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Terreneuvian – Holocene, 542–0 Ma
|The Glass catfish (Kryptopterus vitreolus) is one of the few chordates with a visible backbone. The spinal cord is housed within its backbone.|
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A chordate is an animal belonging to the phylum Chordata; chordates possess a notochord, a hollow dorsal nerve cord, pharyngeal slits, an endostyle, and a post-anal tail, for at least some period of their life cycle. Chordates are deuterostomes, as during the embryo development stage the anus forms before the mouth. They are also bilaterally symmetric coelomates with metameric segmentation and a circulatory system. In the case of vertebrate chordates, the notochord is usually replaced by a vertebral column during development.
Taxonomically, the phylum includes the following subphyla: the Vertebrata, which includes fish, amphibians, reptiles, birds, and mammals; the Tunicata, which includes salps and sea squirts; and the Cephalochordata, which include the lancelets. There are also additional extinct taxa such as the Vetulicolia. The Vertebrata are sometimes considered as a subgroup of the clade Craniata, consisting of chordates with a skull; the Craniata and Tunicata compose the clade Olfactores.
Of the more than 65,000 living species of chordates, about half are bony fish of the superclass Osteichthyes. The world's largest and fastest animals, the blue whale and peregrine falcon respectively, are chordates, as are humans. Fossil chordates are known from at least as early as the Cambrian explosion.
Hemichordata, which includes the acorn worms, has been presented as a fourth chordate subphylum, but it now is usually treated as a separate phylum. The Hemichordata, along with the Echinodermata (which includes starfish, sea urchins, sea cucumbers, and crinoids), form the Ambulacraria, the sister taxon of the Chordates. The Chordata and Ambulacraria form the superphylum Deuterostomia, composed of the deuterostomes.
Attempts to work out the evolutionary relationships of the chordates have produced several hypotheses. The current consensus is that chordates are monophyletic, meaning that the Chordata include all and only the descendants of a single common ancestor, which is itself a chordate, and that craniates' nearest relatives are tunicates.
All of the earliest chordate fossils have been found in the Early Cambrian Chengjiang fauna, and include two species that are regarded as fish, which implies that they are vertebrates. Because the fossil record of early chordates is poor, only molecular phylogenetics offers a reasonable prospect of dating their emergence. However, the use of molecular phylogenetics for dating evolutionary transitions is controversial.
It has also proved difficult to produce a detailed classification within the living chordates. Attempts to produce evolutionary "family trees" shows that many of the traditional classes are paraphyletic.
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.
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.
There are soft constraints that separate chordates from certain other biological lineages, but have not yet been made part of the formal definition:
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.
Craniates, one of the three subdivisions of chordates, 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 are vertebrates, in which the notochord is replaced by the vertebral column. These consist of a series of bony or cartilaginous cylindrical vertebrae, generally with neural arches that protect the spinal cord, and with projections that link the vertebrae. However hagfish have incomplete braincases and no vertebrae, and are therefore not regarded as vertebrates, but as members of the craniates, the group from which vertebrates are thought to have evolved. However the cladistic exclusion of hagfish from the vertebrates is controversial, as they may be degenerate vertebrates who have lost their vertebral columns.
The position of lampreys is ambiguous. They have complete braincases and rudimentary vertebrae, and therefore may be regarded as vertebrates and true fish. However, molecular phylogenetics, which uses biochemical features to classify organisms, has produced both results that group them with vertebrates and others that group them with hagfish. If lampreys are more closely related to the hagfish than the other vertebrates, this would suggest that they form a clade, which has been named the Cyclostomata.
Most tunicates appear as adults in two major forms, 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 Urochorda(ta) (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.
Cephalochordates 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.
The majority of animals more complex than jellyfish and other Cnidarians are split into two groups, the protostomes and deuterostomes, the latter of which contains chordates. It seems very likely the Kimberella was a member of the protostomes. If so, this means the protostome and deuterostome lineages must have split some time before Kimberella appeared—at least , and hence well before the start of the Cambrian . The Ediacaran fossil Ernietta, from about , may represent a deuterostome animal.
Fossils of one major deuterostome group, the echinoderms (whose modern members include starfish, sea urchins and crinoids), are quite common from the start of the Cambrian, . The Mid Cambrian fossil Rhabdotubus johanssoni has been interpreted as a pterobranch hemichordate. Opinions differ about whether the Chengjiang fauna fossil Yunnanozoon, from the earlier Cambrian, was a hemichordate or chordate. Another fossil, Haikouella lanceolata, also from the Chengjiang fauna, is interpreted as a chordate and possibly a craniate, as it shows signs of a heart, arteries, gill filaments, a tail, a neural chord with a brain at the front end, and possibly eyes—although it also had short tentacles round its mouth. Haikouichthys and Myllokunmingia, also from the Chengjiang fauna, are regarded as fish. Pikaia, discovered much earlier (1911) but from the Mid Cambrian Burgess Shale (505 Ma), is also regarded as a primitive chordate. On the other hand, fossils of early chordates are very rare, since invertebrate chordates have no bones or teeth, and only one has been reported for the rest of the Cambrian.
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 and the earliest chordates around . 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.
The following schema is from the third edition of Vertebrate Palaeontology. The invertebrate chordate classes are from Fishes of the World. While it is structured so as to reflect evolutionary relationships (similar to a cladogram), it also retains the traditional ranks used in Linnaean taxonomy.
|Cladogram of the Chordate phylum. Lines show probable evolutionary relationships, including extinct taxa, which are denoted with a dagger, †. Some are invertebrates. The positions (relationships) of the Lancelet, Tunicate, and Craniata clades are as reported in the scientific journal Nature. Note that this cladogram, in showing the extant cyclostomes (hagfish and lamprey) as paraphyletic, is contradicted by nearly all recent molecular data, which support the monophyly of the extant cyclostomes (see Ota and Kurakani 2007 and references therein for a review of evidence).
Hemichordates ("half (½) chordates") have some features similar to those of chordates: branchial openings that open into the pharynx and look rather like gill slits; stomochords, similar in composition to notochords, but running in a circle round the "collar", which is ahead of the mouth; and a dorsal nerve cord—but also a smaller ventral nerve cord.
There are two living groups of hemichordates. The solitary enteropneusts, commonly known as "acorn worms", have long proboscises and worm-like bodies with up to 200 branchial slits, are up to 2.5 metres (8.2 ft) long, and burrow though seafloor sediments. Pterobranchs are colonial animals, often less than 1 millimetre (0.039 in) long individually, whose dwellings are interconnected. Each filter feeds by means of a pair of branched tentacles, and has a short, shield-shaped proboscis. The extinct graptolites, colonial animals whose fossils look like tiny hacksaw blades, lived in tubes similar to those of pterobranchs.
Echinoderms differ from chordates and their other relatives in three conspicuous ways: they possess bilateral symmetry only as larvae - in adulthood they have radial symmetry, meaning that their body pattern is shaped like a wheel; they have tube feet; and their bodies are supported by skeletons made of calcite, a material not used by chordates. Their hard, calcified shells keep their bodies well protected from the environment, and these skeletons enclose their bodies, but are also covered by thin skins. The feet are powered by another unique feature of echinoderms, a water vascular system of canals that also functions as a "lung" and surrounded by muscles that act as pumps. Crinoids look rather like flowers, and use their feather-like arms to filter food particles out of the water; most live anchored to rocks, but a few can move very slowly. Other echinoderms are mobile and take a variety of body shapes, for example starfish, sea urchins and sea cucumbers.
Although I was among the early supporters of vertebrate paraphyly, I am impressed by the evidence provided by Heimberg et al. and prepared to admit that cyclostomes are, in fact, monophyletic. The consequence is that they may tell us little, if anything, about the dawn of vertebrate evolution, except that the intuitions of 19th century zoologists were correct in assuming that these odd vertebrates (notably, hagfishes) are strongly degenerate and have lost many characters over time
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