Chordate

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Not to be confused with Cordate.

Chordates
Temporal range:
Ma[1]
(Possible Ediacaran record, 555 Ma[2])
Example of chordates: Branchiostoma lanceolatum (Cephalochordata), Polycarpa aurata (Tunicata), as well as a Tiger shark and a Siberian tiger (Vertebrata).
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Subkingdom: Eumetazoa
Clade: ParaHoxozoa
Clade: Bilateria
Clade: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Haeckel, 1874[3][4]
Subgroups

And see text

A chordate (/ˈkɔːrdeɪt/ KOR-dayt) is a deuterostomic animal belonging to the phylum Chordata (/kɔːrˈdeɪtə/ kor-DAY-tə). All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (synapomorphies) that distinguish them from other taxa. These five synapomorphies are a notochord, a hollow dorsal nerve cord, an 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 body plan structuring and movements. Chordates are also bilaterally symmetric, have a coelom, possess an enclosed 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.[5] 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.[6]

Chordate fossils have been found from as early as the Cambrian explosion, 539 million years ago.[7] 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[8] 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).

Anatomy

The glass catfish (Kryptopterus vitreolus) is one of the few chordates with a visible backbone. The spinal cord is housed within its backbone.

Chordates form a phylum of animals that are defined by having at some stage in their lives all of the following anatomical features:[9]

There are soft constraints that separate chordates from other biological lineages, but are not part of the formal definition:

1 = bulge in spinal cord ("brain")
4 = post-anal tail
5 = anus
6 =
digestive canal
8 =
atriopore
9 = space above pharynx
11 = pharynx
12 =
vestibule
13 = oral cirri
14 = mouth opening
16 = light sensor
17 =
nerves
19 = hepatic caecum (liver-like sack)
Amphioxus
. Bolded items are components of all chordates at some point in their lifetimes, and distinguish them from other phyla.

Classification

The following schema is from the 2015 edition of

Vertebrate Palaeontology.[13][14] The invertebrate chordate classes are from Fishes of the World.[15] While it is structured so as to reflect evolutionary relationships (similar to a cladogram), it also retains the traditional ranks used in Linnaean taxonomy
.

Subphyla

See also: List of chordate orders
Cephalochordate: Lancelet

Cephalochordata: Lancelets

Main article: Lancelet

Cephalochordates, one of the three subdivisions of chordates, are small, "vaguely fish-shaped" animals that lack brains, clearly defined heads and specialized sense organs.[21] These burrowing filter-feeders compose the earliest-branching chordate sub-phylum.[22][23]

Tunicata (Urochordata)

Main article: Tunicate (Urochordata)
Tunicates: sea squirts

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;[24] 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.[25] However, all tunicate larvae have the standard chordate features, including long, tadpole-like tails; they also have rudimentary brains, light sensors and tilt sensors.[24] 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.[26] 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.[27] The term Tunicata (Lamarck 1816) is recognised as having precedence and is now more commonly used.[24]

Comparison of two invertebrate chordates
syphon, 11. Exhalant syphon, 12. Heart, 13. Stomach, 14. Esophagus, 15. Intestines, 16. Tail, 17. Atrium, 18. Tunic

Craniata (Vertebrata)

Main articles:
Vertebrata
Craniate: Hagfish

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".[28]

Most craniates are

braincases and no vertebrae, and are therefore not regarded as vertebrates,[30]
but as members of the craniates, the group from which vertebrates are thought to have evolved.[31] However the cladistic exclusion of hagfish from the vertebrates is controversial, as they may be degenerate vertebrates who have lost their vertebral columns.[32]

The position of

Cyclostomata.[35]

Phylogeny

Overview

Haikouichthys, from about 518 million years ago in China, may be the earliest known fish.[36]

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.[37]

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. Recent identification of two conserved signature indels (CSIs) in the proteins cyclophilin-like protein and mitochondrial inner membrane protease ATP23, which are exclusively shared by all vertebrates, tunicates and cephalochordates also provide strong evidence of the monophyly of Chordata.[5]

All of the earliest chordate

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 "

paraphyletic
.

Deuterostomes
Ambulacraria

Hemichordates

Echinoderms

Chordates

Cephalochordates

Olfactores

Tunicates

Craniates (vertebrates)

Diagram of the evolutionary relationships of chordates[10]

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.[38]

The majority of animals more complex than

Cnidarians are split into two groups, the protostomes and deuterostomes, the latter of which contains chordates.[39] It seems very likely the 555 million-year-old Kimberella was a member of the protostomes.[40][41] If so, this means 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 538.8 million years ago.[39] The Ediacaran fossil Ernietta, from about 549 to 543 million years ago, may represent a deuterostome animal.[42]

A skeleton of the blue whale, the largest animal, extant or extinct, ever discovered, outside the Long Marine Laboratory at the University of California, Santa Cruz. The largest blue whale ever reliably recorded measured 98ft (29.9m) long.
A peregrine falcon, the world's fastest animal. Peregrines use gravity and aerodynamics to achieve their top speed of around 242mph (389km/h), as opposed to locomotion.

Fossils of one major deuterostome group, the

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.[46] Haikouichthys and Myllokunmingia, also from the Chengjiang fauna, are regarded as fish.[36][47] Pikaia, discovered much earlier (1911) but from the Mid Cambrian Burgess Shale (505 Ma), is also regarded as a primitive chordate.[48] 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.[49]

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.[10] 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.[50] 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.[10][51]

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.[51] However, molecular estimates of dates often disagree with each other and with the fossil record,[51] and their assumption that the molecular clock runs at a known constant rate has been challenged.[52][53]

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.[5]

Cladogram

invertebrates. The positions (relationships) of the lancelets, tunicates, and craniates/vertebrates are based on the following studies:[54][55][56][57]

Chordata
Cephalochordata

Amphioxiformes (lancelets)

Olfactores

†Haikouella

Tunicata

Larvacea
)

"

polyphyletic; sea squirts)

salps)

Vertebrata/
Cyclostomata

hagfishes)

lampreys)

(Agnathans)

†

Conodonta

†Pteraspidomorphi (includes †Arandaspida, †Astraspida and †Heterostraci)

†Cephalaspidomorphi (includes †Galeaspida, †Osteostraci and †Pituriaspida)

Gnathostomata

†"

paraphyletic; includes †Antiarchi, †Petalichthyida, †Ptyctodontida and †Arthrodira)

Crown

†"

paraphyletic)

rat fish)

Euteleostomi/

Actinopterygii (ray-finned fishes)

Sarcopterygii

Actinistia (coelacanths)

Dipnoi (lungfishes)

 Tetrapoda 

 Amphibia

 Amniota 
 
Synapsida
 

 

Mammalia

 Sauropsida 

 Lepidosauromorpha (lizards, snakes, tuatara, and their extinct relatives)

 

extinct relatives)

  Testudinata (turtles and their extinct relatives)

(four‑limbed vertebrates)
(lobe‑finned fishes)
Osteichthyes
Gnathostomata
Craniata

Closest nonchordate relatives

Acorn worms or Enteropneusts are example of hemichordates.

The closest relatives of the Chordates are believed to be the

Echinodermata, which together form the Ambulacraria
. The Chordata and Ambulacraria together form the superphylum
Deuterostomia
.

Hemichordates

Main article: Hemichordate

ventral
nerve cord.

There are two living groups of hemichordates. The solitary

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.[58]

Echinoderms

Protoreaster linckii is an example of Asterozoan Echinoderm
.
Main article: Echinoderm

sea cucumbers.[59]

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.[4]

See also

References

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External links

Wikispecies has information related to Chordata.
The Wikibook Dichotomous Key has a page on the topic of: Chordata
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