'''Chordates''' ([[phylum]] '''Chordata''') are a group of [[animal]]s that includes the [[vertebrate]]s, together with several closely related [[invertebrate]]s. They are united by having, at some time in their life cycle, a [[notochord]], a hollow [[dorsal nerve cord]], [[pharyngeal slit]]s, an [[endostyle]], and a post-anal [[tail]]. The phylum Chordata consists of three subphyla: [[Urochordata]], represented by [[tunicate]]s; [[Cephalochordata]], represented by [[lancelet]]s;and [[Craniata]], which includes [[vertebrate|Vertebrata]]. The [[Hemichordata]] have been presented as a fourth chordate subphylum, but they are now usually treated as a separate phylum. Urochordate larvae have a notochord and a nerve cord but these are lost in adulthood. Cephalochordates have a notochord and a nerve cord but no [[brain]] or specialist sense organs, and a very simple [[circulatory system]]. Craniates are the only sub-phylum whose members have [[skull]]s. In all craniates except for [[Hagfish]], the dorsal hollow nerve cord has been surrounded with [[cartilaginous]] or bony vertebrae and the notochord generally reduced; hence hagfish are not regarded as vertebrates. The chordates and three sister [[phylum|phyla]], the [[Hemichordata]], the [[Echinodermata]] and the [[Xenoturbellida]], make up the [[deuterostomes]], one of the two [[superphylum|superphyla]] which encompass all fairly complex animals.
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Attempts to work out the evolutionary relationships of the chordates have produced several hypotheses, but the current consensus is that chordates are [[monophyletic]], in other words contain all and only the descendants of a single common ancestor ''which is itself a chordate'', and that craniates' nearest relatives are cephalochordates. All of the earliest chordate [[fossil]]s have been found in the Early [[Cambrian]] [[Chengjiang fauna]], and include two species that are regarded as [[fish]], which implies that these are vertebrates. Because the fossil record of 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.
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It has also proved difficult to produce a detailed classification within the living chordates. Attempts to produce evolutionary "[[phylogenetic tree|family trees]]" give results that differ from traditional [[class (biology)|classes]] because several of those classes are not monophyletic. As a result vertebrate classification is in a state of flux.
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== Definition, sub-divisions and closest relatives ==
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=== Definition ===
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{{Annotated image | caption=Anatomy of the [[cephalochordate]] ''[[Amphioxus]]''
Chordates form a [[phylum]] - a grouping of animals with a shared [[bodyplan]]<ref>{{cite book
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| last = Valentine | first = J.W. | year = 2004 | title = On the Origin of Phyla
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| publisher = University Of Chicago Press | location = Chicago | isbn = 0226845486
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| page=7
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}}"<cite>Classifications of organisms in hierarchical systems were in use by the seventeenth and eighteenth centuries. Usually organisms were grouped according to their morphological similarities as perceived by those early workers, and those groups were then grouped according to <strong>their</strong> similarities, and so on, to form a hierarchy.</cite>"
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</ref> - defined by having at some stage in their lives all of the following:<ref name="RychelSmithShimamotoSwalla2006">{{cite journal
*a [[notochord]], in other words a fairly stiff rod of [[cartilage]] that extends along the inside of the body. Among the vertebrate sub-group of chordates the notochord develops into the [[Vertebral column | spine]], and in wholly aquatic species this helps the animal to swim by flexing its tail.
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*a [[Anatomical_terms_of_location#Dorsal_and_ventral|dorsal]] [[neural tube]]. In fish and other [[vertebrate]]s this develops into the [[spinal cord]], the main commmunications trunk of the [[nervous system]].
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*[[pharyngeal slit]]s. The [[pharynx]] is the part of the [[throat]] immediately behind the mouth. In [[fish]] the slits are modified to form [[gills]], but in some other chordates they are part of a [[filter feeding]] system that extracts particles of food from the water in which the animals live.
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*a muscular tail that extends backwards behind the [[anus]].
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*an [[endostyle]]. This is a groove in the [[Anatomical_terms_of_location#Dorsal_and_ventral|ventral]] wall of the pharynx. In [[filter feeding]] species it produces [[mucus]] to gather food particles, which helps in transporting food to the [[esophagus]].<ref name="Ruppert2005">{{cite journal
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| author=Ruppert, E. | journal=Canadian Journal of Zoology | volume=83 | pages=8–23 | year=2005
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| doi = 10.1139/Z04-158
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| title=Key characters uniting hemichordates and chordates: homologies or homoplasies?
}}</ref> It also stores [[iodine]], and may be a precursor of the vertebrate [[thyroid]] gland.<ref name="RychelSmithShimamotoSwalla2006" />
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{{clear}}
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===Sub-divisions===
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There are three major groupings within the chordates:
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{| border="0" cellspacing=3 cellpadding=0
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|- valign="top"
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| colspan="2" | [[Craniate]]s have distinct [[skull]]s. [[Michael J. Benton]] comments that "craniates are characterized by their heads, just as chordates, or possibly all [[deuterostomes]], are by their tails."<ref>{{cite book
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| author=Benton, M.J. | title=Vertebrate Palaeontology: Biology and Evolution
| title=Morphology of the Vertebrates | publisher=University of California Museum of Paleontology
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}}</ref> This consists of a series of bony or cartilaginous [[cylinder|cylindrical]] vertebrae, generally with [[neural arch]]es that protect the [[spinal chord]] and with projections that link the vertebrae. [[Hagfish]] have incomplete [[braincase]]s and no vertebrae, and are therefore not regarded as vertebrates,<ref>{{cite web
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| title=Introduction to the Myxini | publisher=University of California Museum of Paleontology
}}</ref> The position of [[lamprey]]s is ambiguous. They have complete braincases and rudimentary vertebrae, and therefore may be regarded as vertebrates and true [[fish]].<ref>{{cite web
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| title=Introduction to the Petromyzontiformes | publisher=University of California Museum of Paleontology
}}</ref> However [[molecular phylogenetics]], which uses [[biochemistry|biochemical]] features to classify organisms, has produced both results that group them with vertebrates and others that group them with hagfish.<ref>{{cite journal
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| author=Shigehiro Kuraku, S., Hoshiyama, D., Katoh, K., Suga, H, and Miyata, T.
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| title=Monophyly of Lampreys and Hagfishes Supported by Nuclear DNA–Coded Genes
|[[Image:Pacific hagfish Myxine.jpg| thumb | left | 100px | Craniate: [[Hagfish]] ]]
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|- valign="top"
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|colspan="2" | [[Cephalochordate]]s are small, "vaguely fish-shaped" animals that lack brains, clearly-defined heads and specialized sense organs.<ref>{{cite book
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| author=Benton, M.J. | title=Vertebrate Palaeontology: Biology and Evolution
}}</ref> These burrowing filter-feeders may be either the closest living relatives of craniates or surviving members of the group from which all other chordates evolved.<ref name="Gee2008AmphioxusUnleashed">{{cite journal
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| author=Gee, H. | title=Evolutionary biology: The amphioxus unleashed
| title=Branchiostoma | publisher=Lander University
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}}</ref>
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|[[Image:Branchiostoma lanceolatum.png| thumb | left | 100px | Cephalochordate: [[Lancelet]] ]]
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|- valign="top"
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|Most [[tunicate]]s appear as adults in two major forms, both of which are bags of jelly that lack the standard features of chordates: "sea squirts" are sessile and consist mainly of water pumps and filter feeding apparatus;<ref name="Benton2002VertebratePalaeontologyP5" /> [[salp]]s 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 [[Colony (biology)|colonies]].<ref>{{cite web
}}</ref> However all tunicate [[larva]]e have the standard chordate features, including long, [[tadpole]]-like tails; they also have rudimentary brains, light sensors and tilt sensors.<ref name="Benton2002VertebratePalaeontologyP5" /> 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.<ref>{{cite web
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| title=Appendicularia
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| publisher=Australian Government Department of the Environment, Water, Heritage and the Arts
}}</ref> Because of their larvae's long tails tunicates are also called [[urochordate]]s ("tail chordates").<ref name="Benton2002VertebratePalaeontologyP5">{{cite book
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| author=Benton, M.J. | title=Vertebrate Palaeontology: Biology and Evolution
|[[Image:BU Bio.jpg|thumb | left | 100px | Tunicates: sea squirts]]
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|[[Image:Salp.jpg| thumb | right | 100px | Tunicates: floating colony of [[salp]]s ]]
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|}
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===Closest non-chordate relatives===
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{| border="0" cellspacing=3 cellpadding=0
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|- valign="top"
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| [[Hemichordate]]s ("half 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 [[notochord]]s but running in a circle round the "collar", which is ahead of the mouth; and a [[Anatomical_terms_of_location#Dorsal_and_ventral|dorsal]] nerve cord – but also a smaller [[Anatomical_terms_of_location#Dorsal_and_ventral|ventral]] nerve cord. There are two living groups of hemichordates. The solitary [[enteropneust]]s, commonly known as "acorn worms", have long [[proboscis|probosces]] and worm-like bodies with up to 200 branchial slits, are up to {{convert|2.5|m|ft}} long, and burrow though seafloor [[sediment]]s. [[Pterobranch]]s are [[colony (biology)|colonial]] animals, often less than {{convert|1|mm|in}} long individually, whose dwellings are inter-connected. Each [[filter feeding|filter feed]]s by means of a pair of branched tentacles, and has a short, shield-shaped proboscis. The extinct [[graptolite]]s, colonial animals whose fossils look tiny [[hacksaw]] blades, lived in tubes similar to those of pterobranchs.<ref>{{cite web
| publisher=University of California Museum of Paleontology
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}}</ref>
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|[[Image:Balanoglossus 01.png| thumb | left | 100px | [[Enteropneust]] hemichordate: ''[[Balanoglossus]]'' ]]
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| [[Image:Rhabdopleura normani 01.png| thumb | right | 100px | [[Pterobranch]] hemichordate: [[Rhabdopleura]] ]]
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|- valign="top"
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| [[Echinoderm]]s differ from chordates' other relatives in three conspicuous ways: instead of having [[bilateral symmetry]] they have [[radial symmetry]], like wheels; their bodies are supported by [[skeleton]]s made of [[calcite]], a material not used by chordates, and these skeleton enclose their bodies but are also covered by a thin skin; they have [[tube feet]]. The feet are powered by another unique feature of echinoderms, a [[water vascular system]] of canals that also function as a "lung" and are surrounded by muscles that act as pumps. [[Crinoid]]s 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 urchin]]s and [[sea cucumber]]s.<ref name="CowenHistLifeEd3P412">{{ cite book
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| author=Cowen, R. | title=History of Life | year=2000 | page=412
| [[Image:Sandstar 300.jpg| thumb | left | 100px | Echinoderm: [[starfish]] ]]
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| [[Image:Featherx.JPG| thumb | right | 100px | Echinoderm: [[crinoid]] ]]
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|}
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== Origins ==
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The majority of animals more complex than [[jellyfish]] and other [[Cnidarians]] are split into two groups, the [[protostome]]s and [[deuterostome]]s, and chordates are deuterostomes.<ref name=Erwin20002>{{cite journal|author=Erwin, Douglas H.; Eric H. Davidson|year=2002|title=The last common bilaterian ancestor|journal=Development|volume=129|pages=3021–3032|url=http://dev.biologists.org/cgi/content/full/129/13/3021|pmid=12070079}}</ref> It seems very likely that {{ma|555|million years old}} ''[[Kimberella]]'' was a member of the protostomes.<ref name=Fedonkin2007>{{The Rise and Fall of the Ediacaran Biota|New data on ''Kimberella'', the Vendian mollusc-like organism (White sea region, Russia): palaeoecological and evolutionary implications|Fedonkin, M.A.; Simonetta, A; Ivantsov, A.Y.|157|179|12}}</ref><ref name=Butterfield2006>{{cite journal
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| author = Butterfield, N.J.
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| year = 2006
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| title = Hooking some stem-group "worms": fossil lophotrochozoans in the Burgess Shale
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| journal = Bioessays
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| volume = 28
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| issue = 12
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| pages = 1161–6
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| doi = 10.1002/bies.20507
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| accessdate = 2007-05-21
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}}</ref> If so, this means that the protostome and deuterostome lineages must have split some time before ''Kimberella'' appeared - at least {{ma|558}}, and hence well before the start of the Cambrian {{ma|Cambrian}}.<ref name=Erwin20002 /> The [[Ediacaran]] fossil ''[[Ernettia]]'', from about {{ma|549|543}}, may represent a deuterostome animal.<ref>{{cite journal
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| author=Dzik , J. | title=Organic membranous skeleton of the Precambrian metazoans from Namibia
[[Image:Haikouichthys4.png| thumb | right | 200px | ''[[Haikouichthys]]'', from about {{ma|518}} in China, may be the earliest known fish.<ref name="ShuConwayMorrisHan2003Haikouichthys" />]]
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Fossils of one major deuterostome group, the [[echinoderm]]s (whose modern members include [[starfish]], [[sea urchin]]s and [[crinoid]]s) are quite common from the start of the Cambrian, {{ma|542}}.<ref name="Bengtson2004">{{citation | contribution=Early skeletal fossils
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| author = Bengtson, S. | editor=Lipps, J.H., and Waggoner, B.M. | title=Neoproterozoic- Cambrian Biological Revolutions
}}</ref> Opinions differ about whether the [[Chengjiang fauna]] fossil ''[[Yunnanozoon]]'', from the earlier Cambrian, was a hemichordate or chordate.<ref name="ChenHangLi1996">{{cite journal
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| author=Shu, D., Zhang, X. and Chen, L. | month=April | year=1996
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| title= Reinterpretation of Yunnanozoon as the earliest known hemichordate
}}</ref> Another Chenjiang 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.<ref name="ChenHangLi1999" /> ''[[Haikouichthys]]'' and ''[[Myllokunmingia]]'', also from the Chenjiang fauna, are regarded as [[fish]].<ref name="ShuConwayMorrisHan2003Haikouichthys">{{cite journal
}}</ref> ''[[Pikaia]]'', discovered much earlier but from the Mid Cambrian [[Burgess Shale]], is also regarded as a primitive chordate.<ref>{{cite journal
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| author=Shu, D-G., Conway Morris, S., and Zhang, X-L.
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| title=A ''Pikaia''-like chordate from the Lower Cambrian of China
}}</ref> On the other hand fossils of early chordates are very rare, since non-vertebrate chordates have no bones or teeth, and none have been reported for the rest of the Cambrian.
</div>A consensus family tree of the chordates<ref name="Ruppert2005" /><ref name="Perseke2007" /></div>
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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, [[embryology|embryological]], and paleontological data have produced different "family trees". Some closely linked chordates and hemichordates, but that idea is now rejected.<ref name="Ruppert2005" /> Combining such analyses with data from a small set of [[ribosome]] [[RNA]] genes eliminated some older ideas, but open the possibility that tunicates (urochordates) are "basal deuterostomes", in other words surviving members of the group from which echinoderms, hemichordates and chordates evolved.<ref name="WinchellSullivanCameronSwallaMallatt2002">{{cite journal
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| author=Winchell, C.J., Sullivan, J., Cameron, C.B., Swalla, B.J., and Mallatt, J.
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| title=Evaluating Hypotheses of Deuterostome Phylogeny and Chordate Evolution with New LSU and SSU Ribosomal DNA Data
}}</ref> Most researchers agree that, within the chordates, craniates are most closely related to cephalochordates, but there also reasons for regarding tunicates (urochordates) as craniates' closest relatives.<ref name="Ruppert2005" /><ref name="BlairHedges2005">{{cite journal
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| author= Blair, J.E., and S. Blair Hedges, S.B.
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| title=Molecular Phylogeny and Divergence Times of Deuterostome Animals
}}</ref> One other phylum, [[Xenoturbellida]], appears to be basal within the deuterostomes, in other words closer to the original deuterostomes than to the chordates, echinoderms and hemichordates.<ref name="Perseke2007">Perseke M, Hankeln T, Weich B, Fritzsch G, Stadler PF, Israelsson O, Bernhard D, Schlegel M. (2007) "The mitochondrial DNA of ''Xenoturbella bocki'': genomic architecture and phylogenetic analysis". ''Theory Biosci''. 126(1):35-42. Available on-line at [http://www.bioinf.uni-leipzig.de/Publications/PREPRINTS/07-009.pdf]</ref>
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Since chordates have left a poor fossil record, attempts have been made to calculate the key dates in their evolution by [[molecular phylogenetics]] techniques, in other words by analysing biochemical differences, mainly in RNA. One such study suggested that deuterostomes arose before {{ma|900}} and the earliest chordates around {{ma|896}}.<ref name="BlairHedges2005" /> However molecular estimates of dates often disagree with each other and with the fossil record,<ref name="BlairHedges2005" /> and their assumption that the [[molecular clock]] runs at a known constant rate has been challenged.<ref name="Ayala1999MolClockMirages">{{cite journal |author= Ayala, F.J.|year=1999 |title=Molecular clock mirages |journal=[[BioEssays]] |volume=21|issue=1 |pages=71–75|url=http://www3.interscience.wiley.com/cgi-bin/abstract/60000186/ABSTRACT?CRETRY=1&SRETRY=0 |doi=10.1002/(SICI)1521-1878(199901)21:1<71::AID-BIES9>3.0.CO;2-B }}</ref><ref name="SchwartzMaresca20006DoMolClocksRun">{{cite journal |author=Schwartz, J. H. and Maresca, B. |year=2006 |title=Do Molecular Clocks Run at All? A Critique of Molecular Systematics |journal=Biological Theory |volume=1 |pages=357–371|url= |doi=10.1162/biot.2006.1.4.357}}</ref>
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<!-- **************
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The origin of chordates is currently unknown. The first clearly-identifiable chordates are reduced fish- or [[lancelet]]-like specimens from the [[Cambrian]]. Most speculations about their origin fit into one or more of these categories:
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* A sediment-dwelling worm-like animal that evolved a flatter body and/or fins for swimming.
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* A sessile tubular filter-feeder that evolved into a free-swimming animal via usage of fins. ([[Tunicates]], considered a chordate, are sessile filter feeders that have a tadpole-like larvae.)
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* A drifting or swimming larva of some other kind of animal that eventually retained its swimming features into adulthood.
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The notochord's stiffness in many chordates may have evolved to facilitate the effectiveness of alternating muscle contractions for swimming (in S-shaped movements). In other words, in order to bend the body, a muscle needs a rigid structure to pull against, and a notochord (at least before spines) is the main structure to provide this. Lack of a stiff body part would merely result in the shorting of the animal during muscle contractions instead of the bending motions needed for swimming.
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********* -->
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== Classification ==
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=== [[Taxonomy]] ===
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The following schema is from the third edition of ''[[Vertebrate Palaeontology (Benton)|Vertebrate Palaeontology]]''.<ref>Benton, M.J. (2004). ''Vertebrate Palaeontology'', Third Edition. Blackwell Publishing, 472 pp. [http://palaeo.gly.bris.ac.uk/benton/vertclass.html The classification scheme is available online]</ref> While it is structured so as to reflect evolutionary relationships (similar to a [[cladistics|cladogram]]), it also retains the traditional ranks used in [[Linnaean taxonomy]].
* Lines show probable evolutionary relationships, including extinct taxa, which are denoted with a [[Dagger (typography)|dagger]], †. Some are invertebrates. Chordata is a phylum.
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* The positions (relationships) of the Lancelet, Tunicate, and Craniata clades are as reported<ref>The amphioxus genome and the evolution of the chordate karyotype, Nicholas H. Putnam, et al. Nature vol 453 p. 1064-1071, June 19, 2000</ref> in the scientific journal [[Nature (journal)|''Nature'']].
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== References ==
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{{reflist}}
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== External links ==
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* [http://www.globaltwitcher.com/taxa_class.asp?phylaid=1 Chordate on GlobalTwitcher.com]
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* [http://tolweb.org/Chordata/2499 Chordate node at Tree Of Life]
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* [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=7711 Chordate node at NCBI Taxonomy]
Chordates (phylumChordata) are a group of animals that includes the vertebrates, together with several closely related invertebrates. They are united by having, at some time in their life cycle, a notochord, a hollow dorsal nerve cord, pharyngeal slits, an endostyle, and a post-anal tail. The phylum Chordata consists of three subphyla: Urochordata, represented by tunicates; Cephalochordata, represented by lancelets;and Craniata, which includes Vertebrata. The Hemichordata have been presented as a fourth chordate subphylum, but they are now usually treated as a separate phylum. Urochordate larvae have a notochord and a nerve cord but these are lost in adulthood. Cephalochordates have a notochord and a nerve cord but no brain or specialist sense organs, and a very simple circulatory system. Craniates are the only sub-phylum whose members have skulls. In all craniates except for Hagfish, the dorsal hollow nerve cord has been surrounded with cartilaginous or bony vertebrae and the notochord generally reduced; hence hagfish are not regarded as vertebrates. The chordates and three sister phyla, the Hemichordata, the Echinodermata and the Xenoturbellida, make up the deuterostomes, one of the two superphyla which encompass all fairly complex animals.
Attempts to work out the evolutionary relationships of the chordates have produced several hypotheses, but the current consensus is that chordates are monophyletic, in other words contain all and only the descendants of a single common ancestor which is itself a chordate, and that craniates' nearest relatives are cephalochordates. All of the earliest chordate fossils have been found in the Early CambrianChengjiang fauna, and include two species that are regarded as fish, which implies that these are vertebrates. Because the fossil record of 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" give results that differ from traditional classes because several of those classes are not monophyletic. As a result vertebrate classification is in a state of flux.
a notochord, in other words a fairly 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.
pharyngeal slits. The pharynx is the part of the throat immediately behind the mouth. In fish the slits are modified to form gills, but in some other chordates they are part of a filter feeding system that extracts particles of food from the water in which the animals live.
a muscular tail that extends backwards behind the anus.
an endostyle. This is a groove in the ventral wall of the pharynx. In filter feeding species it produces mucus to gather food particles, which helps in transporting food to the esophagus.[3] It also stores iodine, and may be a precursor of the vertebrate thyroid gland.[2]
There are three major groupings within the chordates:
Craniates have distinct skulls. Michael J. Benton comments that "craniates are characterized by their heads, just as chordates, or possibly all deuterostomes, are by their tails."[4] Most are vertebrates, in which the notochord is relaced by the spinal column.[5] This consists of a series of bony or cartilaginous cylindrical vertebrae, generally with neural arches that protect the spinal chord and with projections that link the vertebrae. Hagfish have incomplete braincases and no vertebrae, and are therefore not regarded as vertebrates,[6] but as members of the craniates, the group from which vertebrates are thought to have evolved.[7] The position of lampreys is ambiguous. They have complete braincases and rudimentary vertebrae, and therefore may be regarded as vertebrates and true fish.[8] 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.[9]
Cephalochordates are small, "vaguely fish-shaped" animals that lack brains, clearly-defined heads and specialized sense organs.[10] These burrowing filter-feeders may be either the closest living relatives of craniates or surviving members of the group from which all other chordates evolved.[11][12]
Most tunicates appear as adults in two major forms, both of which are bags of jelly that lack the standard features of chordates: "sea squirts" are sessile and consist mainly of water pumps and filter feeding apparatus;[13]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.[14] However all tunicate larvae have the standard chordate features, including long, tadpole-like tails; they also have rudimentary brains, light sensors and tilt sensors.[13] 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.[15] Because of their larvae's long tails tunicates are also called urochordates ("tail chordates").[13]
Hemichordates ("half 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 probosces and worm-like bodies with up to 200 branchial slits, are up to 2.5 metres (Template:Convert/ft)Template:Convert/test/A long, and burrow though seafloor sediments. Pterobranchs are colonial animals, often less than Template:Convert/mmTemplate:Convert/test/A long individually, whose dwellings are inter-connected. 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 tiny hacksaw blades, lived in tubes similar to those of pterobranchs.[16]
Echinoderms differ from chordates' other relatives in three conspicuous ways: instead of having bilateral symmetry they have radial symmetry, like wheels; their bodies are supported by skeletons made of calcite, a material not used by chordates, and these skeleton enclose their bodies but are also covered by a thin skin; they have tube feet. The feet are powered by another unique feature of echinoderms, a water vascular system of canals that also function as a "lung" and are 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.[17]
The majority of animals more complex than jellyfish and other Cnidarians are split into two groups, the protostomes and deuterostomes, and chordates are deuterostomes.[18] It seems very likely that 555 million years old
Kimberella was a member of the protostomes.[19][20] If so, this means that the protostome and deuterostome lineages must have split some time before Kimberella appeared - at least 558 million years ago
, and hence well before the start of the Cambrian Cambrian million years ago
.[18] The Ediacaran fossil Ernettia, from about 549 543
, may represent a deuterostome animal.[21]
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, 542 million years ago
.[23] The Mid Cambrian fossil Rhabdotubus johanssoni has been interpreted as a pterobranch hemichordate.[24] Opinions differ about whether the Chengjiang fauna fossil Yunnanozoon, from the earlier Cambrian, was a hemichordate or chordate.[25][26] Another Chenjiang 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.[26]Haikouichthys and Myllokunmingia, also from the Chenjiang fauna, are regarded as fish.[22][27]Pikaia, discovered much earlier but from the Mid Cambrian Burgess Shale, is also regarded as a primitive chordate.[28] On the other hand fossils of early chordates are very rare, since non-vertebrate chordates have no bones or teeth, and none have 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.[3] Combining such analyses with data from a small set of ribosomeRNA genes eliminated some older ideas, but open the possibility that tunicates (urochordates) are "basal deuterostomes", in other words surviving members of the group from which echinoderms, hemichordates and chordates evolved.[30] Most researchers agree that, within the chordates, craniates are most closely related to cephalochordates, but there also reasons for regarding tunicates (urochordates) as craniates' closest relatives.[3][31] One other phylum, Xenoturbellida, appears to be basal within the deuterostomes, in other words closer to the original deuterostomes than to the chordates, echinoderms and hemichordates.[29]
Since chordates have left a poor fossil record, attempts have been made to calculate the key dates in their evolution by molecular phylogenetics techniques, in other words by analysing 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
.[31] However molecular estimates of dates often disagree with each other and with the fossil record,[31] and their assumption that the molecular clock runs at a known constant rate has been challenged.[32][33]
The following schema is from the third edition of Vertebrate Palaeontology.[34] While it is structured so as to reflect evolutionary relationships (similar to a cladogram), it also retains the traditional ranks used in Linnaean taxonomy.
Lines show probable evolutionary relationships, including extinct taxa, which are denoted with a dagger, †. Some are invertebrates. Chordata is a phylum.
The positions (relationships) of the Lancelet, Tunicate, and Craniata clades are as reported[35] in the scientific journal Nature.
↑Valentine, J.W. (2004). On the Origin of Phyla, Chicago: University Of Chicago Press."Classifications of organisms in hierarchical systems were in use by the seventeenth and eighteenth centuries. Usually organisms were grouped according to their morphological similarities as perceived by those early workers, and those groups were then grouped according to their similarities, and so on, to form a hierarchy."
↑ 2.02.1Rychel, A.L., Smith, S.E., Shimamoto, H.T., and Swalla, B.J. (2006). Evolution and Development of the Chordates: Collagen and Pharyngeal Cartilage. Molecular Biology and Evolution23 (3): 541–549.
↑ 29.029.1Perseke M, Hankeln T, Weich B, Fritzsch G, Stadler PF, Israelsson O, Bernhard D, Schlegel M. (2007) "The mitochondrial DNA of Xenoturbella bocki: genomic architecture and phylogenetic analysis". Theory Biosci. 126(1):35-42. Available on-line at [1]
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