|Example of chordates: Branchiostoma lanceolatum (Cephalochordata), Polycarpa aurata (Tunicata), as well as a Tiger shark and a Siberian tiger (Vertebrata).|
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A chordate (/ˈkɔːrdeɪt/ KOR-dayt) is an animal belonging to the phylum Chordata (/kɔːrˈdeɪtə/ kor-DAY-tə). 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 synapomorphies include a notochord, dorsal hollow nerve cord, endostyle or thyroid, pharyngeal slits, and a post-anal tail. The name "chordate" comes from the first of these synapomorphies, the notochord, which plays a significant role in chordate structure and movement. Chordates are also bilaterally symmetric, have a coelom, possess a circulatory system, and exhibit metameric segmentation.
In addition to the morphological characteristics used to define chordates, analysis of genome sequences has identified two conserved signature indels (CSIs) in their proteins: cyclophilin-like protein and mitochondrial inner membrane protease ATP23, which are exclusively shared by all vertebrates, tunicates and cephalochordates. These CSIs provide molecular means to reliably distinguish chordates from all other Metazoa.
Chordates are divided into three subphyla: Craniate or Vertebrate (fish, amphibians, reptiles, birds, and mammals); Tunicata or Urochordata (sea squirts, salps and relatives, and larvaceans); and Cephalochordata (which includes lancelets). The Craniata and Tunicata compose the clade Olfactores, which is sister to Cephalochordata. (See diagram under Phylogeny.) Extinct taxa such as Vetulicolia and Conodonta are Chordata, but their internal placement is less certain. Hemichordata (which includes the acorn worms) was previously considered a fourth chordate subphylum, but now is treated as a separate phylum: hemichordates and Echinodermata form the Ambulacraria, the sister phylum of the Chordates. The Chordata and Ambulacraria, together and possibly with the Xenacoelomorpha, are believed to form the superphylum Deuterostomia, although this has recently been called into doubt.
Chordate fossils have been found from as early as the Cambrian explosion, 539 million years ago. Cladistically (phylogenetically), vertebrates – chordates with the notochord replaced by a vertebral column during development – are a subgroup of the clade Craniata, which consists of chordates with a skull. Of the more than 81,000 living species of chordates, about half are ray-finned fishes that are members of the class Actinopterygii and the vast majority of the rest are tetrapods (mostly birds and mammals).
Chordates form a phylum of animals that are defined by having at some stage in their lives all of the following anatomical features:
- A notochord, a stiff rod of cartilage that extends along the inside of the body. Among the vertebrate sub-group of chordates the notochord develops into the spine, and in wholly aquatic species this helps the animal to swim by flexing its tail.
- A dorsal neural tube. In fish and other vertebrates, this develops into the spinal cord, the main communications trunk of the nervous system.
- filter-feeding system that extracts particles of food from the water in which the animals live. In tetrapods, they are only present during embryonic stages of the development.
- Post-anal tail. A muscular tail that extends backwards behind the anus. In some chordates such as humans, this is only present in the embryonic stage.
There are soft constraints that separate chordates from other biological lineages, but are not part of the formal definition:
- All chordates are deuterostomes. This means that, during the embryo development stage, the anus forms before the mouth.
- All chordates are based on a
- All chordates are coelomates, and have a fluid-filled body cavity called a coelom with a complete lining called peritoneum derived from mesoderm (see Brusca and Brusca).
The following schema is from the 2015 edition of
- Phylum Chordata
- Subphylum Cephalochordata(Acraniata) – (lancelets; 30 species)
- Class Leptocardii(lancelets)
- Clade Olfactores
- Subphylum Tunicata(Urochordata) – (tunicates; 3,000 species)
- Class Ascidiacea (sea squirts)
- Class Thaliacea (salps, doliolids and pyrosomes)
- Class Appendicularia (larvaceans)
- Class Sorberacea
- Subphylum Craniata) (vertebrates – animals with backbones; 66,100+ species)
- Superclass 'paraphyletic(jawless vertebrates; 100+ species)
- Class Cyclostomata
- Infraclass Myxini(hagfish; 65 species)
- Infraclass Petromyzontida or Hyperoartia(lampreys)
- Class †Conodonta
- Class †Myllokunmingiida
- Class †Pteraspidomorphi
- Class †Thelodonti
- Class †Anaspida
- Class †Cephalaspidomorphi
- Infraphylum Gnathostomata (jawed vertebrates)
- Class †Placodermi (Paleozoic armoured forms; paraphyletic in relation to all other gnathostomes)
- Class Chondrichthyes (cartilaginous fish; 900+ species)
- Class †Acanthodii (Paleozoic "spiny sharks"; paraphyletic in relation to Chondrichthyes)
- Class Osteichthyes (bony fish; 30,000+ species)
- Subclass Actinopterygii (ray-finned fish; about 30,000 species)
- Subclass Sarcopterygii (lobe-finned fish: 8 species)
- Superclass Tetrapoda (four-limbed vertebrates; 35,100+ species) (The classification below follows Benton 2004, and uses a synthesis of rank-based Linnaean taxonomy and also reflects evolutionary relationships. Benton included the Superclass Tetrapoda in the Subclass Sarcopterygii in order to reflect the direct descent of tetrapods from lobe-finned fish, despite the former being assigned a higher taxonomic rank.)
- Superclass '
Cephalochordates, one of the three subdivisions of chordates, are small, "vaguely fish-shaped" animals that lack brains, clearly defined heads and specialized sense organs. These burrowing filter-feeders compose the earliest-branching chordate sub-phylum.
Most tunicates appear as adults in two major forms, known as "sea squirts" and salps, both of which are soft-bodied filter-feeders that lack the standard features of chordates. Sea squirts are sessile and consist mainly of water pumps and filter-feeding apparatus; salps float in mid-water, feeding on plankton, and have a two-generation cycle in which one generation is solitary and the next forms chain-like colonies. However, all tunicate larvae have the standard chordate features, including long, tadpole-like tails; they also have rudimentary brains, light sensors and tilt sensors. The third main group of tunicates, Appendicularia (also known as Larvacea), retain tadpole-like shapes and active swimming all their lives, and were for a long time regarded as larvae of sea squirts or salps. The etymology of the term Urochordata (Balfour 1881) is from the ancient Greek οὐρά (oura, "tail") + Latin chorda ("cord"), because the notochord is only found in the tail. The term Tunicata (Lamarck 1816) is recognised as having precedence and is now more commonly used.
Craniates all have distinct skulls. They include the hagfish, which have no vertebrae. Michael J. Benton commented that "craniates are characterized by their heads, just as chordates, or possibly all deuterostomes, are by their tails".
Most craniates are
The position of
There is still much ongoing differential (DNA sequence based) comparison research that is trying to separate out the simplest forms of chordates. As some lineages of the 90% of species that lack a backbone or notochord might have lost these structures over time, this complicates the classification of chordates. Some chordate lineages may only be found by DNA analysis, when there is no physical trace of any chordate-like structures.
Attempts to work out the evolutionary relationships of the chordates have produced several hypotheses. The current consensus is that chordates are
All of the earliest chordate
It has also proved difficult to produce a detailed classification within the living chordates. Attempts to produce evolutionary "
While this has been well known since the 19th century, an insistence on only monophyletic taxa has resulted in vertebrate classification being in a state of flux.
The majority of animals more complex than
Fossils of one major deuterostome group, the
The evolutionary relationships between the chordate groups and between chordates as a whole and their closest deuterostome relatives have been debated since 1890. Studies based on anatomical, embryological, and paleontological data have produced different "family trees". Some closely linked chordates and hemichordates, but that idea is now rejected. Combining such analyses with data from a small set of ribosome RNA genes eliminated some older ideas, but opened up the possibility that tunicates (urochordates) are "basal deuterostomes", surviving members of the group from which echinoderms, hemichordates and chordates evolved. Some researchers believe that, within the chordates, craniates are most closely related to cephalochordates, but there are also reasons for regarding tunicates (urochordates) as craniates' closest relatives.
Since early chordates have left a poor fossil record, attempts have been made to calculate the key dates in their evolution by molecular phylogenetics techniques—by analyzing biochemical differences, mainly in RNA. One such study suggested that deuterostomes arose before 900 million years ago and the earliest chordates around 896 million years ago. However, molecular estimates of dates often disagree with each other and with the fossil record, and their assumption that the molecular clock runs at a known constant rate has been challenged.
Traditionally, Cephalochordata and Craniata were grouped into the proposed clade "Euchordata", which would have been the sister group to Tunicata/Urochordata. More recently, Cephalochordata has been thought of as a sister group to the "Olfactores", which includes the craniates and tunicates. The matter is not yet settled.
A specific relationship between Vertebrates and Tunicates is also strongly supported by two CSIs found in the proteins predicted exosome complex RRP44 and serine palmitoyltransferase, that are exclusively shared by species from these two subphyla but not Cephalochordates, indicating Vertebrates are more closely related to Tunicates than Cephalochordates.
Closest nonchordate relatives
The closest relatives of the Chordates are the
There are two living groups of hemichordates. The solitary
History of name
Although the name Chordata is attributed to William Bateson (1885), it was already in prevalent use by 1880. Ernst Haeckel described a taxon comprising tunicates, cephalochordates, and vertebrates in 1866. Though he used the German vernacular form, it is allowed under the ICZN code because of its subsequent latinization.
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