Vertebrates (), also called craniates, are animals with a vertebral column and a cranium. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebrata ( VUR-tə-BRAY-tə) with some 65,000 species, by far the largest ranked grouping in the phylum Chordata. The vertebrates include mammals, birds, amphibians, and various classes of fish and reptiles. The fish include the jawless Agnatha, and the jawed Gnathostomata. The jawed fish include both the cartilaginous fish and the bony fish. Bony fish include the lobe-finned fish, which gave rise to the tetrapods, the animals with four limbs. Despite their success, vertebrates still only make up less than five percent of all described animal species. The first vertebrates appeared in the Cambrian explosion some 518 million years ago. Jawed vertebrates evolved in the Ordovician or Silurian; bony fishes appeared in the Silurian and diversified widely in the Devonian. The first tetrapods appeared towards the end of the Devonian, and the first amphibians appeared on land in the Carboniferous. During the Triassic, mammals and dinosaurs appeared, the latter giving rise to birds in the Jurassic. 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, overexploitation of natural resources, climate change, pollution and the impact of invasive species.
Characteristics
Unique features Vertebrates belong to Chordata, a phylum characterised by five synapomorphies (unique characteristics): namely a notochord, a hollow nerve cord along the back, a post-anal tail, an endostyle (present in vertebrates as the thyroid gland), and pharyngeal gills arranged in pairs. Vertebrates share these characteristics with other chordates. Vertebrates are distinguished from all other animals, including other chordates, by multiple synapomorphies, namely: a vertebral column and a skull made of cartilage or bone; a brain divided into 3 or more sections; a muscular heart with 2 or more chambers; an inner ear with semicircular canals; camera-type eyes; a nose; kidneys with glomeruli; a thyroid; and a liver.
Physical
Vertebrates (and other chordates) belong to the Bilateria, a group of animals with mirror symmetrical bodies. They move, typically by swimming, using muscles along the back, supported by a strong but flexible skeletal structure, the spine or vertebral column. The name 'vertebrate' derives from the Latin vertebratus, 'jointed', from vertebra, 'joint', in turn from Latin vertere, 'to turn'.
As embryos, vertebrates still have a notochord. In all but the jawless fishes, it is replaced with a vertebral column (made of bone or cartilage) during development. Vertebrate embryos have pharyngeal arches; in adult fish, these support the gills, while in adult tetrapods they develop into other structures. In the embryo, a layer of cells along the back folds and fuses into a hollow neural tube. This develops into the spinal cord, and at its front end, the brain. The brain receives information about the world through nerves which carry signals from sense organs 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, 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 to the anus. The vertebral column typically continues beyond the anus to form an elongated tail.
The ancestral vertebrates, and most extant species, are aquatic and carry out gas exchange 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 arches. In jawed vertebrates, the first gill arch pair evolved into the jaws. In amphibians and some primitive bony fishes, the larvae have external gills, branching off from the gill arches. Oxygen is carried from the gills to the body in the blood, and carbon dioxide is returned to the gills, in a closed circulatory system driven by a chambered heart. The tetrapods have lost the gills of their fish ancestors; they have adapted the swim bladder (that fish use for buoyancy) into lungs to breathe air, and the circulatory system is adapted accordingly. At the same time, they adapted the bony fins of the lobe-finned fishes into two pairs of walking legs, carrying the weight of the body via the shoulder and pelvic girdles. Vertebrates vary in size from the smallest frog species such as Brachycephalus pulex, with a minimum adult snout–vent length of 6.45 millimetres (0.254 in) to the blue whale, at up to 33 m (108 ft) and weighing some 150 tonnes.
Molecular Molecular markers known as conserved signature indels in protein sequences 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 translation initiation factor 3, adenosine kinase and a protein related to ubiquitin carboxyl-terminal hydrolase). A specific relationship between vertebrates and tunicates is supported by two molecular markers, the proteins Rrp44 (associated with the exosome complex) and serine C-palmitoyltransferase. These are exclusively shared by species from these two subphyla, but not by cephalochordates.
Evolutionary history
Cambrian explosion
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