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Biology of Vertebrates

Craniates, one of the three sub-divisions of chordates, 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 replaced by the spinal column. [5]

This consists 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. 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[9] features to classify organisms, has produced both results that group them with vertebrates and others that group them with hagfish.

[edit] Cephalochordata: "The Lancelets"

Cephalochordate: Lancelet

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]

[edit] Urochordata: "The Tunicates"

Tunicates: sea squirts

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;[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]

[edit] Closest non-chordate relatives

[edit] The Hemichordates

EnteropneustBalanoglossus hemichordate:

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 (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 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 like tiny hacksaw blades, lived in tubes similar to those of pterobranchs.[16]

[edit] The Echinoderms

Echinoderm: starfish

Echinoderms differ from chordates and their other relatives in three conspicuous ways: instead of having bilateral symmetry they have radial symmetry, meaning 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. The hard calcified shell keeps their bodies well protected from the environment, and these skeletons enclose their bodies but are also covered by a thin skin. 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]

[edit] Origins

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]555 million year 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 542 million years ago.[18] The Ediacaran fossil Ernietta, from about 549 to 543 million years ago, may represent a deuterostome animal.[21] It seems very likely that

Haikouichthys, from about 518 million years ago in China, may be the earliest known fish.[22]

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 johanssonipterobranch hemichordate.[24] Opinions differ about whether the Chengjiang fauna fossil Yunnanozoon, from the earlier Cambrian, was a hemichordate or chordate.[25][26] 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.[26]Haikouichthys and Myllokunmingia, also from the Chengjiang 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 only one has been reported for the rest of the Cambrian.[29] has been interpreted as a






TunicatesUrochordates) (



A consensus family tree of the chordates[3][30]

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 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.[31] Most researchers agree that, within the chordates, craniates are most closely related to cephalochordates, but there are also reasons for regarding tunicates (urochordates) as craniates' closest relatives.[3][32] 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.[30]

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.[32] However molecular estimates of dates often disagree with each other and with the fossil record,[32] and their assumption that the molecular clock[33][34] runs at a known constant rate has been challenged.

[edit] Classification

[edit] Taxonomy

The following schema is from the third edition of Vertebrate Palaeontology.[35] The invertebrate chordate classes are from Fishes of the World.[36] While it is structured so as to reflect evolutionary relationships (similar to a cladogram), it also retains the traditional ranks used in Linnaean taxonomy.

  • Phylum Chordata
    • Subphylum Tunicata (Urochordata) — (tunicates; 3,000 species)
      • Class Ascidiacea
      • Class Thaliacea (salps)
      • Class Appendicularia (larvacea)
    • Subphylum Cephalochordata (Acraniata) — (lancelets; 30 species)
    • Subphylum Vertebrata (Craniata) (vertebrates — animals with backbones; 57,674 species)
        • Class 'Agnatha' paraphyletic (jawless vertebrates; 100+ species)
          • Subclass Myxinoidea (hagfish; 65 species)
          • Subclass Petromyzontida (lampreys)
          • Subclass Conodonta
          • Subclass Pteraspidomorphi (Paleozoic jawless fish)
            • Order Anaspida
            • Order Thelodonti (Paleozoic jawless fish)
      • Infraphylum Gnathostomata (jawed vertebrates)
          • Class Placodermi (Paleozoic armoured forms)
          • Class Chondrichthyes (cartilaginous fish; 900+ species)
          • Class Acanthodii (Paleozoic "spiny sharks")
          • Class Osteichthyes (bony fish; 30,000+ species)
            • Subclass Actinopterygii (ray-finned fish; about 30,000 species)
            • Subclass Sarcopterygii (lobe-finned fish)
        • Superclass Tetrapoda (four-legged vertebrates; 28,000+ species)
          • Class Amphibia (amphibians; 6,000 species)
            • Series Amniota (with amniotic egg)
          • Class Reptilia (reptiles; 8,225+ species)
            • Subclass Anapsida (extinct "proto-reptiles" and possibly turtles)
            • Subclass Synapsida (mammal-like "reptiles"; 4,500+ species, progenitors of mammals)
            • Subclass Diapsida (majority of reptiles, progenitors of birds)
          • Class Mammalia (mammals; 5,800 species)
          • Class Aves (birds; 8,800–10,000 species)

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