Etymology: Vertebrates (

Vertebrates (pronounced /vrtbrts/) are animals that are members of the
subphylum Vertebrata (chordates with backbones and spinal columns).

Vertebrates include the overwhelming majority of the phylum chordata, with
currently about 58,000 species described.[2] Vertebrates include the jawless fishes,
bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds. Extant
vertebrates range in size from the frog species Paedophryne amauensis, at as little
as 7.7 mm (0.3 inch), to the blue whale, at up to 33 m (110 ft). Vertebrates make
up about 5% of all described animal species; the rest are invertebrates, which lack
backbones.
The vertebrates traditionally include the hagfishes, which do not have proper
vertebrae, though their closest living relatives, the lampreys, do have vertebrae.[3]
Hagfishes do, however, possess a cranium. For this reason, the vertebrate
subphylum is sometimes referred to as "Craniata" when discussing morphology.
Molecular analysis since 1992 has suggested that the hagfishes are most closely
related to lampreys[4], and so also are vertebrates in a monophyletic sense. Others
consider them a sister group of vertebrates in the common taxon of Craniata.[5]
Etymology
The word vertebrate derives from the Latin word vertebratus (Pliny), meaning
joint of the spine.[6] It is closely related to the word vertebra, which refers to any
of the bones or segments of the spinal column.[7]
[edit] Anatomy and morphology

All vertebrates are built along the basic chordate body plan: a stiff rod running
through the length of the animal (vertebral column or notochord),[8] with a hollow
tube of nervous tissue (the spinal cord) above it and the gastrointestinal tract
below. In all vertebrates, the mouth is found at, or right below, the anterior end of
the animal, while the anus opens to the exterior before the end of the body. The
remaining part of the body continuing aft of the anus forms a tail with vertebrae
and spinal cord, but no gut.[9]
[edit] Vertebral column
The defining characteristic of a vertebrate is the vertebral column, in which the

notochord (a stiff rod of uniform composition) found in all chordates has been
replaced by a segmented series of stiffer elements (vertebrae) separated by mobile
joints (intervertebral discs, derived embryonically and evolutionarily from the
notochord). However, a few vertebrates have secondarily lost this anatomy,
retaining the notochord into adulthood, such as the sturgeon[10] and the Latimeria.
Jawed vertebrates are typified by paired appendages (fins or legs, which may be
secondarily lost), but this is not part of the definition of vertebrates as a whole.
Fossilized skeleton of Diplodocus carnegii, showing an extreme example of the
backbone that characterizes the vertebrates. Exhibited at the Museum fr
Naturkunde (Museum of Natural Science), Berlin.
[edit] Gills
Gill arches bearing gills in a pike
All basal vertebrates breathe with gills. The gills are carried right behind the head,
bordering the posterior margins of a series of openings from the esophagus to the
exterior. Each gill is supported by a cartilagenous or bony gill arch.[11] The bony
fish have three pairs of arches, cartilaginous fish have five to seven pairs, while
the primitive jawless fish have seven. The vertebrate ancestor no doubt had more
arches, as some of their chordate relatives have more than 50 pairs of gills.[9]
In amphibians and some primitive bony fishes, the larvae bear external gills,
branching off from the gill arches.[12] These are reduced in adulthood, their
function taken over by the gills proper in fishes and by lungs in most amphibians.
Some amphibans retain the external larval gills in adulthood, the complex internal
gill system as seen in fish apparently being irrevocably lost very early in the
evolution of tetrapods.[13]
While the higher vertebrates do not have gills, the gill arches form during fetal
developement, and lay the basis of essential structures such as jaws, the thyroid
gland, the larynx, the columella (corresponding to the stapes in mammals) and in
mammals the malleus and incus.[9]
[edit] Central nervous system
The vertebrates are the only chordate group to exhibit a proper brain. A slight
swelling of the anterior end of the nerve cord is found in the lancelet, though it
lacks the eyes and other complex sense organs comparable to those of vertebrates.
Other chordates do not show any trends towards cephalisation.[9]
The central nervous system is based on a hollow nerve tube running along the
length of the animal, form which the peripheral nervous system branches out to
enervate the various systems. The front end of the nerve tube is expanded by a
thickening of the walls and expansion of the central canal of spinal cord into three
primary brain vesicles: The prosencephalon (forebrain), mesencephalon
(midbrain) and rhombencephalon (hindbrain), further differentiated in the various
vertebrate groups.[14] Two laterally placed eyes form around outgrows from the
midbrain, except in hagfish, though this may be a secondary loss.[15][16] The
forebrain is well developed and subdivided in most tetrapods, while the midbrain
dominate in many fish and some salamanders. Vesicles of the forebrain are
usually paired, giving rise to hemispheres like the cerebral hemispheres in
mammals.[14] The resulting anatomy of the central nervous system, with a single,
hollow nerve cord topped by a series of (often paired) vesicles is unique to
vertebrates. All invertebrates with well developed brains, like insects, spiders and
squids have a ventral rather than dorsal system of ganglions, with a split brain
stem running on each side of the mouth/gut.[9]
[edit] Evolutionary history

[edit] First vertebrates
The early vertebrate Haikouichthys
Vertebrates originated about 525 million years ago during the Cambrian
explosion, which saw the rise in organism diversity. The earliest known vertebrate
is believed to be the Myllokunmingia.[1] Another early vertebrate is Haikouichthys
ercaicunensis. Unlike the other fauna that dominated the Cambrian, these groups
had the basic vertebrate body plan: a notochord, rudimentary vertebrae, and a
well-defined head and tail.[17] All of these early vertebrates lacked jaws in the
common sense and relied on filter feeding close to the seabed.[18]
[edit] From fishes to amphibians
Acanthostega, a fish-like early labyrinthodont.
The first jawed vertebrates appeared in the latest Ordovician and became common
in the Devonian, often known as the "Age of Fishes".[19] The two groups of bony
fishes, the actinopterygii and sarcopterygii, evolved and became common.[20] The
Devonian also saw the demise of virtually all jawless fishes, save for lampreys
and hagfish, as well as the Placodermi, a group of armoured fish that dominated
much of the late Silurian. The Devonian also saw the rise of the first
labyrinthodonts, which was a transitional between fishes and amphibians.
[edit] Mesozoic vertebrates
Silesaurus, an archosaur
The reptiles appeared from labyrinthodonts in the subsequent Carboniferous

period. The anapsid and synapsid reptiles were common during the late Paleozoic,
while the diapsids became dominant during the Mesozoic. In the sea, the bony
fishes became dominant. The dinosaurs gave rise to the birds in the Jurassic.[21]
The demise of the dinosaurs at the end of the Cretaceous promoted expansion of
the mammals, which had evolved from the therapsids, a group of synapsid
reptiles, during the late Triassic Period.
[edit] After the Mesozoic

Frogs like Palaeobatrachus were among the animals to successfully diversify in
the post-Mesozoic world
The Cenozoic world has seen great diversification of bony fishes, frogs, birds and
mammals.
Over half of all living vertebrate species (about 32,000 species) are fishes (non-
tetrapod craniates), a diverse set of lineages that inhabit all the world's aquatic
ecosystems, from snow minnows (Cypriniformes) in Himalayan lakes at
elevations over 4,600 metres (15,000 feet) to flatfishes (order Pleuronectiformes)
in the Challenger Deep, the deepest ocean trench at about 11,000 metres (36,000
feet). Fishes of myriad varieties are the main predators in most of the worlds
water bodies, both freshwater and marine. The rest of the vertebrate species are
tetrapods, a single lineage that includes amphibians (frogs, with more than 5,800
species; salamanders, with about 580 species; and caecilians, with about 175
species); mammals (with over 5,400 species); and reptiles and birds (with more
than 18,000 species). Tetrapods dominate the megafauna of most terrestrial
environments (including fossorial and arboreal realms) and also include many
partially or fully aquatic groups (e.g., sea snakes, penguins, cetaceans).
[edit] Classification
There are several ways of classifying animals. Evolutionary systematics relies on
anatomy, physiology and evolutionary history, which is determined through
similarities in anatomy and, if possible, the genetics of organisms. Phylogenetic
classification is based solely on phylogeny.[22] Evolutionary systematics gives an
overview; phylogenetic systematics gives detail. The two systems are thus
complementary rather than opposed.[23]
[edit] Traditional classification
Traditional spindle diagram of the evolution of the vertebrates at class level
Conventional classification has living vertebrates grouped into seven classes

based on traditional interpretations of gross anatomical and physiological traits.
This classification is the one most commonly encountered in school textbooks,
overviews, non-specialist, and popular works. The extant vertebrates are:[9]
Subphylum Vertebrata
o Class Agnatha (jawless fishes)
o Class Chondrichthyes (cartilaginous fishes)
o Class Osteichthyes (bony fishes)
o Class Amphibia (amphibians)
o Class Reptilia (reptiles)
o Class Aves (birds)
o Class Mammalia (mammals)
In addition to these comes two classes of extinct armoured fishes, the Placodermi
and the Acanthodii. Other ways of classifying the vertebrates have been devised,
particularly with emphasis on the phylogeny on early amphibians and reptiles. An
example based on Janvier (1981, 1997), Shu et al. (2003), and Benton (2004)[24] is
given here:
o Superclass Agnatha or Cephalaspidomorphi (lampreys and other
jawless fishes)
o Infraphylum Gnathostomata (vertebrates with jaws)

Class Placodermi (extinct armoured fishes)
Class Chondrichthyes (cartilaginous fishes)
Class Acanthodii (extinct spiny "sharks")
Superclass Osteichthyes (bony fishes)
Class Actinopterygii (ray-finned bony fishes)
Class Sarcopterygii (lobe-finned fishes, some
ancestral to tetrapods)
Superclass Tetrapoda (four-limbed vertebrates)
Class Amphibia (amphibians, some ancestral to the
amniotes)
Class Synapsida (extinct mammal-like "reptiles",
some ancestral to mammals, sometimes classed with
Reptilia)
Class Reptilia (reptiles, some ancestral to birds)
Class Aves (birds)
Class Mammalia (mammals)
While this traditional classification is orderly, most of the groups are paraphyletic,
i.e. do not contain all descendants of the class's common ancestor.[24] For instance,
descendants of the first reptiles include modern reptiles as well as birds. Most of
the classes listed are not "complete" taxa, meaning that they do not include all the
descendants of the first representative of the group. For example, the agnathans
have given rise to the jawed vertebrates; the bony fishes have given rise to the
land vertebrates; the traditional "amphibians" have given rise to the reptiles
(traditionally including the synapsids, or "mammal-like "reptiles"), which in turn
have given rise to the mammals and birds. Most scientists working with
vertebrates use a classification based purely on phylogeny, organized by their
known evolutionary history and sometimes disregarding the conventional
interpretations of their anatomy and physiology.
[edit] Phylogenetic relationships
In phylogenetic taxonomy, the relationships between animals are not typically

divided into ranks, but illustrated as a nested "family tree" known as a cladogram.
Phylogenetic groups are given definitions based on their relationship to one
another, rather than purely on physical traits such as the presence of a backbone.
This nesting pattern is often combined with traditional taxonomy (as above), in a
practice known as evolutionary taxonomy.
The cladogram presented below is based on studies compiled by Philippe Janvier

and others for the Tree of Life Web Project.[25]
Vert Hyperoartia (lampreys)
ebrat
a ?Euconodonta
unn Pteraspidomorphi
ame ?Thelodonti
d unn ?Anaspida
ame unn Galeaspida
d ame unn ?Pituriaspida
d ame Osteostraci
d Gnatho Placodermi (armoured fishes)
stomat unn Chondrichthyes (cartilaginous
a ame fishes)
d Tele Acanthodii
osto Osteic Actinopterygii
mi hthye (ray-finned
s fishes)
Sarco ?Onychod
pteryg ontiformes
ii Coelacanthi
morpha
(coelacanth
s)
unn Porolepif
ame ormes
d Dipnoi
(lungfishe
s)
unn Rhizodon
ame timorpha
d Tristich
opteridae
Four
limbed
vertebrat
es
[edit] See also

Invertebrate
Marine vertebrates
[edit] References
1. ^ a b Shu et al.; Luo, H-L.; Conway Morris, S.; Zhang, X-L.; Hu, S-X.;
Chen, L.; Han, J.; Zhu, M. et al (November 4, 1999). "Lower Cambrian
vertebrates from south China". Nature 402 (6757): 4246. Bibcode
1999Natur.402...42S. doi:10.1038/46965.
2. ^ Jonathan E.M. Baillie, et al. (2004). "A Global Species Assessment".
World Conservation Union.
3. ^ Kuraku et al.; Hoshiyama, D; Katoh, K; Suga, H; Miyata, T (December
1999). "Monophyly of Lampreys and Hagfishes Supported by Nuclear
DNACoded Genes". Journal of Molecular Evolution 49 (6): 72935.
doi:10.1007/PL00006595. PMID 10594174.
4. ^ Stock, David; Whitt GS (7). "Evidence from 18S ribosomal RNA
sequences that lampreys and hagfishes form a natural group". Science 257
(5071): 7879. doi:10.1126/science.1496398. PMID 1496398. Retrieved
22 November 2011.
5. ^ Nicholls, Henry (10 September 2009). "Mouth to Mouth". Nature 461
(7261): 164166. doi:10.1038/461164a. PMID 19741680.
6. ^ Douglas Harper, Historian. "vertebrate". Online Etymology Dictionary.
Dictionary.com..
7. ^ Douglas Harper, Historian. "vertebra". Online Etymology Dictionary.
Dictionary.com..
8. ^ Waggoner, Ben. "Vertebrates: More on Morphology". UCMP. Retrieved
13 July 2011.
9. ^ a b c d e f Romer, A.S. (1949): The Vertebrate Body. W.B. Saunders,
Philadelphia. (2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970)
10. ^ Liem, Karel F.; Warren Franklin Walker (2001). Functional anatomy of
the vertebrates: an evolutionary perspective. Harcourt College Publishers.
p. 277. ISBN 9780030223693.
11. ^ Scott, Thomas (1996). Concise encyclopedia biology. Walter de Gruyter.
p. 542. ISBN 9783110106619.
12. ^ The American Naturalist (Essex Institute) 91: 287. 1957.
13. ^ Clack, J. A. (2002): Gaining ground: the origin and evolution of
tetrapods. Indiana University Press, Bloomington, Indiana. 369 pp
14. ^ a b Hildebrand, M. & Gonslow, G. (2001): Analysis of Vertebrate
Structure. 5th edition. John Wiley & Sons, Inc. New York
15. ^ "Keeping an eye on evolution". PhysOrg.com. 2007-12-03. Retrieved
2007-12-04.
16. ^ Hyperotreti - Hagfishes
17. ^ Waggoner, Ben. "Vertebrates: Fossil Record". UCMP. Retrieved 15 July
2011.
18. ^ Tim Haines, Paul Chambers (2005). The Complete Guide to Prehistoric
Life. Firefly Books.
19. ^ Encyclopaedia Britannica: a new survey of universal knowledge,
Volume 17. Encyclopdia Britannica. 1954. p. 107.
20. ^ Berg, Linda R.; Eldra Pearl Solomon, Diana W. Martin (2004). Biology.
Cengage Learning. p. 599. ISBN 9780534492762.
21. ^ Cloudsley-Thompson, J. L. (2005). Ecology and behaviour of Mesozoic
reptiles. 9783540224211: Springer. p. 6.
22. ^ Andersen, Nils Mller; Tom A. Weir (2004). Australian water bugs:
their biology and identification (Hemiptera-Heteroptera, Gerromorpha &
Nepomorpha). Apollo Books. p. 38. ISBN 9788788757781.
23. ^ Hildebran, M. & Gonslow, G. (2001): Analysis of Vertebrate Structure.
5th edition. John Wiley & Sons, Inc. New York, page 33: Comment: The
problem of naming sister groups
24. ^ a b Benton, Michael J. (2004-11-01). Vertebrate Palaeontology (Third
ed.). Blackwell Publishing. pp. 33, 455 pp.. ISBN 0632056371/978-
0632056378.
25. ^ Janvier, Philippe. 1997. Vertebrata. Animals with backbones. Version
01 January 1997 (under construction).
http://tolweb.org/Vertebrata/14829/1997.01.01 in The Tree of Life Web
Project, http://tolweb.org/
[edit] Bibliography
Kardong, Kenneth V. (1998). Vertebrates: Comparative Anatomy,
Function, Evolution (second ed.). USA: McGraw-Hill. pp. 747 pp..
ISBN 0-07-115356-X/0-697-28654-1.
"Vertebrata". Integrated Taxonomic Information System. Retrieved 6
August 2007.
[edit]
Vertebrata
Animals with backbones

Philippe Janvier
Interrelationships of the fossil and Recent Vertebrata. (All terminal taxa are clades,
except for the Thelodonti, which are possibly paraphyletic - see Thelodonti page).
The main characteristics supporting the nodes of this phylogeny are:
Node 1: Mineralized exoskeleton, sensory-line canals and grooves

Node 2: Perichondral bone or calcification, externally open endolymphatic duct
Node 3: Paired fins containing musculature and concentrated in pectoral position,
two dorsal fins, epicercal (i.e. upwardly tappering) tail, sclerotic ring and scleral
ossification, cellular dermal bone
Containing group: Craniata
Introduction
The Vertebrata, or vertebrates, is a very diverse group, ranging from lampreys to

Man. It includes all craniates, except hagfishes, and are characterized chiefly by a
vertebral column, hence their name. The majority of the extant vertebrates are the
jawed vertebrates, or gnathostomes, but lampreys are jawless vertebrates.
However, in Late Silurian or Early Devonian times, about 420 to 400 million
years ago, the situation was reverse, and the majority of the vertebrate species
were jawless fishes (the "ostracoderms", presumably more closely related to the
gnathostomes than to lampreys). The decline of the jawless vertebrates and the
subsequent rise of the gnathostomes took place about 380 million years ago.
Extant vertebrates comprise two clades: the Hyperoartia, or lampreys, and the
Gnathostomata, or jawed vertebrates. In addition, there is a number of taxa of
fossil jawless vertebrates which were formerly referred to as the "ostracoderms"
("shell-skinned") because most of them possess an extensive, bony endo- and
exoskeleton. The "ostracoderms" lived from the Early ordovician (about 480
million years ago) to the Late Devonian (about 370 million years ago). The
relationships of the various groups of "ostracoderms" has been the subject of
considerable debate since the mid-nineteenth Century, and the theory of
relationship proposed here is far from definitive, yet the best supported by the
currently available data. The "ostracoderms" are represented by five major groups,
four of which are almost certainly clades: the Heterostraci, Osteostraci,
Galeaspida, Anaspida, and Thelodonti (the monophyly of the latter being debated,
Thelodonti page). In addition, there are minor groups which only include a few
species: the Arandaspida, Astraspida, Eriptychiida, and Pituriaspida. The
Arandaspida, Astraspida, Eriptychiida, and Heterostraci are regarded as forming a
clade, the Pteraspidomorphi. Some monospecific genera, Jamoytius, Endeiolepis,
and Euphanerops, formerly referred to the Anaspida, are now removed from that
clade and may be more closely related to lampreys (see Hyperoartia). A large but
still poorly known group, the Euconodonta, has recently been included in the
Craniata, and possibly the Vertebrata. It is currently referred to as 'conodonts', but
the only forms that can reliably be regarded as craniates belong to a subgroup of
conodonts known as euconodonts.
Characteristics
The Vertebrata have all the characteristics of the Craniata but share, in addition, a
number of unique characteristics which do not occur in hagfishes (Hyperotreti).
These characteristics are:
Metamerically arranged endoskeletal elements flanking the spinal cord. There

are primitively two pairs of such elements in each metamere and on each side:
the interdorsals and basidorsals. In the gnathostomes, there are two additional
pairs ventrally to the notochord: the interventrals and basiventrals. These
elements are called arcualia and can fuse to a notochordal calcification, the
centrum. The ensemble of the arcualia , centrum is the vertebra, and the
ensemble of the vertebrae is the vertebral column.
The vertebrates are characterized by a vertebral column; that is, a variable
number of endoskeletal elements aligned along the notochord (green) and
flanking the spinal cord (yellow). In lampreys (top), the vertebral elements
are only the basidorsal (red) and the interdorsals (blue). In the
gnathostomes, there are in addition ventral elements, the basiventrals
(purple) and interventrals (orange), and the notochord may calcify into
centra (pink). (After Janvier 1996).
Extrinsic eye muscles. These muscles are attached to the eyeball and orbital
wall, and ensure eye movements
Radial muscles in fins. These are small muscles associated with each of the
cartilaginous radials of the unpaired and paired fins. They ensure the undulatory
movements of the fin web.
Atrium and ventricle of heart closely-set.
Nervous regulation of heart. The heart in the embryo of the vertebrates is
aneural, like the heart of adult hagfishes. In adult vertebrates, however, the
heart is innervated by a branch of the vagus nerve.
Typhlosole in the intestine. This is a spirally coiled fold of the intestinal wall. In
the Gnathostomes, it can be developed into a complex spiral valve.
At least two vertical semicircular canals in the labyrinth
True neuromasts in the sensory-line system
There are many other vertebrate characteristics, both anatomical and

physiological.
Discussion of Phylogenetic Relationships
As for extant vertebrates, the main question is whether lampreys are the sister-
group of the gnathostomes, or that of hagfishes. In the latter case there would be
no reason to distinguish the Vertebrata from the Craniata, as it was formerly done.
Although there is good evidence for the lamprey-gnathostome sister-group
relationship, the theory that the cyclostomes (lampreys,hagfishes) are a clade is
still supported by a number of zoologists. Considering the large number of
anatomical, physiological and molecular data that are available now to test these
theories, one can expect a definitive clue in a near future (for discussion, see
Craniata).
The question of the relationships of the numerous extinct vertebrate groups is, in
contrast, far from being resolved. This chiefly concerns the Palaeozoic taxa
formerly referred to as "ostracoderms"; that is, armored jawless craniates, which
are likely to be vertebrates and are now considered as being all more closely
related to the gnathostomes than to lampreys.
During most of the nineteenth century, the "ostracoderms" known at that time (i.e.
the Heterostraci and Osteostraci) were regarded as bony fishes, until Cope (1889)
suggested to include them with lampreys and hagfishes in the taxon Agnatha
("jawless"). In the beginning of the twentieth century, Kiaer (1924) and Stensi
(1927) showed that the Anaspida and Osteostraci share with lampreys a median,
dorsally placed "nostril" (in fact a nasohypophysial opening) and suggested to
include these three groups in a clade Cephalaspidomorphi. In addition, Stensi
(1927) proposed that hagfishes were derived from the Heterostraci and should be
grouped with them in the Pteraspidomorphi. At that time, however, the Agnatha
were regarded as a clade, whose sister-group was the Gnathostomata, as
illustrated by Stensi's (1927) diagram:
This theory implied the diphyletic origin of the Recent "cyclostomes" (hagfishes
and lampreys). Although they accepted the monophyly of the
Cephalaspidomorphi, most paleontologists rejected that of the Pteraspidomorphi
(as including hagfishes). In contrast, until the 1970's, it was widely accepted that
the Heterostraci are more closely related, or ancestral to the gnathostomes, mainly
because they lacked the specializations of the Cephalaspidomorphi and because
they had paired olfactory capsules, like the gnathostomes. With the rise of
cladistics, in the late 1970's and the 1980's, and following Lvtrup's (1977)
suggestion that extant cyclostomes were paraphyletic, a number of trees were
published, which all showed the "ostracoderms" (and the Agnatha as a whole) as
paraphyletic. However, all these trees implied that lampreys had lost several
characteristics, in particular the paired fins, mineralized skeleton, and sensory-line
canals. A major change was Gagnier's (1993) first computer-generated tree, in
which these reversions were avoided by considering all "ostracoderms" as more
closely related to the gnathostomes than to either lampreys and hagfishes. Further
analyses (Forey & Janvier 1994, Janvier 1996b) largely confirmed the higher
degree of parsimony of this phylogeny. Although there are variations as to the
position of certain taxa, the Galeaspida and Osteostraci constantly group together
with the Gnathostomes, whereas the Astraspida, Eriptychiida, Arandaspida, and
Heterostraci form a clade, the Pteraspidomorphi, albeit poorly supported. One of
the consequences of this tree is that the dorsal nasohypophysial opening (formerly
the characteristic of the Cephalaspidomorphi) either occurred more than once, or
is a general feature of the Vertebrata.
In this tree, four fossil groups are positioned with a question mark. In the case of
the Euconodonta, Anaspida and Pituriaspida, this uncertainty is largely due to the
scarcity of the characters available from the material (in particular as to the
internal anatomy). In the case of the Thelodonti, it is due to their controversial
status, as they are likely to be a paraphyletic assemblage of stem Heterostraci and
possibly stem forms of other "ostracoderm" groups, yet some authors regard them
as a clade (see Thelodonti page).
Other Names for Vertebrata
vertebrates
Animals with backbones
References
Forey, P. L. (1984). Yet more reflections on agnathan-gnathostome relationships.

Journal of Vertebrate Paleontology, 4, 330-343.
Forey, P. L., and Janvier, P. (1993). Agnathans and the origin of jawed
vertebrates. Nature, 361, 129-134.
Forey, P. L., and Janvier, P. (1994). Evolution of the early vertebrates. American
Scientist, 82, 554-565.
Hardisty, M. W. (1982). Lampreys and hagfishes: Analysis of cyclostome

relationships. In The Biology of Lampreys, (ed. M. W. Hardisty and I. C. Potter),
Vol.4B, pp. 165-259. Academic Press, London.
Janvier, P. (1993). Patterns of diversity in the skull of jawless fishes. In The skull
(ed. J. Hanken and B. K. Hall), Vol. 2, pp. 131-188. The University of Chicago
Press.
Janvier, P. (1996a). Early vertebrates. Oxford Monographs in Geology and

Geophysics, 33, Oxford University Press, Oxford.
Janvier, P. (1996b). The dawn of the vertebrates: characters versus common

ascent in current vertebrate phylogenies. Palaeontology, 39, 259-287.
Lvtrup, S. (1977). The Phylogeny of Vertebrata. Wiley, New York.

Stensi, E. A. (1927). The Devonian and Downtonian vertebrates of Spitsbergen.
1. Family Cephalaspidae. Skrifter om Svalbard og Ishavet, 12, 1-391.
Wang, N. Z. (1991). Two new Silurian galeaspids (Jawless craniates) from

Zhejiang province, China, with a discussion of galeaspid-gnathostome
relationships.In Early vertebrates and related problems of evolutionary biology
(ed. M. M. Chang, Y. H. Liu, and G. R. Zhang), pp. 41-65. Science Press, Beijing.
Information on the Internet
DigiMorph. The Digital Morphology library is a dynamic archive of information

on digital morphology and high-resolution X-ray computed tomography of
biological specimens. A National Science Foundation Digital Library at the
University of Texas at Austin.
Title Illustrations
Scientific Name Selene vomer
Specimen Condition Dead Specimen
Body Part skeleton
Copyright 1995 Tierney Thys
About This Page
Philippe Janvier
Musum National d'Histoire Naturelle Paris, France
Page copyright 1997 Philippe Janvier
Page: Tree of Life Vertebrata. Animals with backbones.

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Vertebrates, which include fishes, reptiles, amphibians, birds, and mammals, all
share a vertebral column, or a chain of bony elements (vertebrae) that run along
the dorsal surface from head to tail and form the main skeletal axis of the body.
The vertebral column surrounds and more or less replaces the notochord as the
chief "stiffener" of the body in locomotion. Some characteristics shared by most
or all vertebrates (in addition to those traits shared among all chordates) include
the following (after Hickman, 1994):
integument of two divisions, including an outer epidermis and an inner

dermis; integument often modified to produce hair, scales, feathers,
glands, horn, etc.
replacement of notochord by vertebral column more or less complete,
depending on group
bony or cartilaginous endoskeleton consisting of cranium, visceral arches,
limb girdles, and 2 pairs of appendages
muscular, perforated pharynx; this structure is the site of gills in fishes but
is much reduced in adult land-dwelling forms (although it is extremely
important in embryonic development of all vertebrates)
movements provided by muscles attached to endoskeleton
digestive system with large digestive glands, liver, and pancreas
ventral heart with 2-4 chambers
blood with red blood corpuscles containing hemoglobin, and in addition,
white corpuscles
well developed body cavity (coelom) containing visceral systems
paired kidneys with ducts to drain waste to exterior
most vertebrates with two sexes, each with paired gonads (there are some
exceptions)
general body plan consisting of head, trunk, 2 pairs of appendages, and
postanal tail (but these structures are highly modified in many vertebrates
and sometimes absent).
For More Information

Find Vertebrata information at
Encyclopedia of Life
Contributors
Phil Myers (author), Museum of Zoology, University of Michigan.
To cite this page: Myers, P. 2001. "Vertebrata" (On-line), Animal Diversity Web.
Accessed April 21, 2012 at
http://animaldiversity.ummz.umich.edu/site/accounts/information/Vertebrata.html
Disclaimer: The Animal Diversity Web is an educational resource written

largely by and for college students. ADW doesn't cover all species in the world,
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information in those accounts. While ADW staff and contributors provide
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Identification
Ablabys taenianotus (Cockatoo fish)
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permission of the copyright holder.
Contributors
Michigan Science Art (copyright holder)
To cite this page: Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond,

and T. A. Dewey. 2012. The Animal Diversity Web (online). Accessed at
http://animaldiversity.org.

Identification
Trachylepis aurata (Golden Grass Mabuya)
Ablepharus pannonicus
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Location
Farhang Torki Ecology and Herpetology Center for Research, Nourabad City,
Lorestan Province, Iran
Caption
Transcaucasian golden grass mabuya (Trachylepis aurata transcaucasica) eating

an Asian snake-eye skink (Ablepharus pannonicus), captive.
Contributors
Farhang Torki (photographer; copyright holder; identification)


Identification
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Asian snake-eye skink (Ablepharus pannonicus), captive.
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Identification
Abrocoma bennettii (Bennett's chinchilla rat)

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Abrocoma boliviensis (Bolivian chinchilla rat)
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Abrocoma cinerea (ashy chinchilla rat)
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Abudefduf sexfasciatus (Scissor-tail sergeant)
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Scissortail sergeant (Abudefduf sexfasciatus)
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Klaus Jost (photographer; copyright holder; identification)
Identification
Abudefduf vaigiensis (Common sergeant)

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Caption
Sergeant Major Damselfish, or Indo-Pacific Damsel. Location: Sail Rock (Hin

Bai), Koh Samui, Thailand. Depth: 12m.
Contributors
Martin Taylor (photographer; copyright holder; identification)
Identification
Acantharchus pomotis (Mud sunfish)
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Mud sunfish (Acantharchus pomotis)
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United States Fish and Wildlife Service (copyright holder)
Identification
Acanthemblemaria maria (Secretary Blenny)

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Acanthisitta chloris (rifleman)
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female
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Identification
Acanthisitta chloris (rifleman)
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male
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Identification
Acanthixalus spinosus
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Acanthiza chrysorrhoa (yellow-rumped thornbill)
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Acanthocybium solandri (Barracuda)
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Acanthophis antarcticus (Common death adder)
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Acanthorhynchus superciliosus (western spinebill)
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Acanthostracion polygonius (Cowfish)
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Barbara L. Lundrigan (photographer; copyright holder; identification)
Identification
Acanthurus leucosternon (Blue surgeonfish)
Acanthuridae (Surgeonfishes, tangs, unicornfishes)
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Caption
Powderblue surgeonfish (Acanthurus leucosternon) showing the location of the

"scalpel", a modified spine on the caudal peduncle.
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Identification
Acanthurus dussumieri (Dussumier's surgeonfish)
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medium - large
Date Taken
November 2008
Location
Near Cotton Bay, Rodrigues Island, Mauritius
Caption
Eyestripe surgeonfish or Poisson Chirurgien oeil ray (Acanthurus dussumieri).
Identification
Acanthurus dussumieri (Dussumier's surgeonfish)
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Near Cotton Bay, Rodrigues Island, Mauritius.
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Eyestripe surgeonfish or Poisson Chirurgien oeil ray (Acanthurus dussumieri).
Identification
Acanthurus guttatus
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Date Taken
August 28, 2006
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Maui, Hawaii
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Allison Poor (photographer; identification), University of Michigan
Identification
Acanthurus lineatus (Blue banded surgeonfish)

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Identification
Acanthurus nigrofuscus (Blackspot surgeonfish)
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8/28/2006
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Maui, Hawaii
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Identification
Acanthurus sohal (Red Sea clown surgeon)
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Arabian tang (Acanthurus sohal)
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Identification
Acanthurus sohal (Red Sea clown surgeon)
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Arabian tang (Acanthurus sohal)

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Identification
Acanthurus triostegus (Convict surgeon)
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8/28/2006
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Maui, Hawaii
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Acanthurus triostegus (Convict surgeon)
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Vertebrata
vertebrates
Information
Pictures
Specimens
Sounds
Classification
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available for that taxon.
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directly to that feature.
Kingdom Animalia (animals)
Eumetazoa (metazoans)

Bilateria (bilaterally symmetrical animals)
Deuterostomia (deuterostomes)
Phylum Chordata (chordates)
Craniata (craniates)
(vertebrates)
Cephalaspidomorphi
Superclass
Gnathostomata (jawed
vertebrates)
Photos
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Taxonomy
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The Subphylum Vertebrata is a member of the Phylum Chordata. Here is the

complete "parentage" of Vertebrata:
Domain: Eukaryota - eukaryotes

o Kingdom: Animalia C. Linnaeus, 1758 - animals
Subkingdom: Bilateria (Hatschek, 1888) Cavalier-Smith,
1983 - bilaterians
Branch: Deuterostomia Grobben, 1908 -
Deuterostomes
Infrakingdom: Chordonia (Haeckel, 1874)
Cavalier-Smith, 1998
Phylum: Chordata Bateson, 1885 -
Chordates
Subphylum: Vertebrata
Cuvier, 1812 - Vertebrates
The Subphylum Vertebrata is further organized into finer groupings including:
Infraphylum (1): Gnathostomata

Superclass (2): Osteichthyes Tetrapoda
Series (3): Amniota Atherinomorpha Percomorpha
Class (14): Actinopterygii Agnatha Amphibia Aves
Cephalaspidomorphi Chondrichthyes Gastropoda Mammalia
Osteichthyes Placodermi Reptilia Sauropsida Secernentea
Synapsida
Classes
[ Back to top ]
Actinopterygii
The Actinopterygii (), or ray-finned fishes, constitute a class or sub-class of the

bony fishes. [more]
Agnatha
[more]
Amphibia
Amphibians are members of the class Amphibia (meaning "on both sides"), a
group of vertebrates whose forms include toads, frogs, newts, caecilians, and
salamanders. They are characterized as non-amniote ectothermic tetrapods. Most
amphibians undergo metamorphosis from a juvenile water-breathing form to an
adult air-breathing form, but some are paedomorphs that retain the juvenile water-
breathing form throughout life. Mudpuppies, for example, retain juvenile gills in
adulthood. The three modern orders of amphibians are Anura (frogs and toads),
Caudata (salamanders and newts), and Gymnophiona (caecilians, limbless
amphibians that resemble snakes), and in total they number approximately 6,500
species. Many amphibians lay their eggs in water. Amphibians are superficially
similar to reptiles, but reptiles are amniotes, along with mammals and birds.
Amphibians are ecological indicators, and in recent decades there has been a
dramatic decline in amphibian populations around the globe. Many species are
now threatened or extinct. The study of amphibians is called batrachology. [more]
Aves
[more]
Cephalaspidomorphi
Cephalaspidomorphs are a group of jawless fishes named for the cephalaspids, a

group of osteostracans. Most biologists regard this taxon as extinct, but the name
is sometimes used in the classification of lampreys because lampreys were once
thought to be related to cephalaspids. If lampreys are included, they would extend
the known range of the group from the Silurian and Devonian periods to the
present day. [more]
Chondrichthyes
Chondrichthyes (; from Greek ???d?- chondr- 'cartilage', ????? ichthys 'fish') or

cartilaginous fishes are jawed fish with paired fins, paired nares, scales, two-
chambered hearts, and skeletons made of cartilage rather than bone. The class is
divided into two subclasses: Elasmobranchii (sharks, rays and skates) and
Holocephali (chimaeras, sometimes called ghost sharks, which are sometimes
separated into their own class). [more]
Gastropoda
The Gastropoda or gastropods, more commonly known as snails and slugs, are
a large taxonomic class within the phylum Mollusca. The class Gastropoda
includes snails and slugs of all kinds and all sizes from microscopic to quite large.
There are huge numbers of sea snails and sea slugs, as well as freshwater snails
and freshwater limpets, and land snails and land slugs. [more]
Mammalia
Mammals (formally Mammalia ) are members of a class of air-breathing

vertebrate animals characterised by the possession of endothermy, hair, three
middle ear bones, and mammary glands functional in mothers with young. Most
mammals also possess sweat glands and specialised teeth, and the largest group of
mammals, the placentals, have a placenta which feeds the offspring during
gestation. The mammalian brain, with its characteristic neocortex, regulates
endothermic and circulatory systems, the latter featuring red blood cells lacking
nuclei and a large four-chambered heart maintaining the very high metabolism
rate they have. Mammals range in size from the 30?40 millimeter (1- to 1.5-inch)
Bumblebee Bat to the 33-meter (108-foot) Blue Whale. [more]
Osteichthyes
Osteichthyes (), also called bony fish, are a taxonomic group of fish that have
bony, as opposed to cartilaginous, skeletons. The vast majority of fish are
osteichthyes, which is an extremely diverse and abundant group consisting of over
29,000 species. It is the largest class of vertebrates in existence today.
Osteichthyes is divided into the ray-finned fish (Actinopterygii) and lobe-finned
fish (Sarcopterygii). The oldest known fossils of bony fish are about 420 million
years ago, which are also transitional fossils, showing a tooth pattern that is in
between the tooth rows of sharks and bony fishes. [more]
Placodermi
Placodermi (from the Greek p??? = plate and d???a = skin, literally "plate-
skinned") is a class of armoured prehistoric fish, known from fossils, which lived
from the late Silurian to the end of the Devonian Period. Their head and thorax
were covered by articulated armoured plates and the rest of the body was scaled or
naked, depending on the species. Placoderms were among the first jawed fish;
their jaws likely evolved from the first of their gill arches. A 380-million-year-old
fossil of one species represents the oldest known example of live birth. [more]
Reptilia
Reptiles (Reptilia) are members of a group of air-breathing, ectothermic (cold-

blooded) vertebrates which are characterized by laying shelled eggs (except for
some vipers and constrictor snakes that give live birth), and having skin covered
in scales and/or scutes. They are tetrapods, either having four limbs or being
descended from four-limbed ancestors. Modern reptiles inhabit every continent
with the exception of Antarctica. Reptiles originated around 320-310 million
years ago during the Carboniferous period, having evolved from advanced reptile-
like amphibians that became increasingly adapted to life on dry land. Four living
orders are typically recognized: [more]
Sauropsida
Sauropsida ("lizard faces") is a group of amniotes that includes all existing

reptiles and birds and their fossil ancestors, including the dinosaurs, the
immediate ancestors of birds. Sauropsida is distinguished from Synapsida, which
includes mammals and their fossil ancestors. [more]
Secernentea
Secernentea are the main class of nematodes, characterised by numerous and an

excretory system possessing lateral canals. Like all nematodes, they have no
circulatory or respiratory system. [more]
Synapsida
[more]
At least 4 species and subspecies belong to the Class Synapsida.

More info about the Class Synapsida may be found here.
References
[ Back to top ]
1. ^ a b Shu et al.; Luo, H-L.; Conway Morris, S.; Zhang, X-L.; Hu, S-X.;
Chen, L.; Han, J.; Zhu, M. et al (November 4, 1999). "Lower Cambrian
vertebrates from south China". Nature 402 (6757): 42?46. Bibcode
1999Natur.402...42S. doi:10.1038/46965.
2. ^ Jonathan E.M. Baillie, et al. (2004). "A Global Species Assessment".
World Conservation Union. http://www.iucn.org/bookstore/HTML-
books/Red%20List%202004/completed/table2.1.html.
3. ^ Kuraku et al.; Hoshiyama, D; Katoh, K; Suga, H; Miyata, T (December
1999). "Monophyly of Lampreys and Hagfishes Supported by Nuclear
DNA?Coded Genes". Journal of Molecular Evolution 49 (6): 729?35.
doi:10.1007/PL00006595. PMID 10594174.
4. ^ Stock, David; Whitt GS (7). "Evidence from 18S ribosomal RNA
sequences that lampreys and hagfishes form a natural group". Science 257
(5071): 787?9. doi:10.1126/science.1496398. PMID 1496398.
http://www.sciencemag.org/content/257/5071/787.short. Retrieved 22
November 2011.
5. ^ Nicholls, Henry (10 September 2009). "Mouth to Mouth". Nature 461
(7261): 164?166. doi:10.1038/461164a. PMID 19741680.
6. ^ Douglas Harper, Historian. "vertebrate". Online Etymology Dictionary.
Dictionary.com.. http://dictionary.reference.com/browse/vertebrate.
7. ^ Douglas Harper, Histo rian. "vertebra". Online Etymology Dictionary.
Dictionary.com.. http://dictionary.reference.com/browse/vertebra.
8. ^ Waggoner, Ben. "Vertebrates: More on Morphology". UCMP.
http://www.ucmp.berkeley.edu/vertebrates/vertmm.html. Retrieved 13
July 2011.
9. ^ a b c d e f Romer, A.S. (1949): The Vertebrate Body. W.B. Saunders,
Philadelphia. (2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970)
10. ^ Liem, Karel F.; Warren Franklin Walker (2001). Functional anatomy of
the vertebrates: an evolutionary perspective. Harcourt College Publishers.
p. 277. ISBN 9780030223693.
11. ^ Scott, Thomas (1996). Concise encyclopedia biology. Walter de Gruyter.
p. 542. ISBN 9783110106619.
12. ^ The American Naturalist (Essex Institute) 91: 287. 1957.
13. ^ Clack, J. A. (2002): Gaining ground: the origin and evolution of
tetrapods. Indiana University Press, Bloomington, Indiana. 369 pp
14. ^ a b Hildebrand, M. & Gonslow, G. (2001): Analysis of Vertebrate
Structure. 5th edition. John Wiley & Sons, Inc. New York
15. ^ "Keeping an eye on evolution". PhysOrg.com. 2007-12-03.
http://www.physorg.com/news115919015.html. Retrieved 2007-12-04.
16. ^ Hyperotreti - Hagfishes
17. ^ Waggoner, Ben. "Vertebrates: Fossil Record". UCMP.
http://www.ucmp.berkeley.edu/vertebrates/vertfr.html. Ret rieved 15 July
2011.
18. ^ Tim Haines, Paul Chambers (2005). The Complete Guide to Prehistoric
Life. Firefly Books.
19. ^ Encyclopaedia Britannica: a new survey of universal knowledge,
Volume 17. Encyclop?dia Britannica. 1954. p. 107.
20. ^ Berg, Linda R.; Eldra Pearl Solomon, Diana W. Martin (2004). Biology.
Cengage Learning. p. 599. ISBN 9780534492762.
21. ^ Cloudsley-Thompson, J. L. (2005). Ecology and behaviour of Mesozoic
reptiles. 9783540224211: Springer. p. 6.
22. ^ Andersen, Nils M?ller; Tom A. Weir (2004). Australian water bugs:
their biology and identification (Hemiptera-Heteroptera, Gerromorpha &
Nepomor pha). Apollo Books. p. 38. ISBN 9788788757781.
23. ^ Hildebran, M. & Gonslow, G. (2001): Analysis of Vertebrate Structure.
5th edition. John Wiley & Sons, Inc. New York, page 33: Comment: The
problem of naming sister groups
24. ^ a b Benton, Michael J. (2004-11-01). Vertebrate Palaeontology (Third
ed.). Blackwell Publishing. pp. 33, 455 pp.. ISBN 0632056371/978-
0632056378. http://palaeo.gly.bris.ac.uk/benton/vertclass.html.
25. ^ Janvier, Philippe. 1997. Vertebrata. Animals with backbones. Version
01 January 19 97 (under construction).
http://tolweb.org/Vertebrata/14829/1997.01.01 in The Tree of Life Web
Project, http://tolweb.org/
Further Reading
[ Back to top ]
Kardong, Kenneth V. (1998). Vertebrates: Comparative Anatomy,

Function, Evolution (second ed.). USA: McGraw-Hill. pp. 747 pp.. ISBN
0-07-115356-X/0-697-28654-1. http://www.amazon.com/Vertebrates-
Comparative-Anatomy-Function-Evolution/dp/0072909560.
Vertebrata (TSN 331030). Integrated Taxonomic Information System.
Retrieved on 6 August 2007.
Sources
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February 22, 2012.
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Last Revised: February 23, 2012
Vertebrates constitute the vast majority of living chordates, and they have evolved
an enormous variety of forms. The backbone of vertebrates protects the nerve cord
and serves as the axis of the internal skeleton. The skeleton provides strength and
rigidity to the body and is an attachment site for muscles. The vertebrae in the
middle region of the trunk give rise to pairs of ribs, which surround and protect
the internal organs. A cartilaginous or bony case encloses the brain. Bone is a
substance unique to vertebrates. It was formerly thought that vertebrates with
cartilage skeletons (cyclostomes and sharklike fishes) were descended from early
vertebrates that had not yet developed bone. However, very primitive fishes with
bone skeletons are known from the fossil record, so lack of bone is now believed
to be a degenerate rather than a primitive feature. All but the most primitive
vertebrates, known as jawless fishes, have jaws and paired appendages. The fishes
and, to a lesser extent, the amphibians and reptiles show a segmental arrangement
of the muscles of the body wall and of the nerves leading to them.
There are eight vertebrate classes. Four are aquatic, and may be grouped together
as the superclass Pisces, or fish; four are terrestrial or (in the case of amphibians)
semiterrestrial, and may be grouped as the superclass Tetrapoda, or four-footed
animals. Fishes breathe water by means of gills located in internal passages,
although they may also have lungs as supplementary air-breathing organs. Most
move through the water by weaving movements of the trunk and tail. All have
fins, and most have two sets of paired fins (pelvic and pectoral). Tetrapods breath
air, usually by means of lungs, and never have gills as adults, although the
amphibians go through a gilled, water-breathing stage. Except where the
appendages have been lost, as in snakes, all have two pairs of limbs, generally
used for locomotion; these are homologous to the pelvic and pectoral fins of fish.
Class Agnatha
The Agnatha, or jawless fishes, are the oldest known vertebrates. The only
surviving members of this class are the hagfish and lampreys, known as
cyclostomes. Cyclostomes have long, slender bodies with dorsal, ventral, and
caudal (tail) fins, all in the median plane. Although in their lack of jaws or paired
lateral appendages they represent a very primitive stage of vertebrate
development, the modern cyclostomes are highly adapted for their particular ways
of life. The hagfish is a specialized scavenger, and the lamprey is a parasite on
other fishes. The lamprey has a round mouth without skeletal supports, a rasping
tongue, and a single, dorsally located nostril. The gill passages are enlarged to
form pouches and are lined with gill filaments that serve as a surface for the
exchange of respiratory gases; in vertebrates the gill passages have acquired a
respiratory function. In cyclostomes, as in all fishes, water is taken in through the
mouth and expelled through the gill passages; as water passes over the thin-walled
gill filaments, dissolved oxygen diffuses into the blood, and carbon dioxide
diffuses out. The lamprey has a notochord extending from the head to near the tip
of the tail. A few cartilaginous blocks around the notochord constitute the bare
rudiments of a backbone; a cartilage framework supports the gill region, and there
is a rudimentary cartilage braincase. The meagerness of the skeleton is considered
a degenerate, not a primitive condition. The larva of the marine lamprey is a small
animal, resembling a lancelet, that uses the pharynx and gill passages for filter-
feeding. It metamorphoses into the adult form before migrating to the sea. The
extinct relatives of the cyclostomes, called ostracoderms, were jawless fishes with
bony armor and in some cases a well-developed bony skeleton.
Class Placodermi
The placoderms, an entirely extinct group of armored fishes, were the first jawed
vertebrates. Jaws enabled vertebrates to become predators, an important factor in
the later development of active, complex forms. The placoderms were also the
first vertebrates to have the two pairs of lateral appendages (supported by pelvic
and pectoral girdles) that characterized all later vertebrate groups. These primitive
paired fins gave rise to the pelvic and pectoral fins of modern fishes and to the
limbs of four-footed animals. The ostracoderms are thought to have given rise to
both the sharklike and the bony fishes.
Class Chondrichthyes
The almost exclusively marine sharks, rays, and chimaeras of the class
Chondrichthyes have skeletons made of cartilage. The mouth, equipped in most
sharks with numerous sharp teeth, is located on the underside of the head.
Passages called gill arches lead from the pharynx to the exterior and are lined with
gill filaments. The gill arches are supported by gill bars. Except in chimaeras, the
external gill slits are not covered and are conspicuous on the surface of the body.
The jaw consists of two distinct pieces; the upper part is not fused to the braincase
as in higher vertebrates. The tail is asymmetrical, curving upward in a shape
found in early fossil fishes and thought to be primitive. There is no lung or swim
bladder. The skin is studded with toothlike structures called denticles. Sharks have
typical vertebrate kidneys that excrete a very dilute urine consisting mostly of
water; presumably the earliest vertebrates (ancestral to sharks) evolved in
freshwater, where this function is necessary to maintain the correct concentration
of the physiologically important salts in the tissues against the tendency for them
to be diluted by the inward diffusion of water. In marine species, on the other
hand, it is necessary to prevent the concentration of those salts from increasing.
Although the kidneys of sharks pump out water, their body fluids contain
ammonia in concentrations high enough to make the osmotic pressure equal to
that of seawater; this prevents the inward diffusion of salts. Sharks have internal
fertilization and lay large eggs, well supplied with yolk and protected by leathery
shells. In a few species the eggs are hatched within the body.
Class Osteichthyes
The bony fish of the class Osteichthyes are the predominant class of living fishes.
In this group the bony skeleton has been retained and lungs and swim bladders
have evolved. Early bony fishes evolved in freshwater under conditions of
periodic drought and stagnation and developed an internal, moisture-retaining
organ, the lung, for gas exchange. Those fishes gave rise to two lines of
descendants.
Members of one line, the fleshy-finned fish, had thick fins with supporting bones,
used for crawling. The only survivors of that group are the coelacanth, or lobefin,
which has a vestigial lung and crawls on the seafloor, and the freshwater
lungfishes of drought-ridden areas, which can crawl over land in search of water
and even live out of water for several years. Early fleshy-finned fish gave rise to
the first land vertebrates, the amphibians.
The second line, the ray-finned fish, constitutes the predominant modern group.
Ray-finned fish are highly specialized for aquatic life; they have developed thin,
lightweight fins supported by slender rays, and used only for balance and steering.
The lung, a ventral outpocketing of the pharynx, was no longer necessary as these
fish invaded freshwaters and oceans throughout the world; it shifted to a dorsal
position and evolved into a hydrostatic organ called the swim bladder, or air float.
The swim bladder, along with the strong, lightweight skeletal construction, makes
ray-finned fishes much lighter-bodied than sharks. The gill passages of ray-finned
fishes resemble those of sharks, but have a bony covering, called the operculum,
over the external gill slits. Ray-fins have a typical vertebrate kidney which, in
freshwater forms, maintains the proper salt concentration in the tissues by
excreting excess water. In the marine forms the activity of the kidney is offset by
the activity of salt-secreting glands; in addition, the kidney may be modified so as
to produce a more concentrated urine. The heart, like that of sharks, has two
chambers, and there is no separation of oxygenated and deoxygenated blood in the
circulatory system. A few primitive ray-fins (the sturgeon, the paddle fish, and the
bowfin) have asymmetrical tails and thick scales regarded as primitive in
construction.
The higher ray-fins, or teleosts, have more or less symmetrical tail fins extending
above and below the vertebral column, and typical fish scales made of very thin
layers of bone. Most marine teleosts produce enormous numbers of small eggs
that are externally fertilized and float in plankton; only a few of these survive. In
many species there is a larval stage that is quite dissimilar to the adult. Teleosts
have evolved a tremendous variety of forms and occupy very diverse ecological
niches, both freshwater and marine.
Class Amphibia
The amphibians, the first vertebrates to have limbs, evolved during the Devonian
period. They are only partially terrestrial: Their externally fertilized eggs are laid
in freshwater, and they go through a gilled, aquatic larval stage (the tadpole stage)
before metamorphosing into land-living adults. The skin of the adult is water-
permeable, and the animal must live in a moist environment to prevent
desiccation. The adult usually breathes by means of lungs, although some breathe
directly through the skin. The heart is a three-chambered structure that creates a
partial separation between oxygenated blood, destined for the body tissues, and
depleted blood, destined for the lungs; this provides better oxygenation than does
a system in which the two kinds of blood mix. There are only three groups of
amphibians living today. The salamanders are closest to the basic amphibian stock
in form and in method of locomotion. Although supported by limbs, they move
with a wriggling motion similar to that of a fish. The frogs and toads are
specialized for jumping, with long, muscular hind legs, while the tropical
caecilians are burrowing forms that have lost all but vestigial traces of their limbs.
Class Reptilia
The reptiles, which evolved from amphibians during the Carboniferous period,
were the first vertebrate group to become entirely independent of water. This was
made possible by the development of a scaly, water-resistant skin and of the
terrestrial, or amniote, type of egg found in all higher land vertebrates. The
amniote egg has an elaborate series of internal membranes (one of which is called
the amnion) surrounding a pool of liquid in which the embryo develops; the
membranes prevent desiccation and allow inward diffusion of oxygen. Reptilian
eggs have porous shells and large amounts of yolk. Fertilization is internal. In
most cases the eggs are laid unhatched; in a few species they are retained and
hatched in the body. Reptiles, including such forms as turtles and sea snakes that
have returned to an aquatic life, are air-breathing at all stages, and nearly all lay
their eggs on land. Gill passages appear, as in birds and mammals, only in the
embryo.
During the Mesozoic era, reptiles were exceedingly diverse and numerous. The
reptilian dinosaurs included the largest terrestrial animals that have ever lived, as
well as many smaller forms. There were also flying and aquatic reptiles. With the
rise of the early mammals the decline of the reptiles began. The only large and
successful modern group of reptiles is the order of lizards and snakes. Snakes are
descended from lizards, but have lost their limbs. Reptiles, like fish and
amphibians, are cold-blooded, that is, they have little ability to regulate their body
temperature, which approaches that of the environment. The reptiles gave rise to
the two warm-blooded vertebrate groups, the birds and the mammals.
Class Aves
The birds evolved from reptiles in the Jurassic period. Their front limbs are
modified into wings, and the breastbone is greatly enlarged to support flight
muscles. They have an insulating covering of feathers, which has been an
important factor in their ability to regulate body temperature. The other advance
that enabled birds to become warm-blooded was the evolution of a four-
chambered heart, making the circulatory system a complete double circuit:
oxygenated blood is pumped from the lungs to the tissues, and deoxygenated
blood is pumped from the tissues to the lungs. The only major group besides
insects to invade the air, birds are much less restricted by external temperature
requirements than cold-blooded animals, and they have spread throughout every
part of the world. They live in many kinds of habitat and have evolved a diversity
of forms. Some have become flightless terrestrial animals, while others are
aquatic, using their wings for swimming instead of or in addition to flying.
Fertilization is internal. The eggs of birds are similar to those of reptiles, but
parental care of the eggs and young is highly developed.
Class Mammalia
The mammals also arose from reptiles in the Jurassic period and are now the
dominant form of terrestrial vertebrate life. Like the birds, they have a four-
chambered heart and a double-circuit circulatory system and are able to regulate
body temperature. In the case of mammals the insulating covering is provided by
hair, a feature unique to the class, although in a few forms (particularly in marine
species) nearly all the hair is lost, and insulation is provided by fat. A second
distinguishing characteristic of mammals is the production of milk by the females
for the nourishment of the young. All mammals have internal fertilization, and all
but the most primitive (the egg-laying monotremes of Australia) bear live young.
The mammalian egg contains little yolk. In the marsupials the young are born at
an extremely undeveloped stage and continue to develop in a milk-supplied
pouch. In the vastly more numerous placental mammals nourishment is passed
from the circulatory system of the mother to that of the embryo by means of a
placenta, and the young are born well-developed. Most mammals have highly
evolved sense organs and larger brains than other vertebrates. As a group they
display great adaptability to a variety of conditions and have spread to all regions
of the world.
The earliest placental mammals were small animals of the insectivore type, but
adaptive radiation has resulted in great diversity of forms and ways of life. Some
mammals are predators; others are herbivores with specialized digestive systems.
Some have taken up an aquatic existence and a few marine forms (whales and
sirenians) even give birth at sea. Members of one group, the bats, have developed
membranous wings supported by elongated fingers and lead an aerial existence.
The primates, the group that includes humans, are fairly close to the original
mammalian type in general structure (for example, they have five fingers and toes
and walk flat on the sole of the foot), but they have undergone great evolutionary
advances in the development of the brain, vision, and manual dexterity.
Sections in this article:

Introduction
Subphylum Urochordata
Subphylum Cephalochordata
Bibliography
The Columbia Electronic Encyclopedia, 6th ed. Copyright 2007, Columbia

University Press. All rights reserved.
Read more: Chordata: Subphylum Vertebrata Infoplease.com

http://www.infoplease.com/ce6/sci/A0857332.html#ixzz1sV9mfSN6

Etymology: Vertebrates (

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Vertebrates (pronounced /vrtbrts/) are animals that are members of the

subphylum Vertebrata (chordates with backbones and spinal columns).

[edit] Anatomy and morphology

[edit] Vertebral column

The defining characteristic of a vertebrate is the vertebral column, in which the

Gill arches bearing gills in a pike

[edit] Central nervous system

[edit] Evolutionary history

The early vertebrate Haikouichthys

Acanthostega, a fish-like early labyrinthodont.

[edit] Mesozoic vertebrates

The reptiles appeared from labyrinthodonts in the subsequent Carboniferous

[edit] After the Mesozoic

[edit] Traditional classification

Traditional spindle diagram of the evolution of the vertebrates at class level

Conventional classification has living vertebrates grouped into seven classes

[edit] Phylogenetic relationships

In phylogenetic taxonomy, the relationships between animals are not typically

The cladogram presented below is based on studies compiled by Philippe Janvier

[edit] See also

Animals with backbones

The main characteristics supporting the nodes of this phylogeny are:

Node 1: Mineralized exoskeleton, sensory-line canals and grooves

Containing group: Craniata

The Vertebrata, or vertebrates, is a very diverse group, ranging from lampreys to

Metamerically arranged endoskeletal elements flanking the spinal cord. There

There are many other vertebrate characteristics, both anatomical and

Discussion of Phylogenetic Relationships

Other Names for Vertebrata

Forey, P. L. (1984). Yet more reflections on agnathan-gnathostome relationships.

Hardisty, M. W. (1982). Lampreys and hagfishes: Analysis of cyclostome

Janvier, P. (1996a). Early vertebrates. Oxford Monographs in Geology and

Janvier, P. (1996b). The dawn of the vertebrates: characters versus common

Lvtrup, S. (1977). The Phylogeny of Vertebrata. Wiley, New York.

Wang, N. Z. (1991). Two new Silurian galeaspids (Jawless craniates) from

Information on the Internet

DigiMorph. The Digital Morphology library is a dynamic archive of information

Scientific Name Selene vomer

Specimen Condition Dead Specimen

Body Part skeleton

Copyright 1995 Tierney Thys

About This Page

Page copyright 1997 Philippe Janvier

Page: Tree of Life Vertebrata. Animals with backbones.

integument of two divisions, including an outer epidermis and an inner

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