other structures. In the embryo, a layer of cells along the back folds and fuses into a hollow

neural tube In the developing chordate (including vertebrates), the neural tube is the embryonic precursor to the central nervous system, which is made up of the brain and spinal cord. The neural groove gradually deepens as the neural folds become elevated, ...

. This develops into the

, and at its front end, the

. The brain receives information about the world through nerves which carry signals from

sense organ A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of Stimulus (physiology), stimuli. Although, in some cultures, five human senses were traditio ...

s in the skin and body. Because the ancestors of vertebrates usually moved forwards, the front of the body encountered stimuli before the rest of the body, favouring

cephalisation Cephalization is an evolutionary trend in animals that, over a sufficient number of generations, concentrates the special sense organs and nerve ganglia towards the front of the body where the mouth is located, often producing an enlarged head. ...

, the evolution of a head containing sense organs and a brain to process the sensory information. Vertebrates have a tubular gut that extends from the

mouth A mouth also referred to as the oral is the body orifice through which many animals ingest food and animal communication#Auditory, vocalize. The body cavity immediately behind the mouth opening, known as the oral cavity (or in Latin), is also t ...

to the

anus In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...

. The vertebral column typically continues beyond the anus to form an elongated

tail The tail is the elongated section at the rear end of a bilaterian animal's body; in general, the term refers to a distinct, flexible appendage extending backwards from the midline of the torso. In vertebrate animals that evolution, evolved to los ...

. The ancestral vertebrates, and most extant species, are aquatic and carry out

gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a b ...

in their gills. The gills are finely-branched structures which bring the blood close to the water. They are positioned just behind the head, supported by cartilaginous or bony

branchial arch Branchial arches or gill arches are a series of paired bony/ cartilaginous "loops" behind the throat ( pharyngeal cavity) of fish, which support the fish gills. As chordates, all vertebrate embryos develop pharyngeal arches, though the eve ...

es. In

jawed vertebrate Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant vertebrates, including all living bony fishes (both ray-finned ...

s, the first gill arch pair evolved into the jaws. In

s and some primitive bony fishes, the larvae have

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 fri ...

, branching off from the gill arches.

Oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...

is carried from the gills to the body in the

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 is com ...

, and

carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...

is returned to the gills, in a closed

circulatory system In vertebrates, the circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the body. It includes the cardiovascular system, or vascular system, that consists of the heart ...

driven by a chambered

heart The heart is a muscular Organ (biology), organ found in humans and other animals. This organ pumps blood through the blood vessels. The heart and blood vessels together make the circulatory system. The pumped blood carries oxygen and nutrie ...

. The

s have lost the gills of their fish ancestors; they have adapted the

swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ (anatomy), organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift ...

(that fish use for buoyancy) into

lung The lungs are the primary Organ (biology), organs of the respiratory system in many animals, including humans. In mammals and most other tetrapods, two lungs are located near the Vertebral column, backbone on either side of the heart. Their ...

s to breathe air, and the circulatory system is adapted accordingly. At the same time, they adapted the bony fins of the

lobe-finned fishes Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class (biology), class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. The ...

into two pairs of walking

leg A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element cap ...

s, carrying the weight of the body via the shoulder and pelvic girdles. Vertebrates vary in size from the smallest

frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...

species such as '' Brachycephalus pulex'', with a minimum adult

snout–vent length Snout–vent length (SVL) is a morphometric measurement taken in herpetology from the tip of the snout to the most posterior opening of the cloacal slit (vent)."direct line distance from tip of snout to posterior margin of vent" It is the mos ...

of 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 ever to have existed. The blue whale's long and slender body can ...

, at up to and weighing some 150 tonnes.

Molecular

Molecular marker In molecular biology and other fields, a molecular marker is a molecule, sampled from some source, that gives information about its source. For example, DNA is a molecular marker that gives information about the organism from which it was taken. ...

s 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 ...

protein sequence Protein primary structure is the linear sequence of amino acids in a peptide or protein. By convention, the primary structure of a protein is reported starting from the amino-terminal (N) end to the carboxyl-terminal (C) end. Protein biosynthe ...

s have been identified and provide distinguishing criteria for the vertebrate subphylum. Five molecular markers are exclusively shared by all vertebrates and reliably distinguish them from all other animals; these include 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 GTP-binding translation elongation factor family. Th ...

, eukaryotic translation initiation factor 3,

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 ...

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 Described and extant vertebrate species are split roughly evenly but non-phylogenetically between non-tetrapod "fish" and tetrapods. The following table lists the number of described Extant taxa, extant species for each vertebrate Class (biology), class as estimated in the IUCN Red List of Threatened Species, 2014.3.The World Conservation Union. 2014. ''IUCN Red List of Threatened Species'', 2014.3. Summary Statistics for Globally Threatened Species
Table 1: Numbers of threatened species by major groups of organisms (1996–2014)
Paraphyletic groups are shown in quotation marks. The IUCN estimates that 1,305,075 Invertebrate#Number of extant species, extant invertebrate species have been described, which means that less than 5% of the Animal#Number of extant species, described animal species in the world are vertebrates.

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 World Wide Fund for Nature , WWF, this could lead to a sixth Extinction event , major extinction event. The five main causes of biodiversity loss are

and

Notes

References

Bibliography

* *

External links

Tree of Life

* [http://entomology.ifas.ufl.edu/fasulo/vector/chapter_07.htm Vertebrate Pests] chapter in United States Environmental Protection Agency and University of Florida/Institute of Food and Agricultural Sciences National Public Health Pesticide Applicator Training Manual
The Vertebrates

Marc W. Kirschner, ''iBioSeminars'', 2008. {{Authority control Vertebrates, * Terreneuvian first appearances Extant Cambrian first appearances