50-485: Carnivoramorpha ("carnivoran-like forms") is a clade of placental mammals of clade Pan-Carnivora from mirorder Ferae , that includes the modern order Carnivora and its extinct stem-relatives. The common feature for members of this clade is the presence of the carnassial teeth. The carnassial teeth of the Carnivoramorpha are upper premolar P4 and lower molar m1. Recent phylogenetic studies indicate that
100-549: A backbone or spine, consisting of vertebrae and intervertebral discs . The vertebrae are irregular bones , and the intervertebral discs are of fibrocartilage . The vertebral column surrounds and protects the spinal cord . The other feature unique to vertebrates is the presence of a cranium , or skull. The vertebrates make up the subphylum Vertebrata with some 65,000 species in the phylum Chordata . The vertebrates include mammals , birds , amphibians , and various classes of reptiles and fish . Classes of fish include
150-491: A vestigial coccyx . A dorsal nerve cord , which folds and fuses into a hollow neural tube during embryonic development and eventually gives rise to the brain and spinal cord , runs more dorsally to the axial endoskeleton, enclosed by neural arches , with a fore-end enlargement within a skeletonized braincase . This provides an alternative name for vertebrates, the craniates . All vertebrate embryos develop transient pharyngeal arches ; in fish, these develop into
200-479: A "ladder", with supposedly more "advanced" organisms at the top. Taxonomists have increasingly worked to make the taxonomic system reflect evolution. When it comes to naming , this principle is not always compatible with the traditional rank-based nomenclature (in which only taxa associated with a rank can be named) because not enough ranks exist to name a long series of nested clades. For these and other reasons, phylogenetic nomenclature has been developed; it
250-417: A central nervous system arising from a single nerve cord dorsal to the gut tube , headed by a series of (typically paired) brain vesicles, is unique to vertebrates. This is in stark contrast to invertebrates with well-developed central nervous systems such as arthropods and cephalopods , which have an often ladder-like ventral nerve cord made of paired segmental ganglia on the opposite ( ventral ) side of
300-623: A clade can be described based on two different reference points, crown age and stem age. The crown age of a clade refers to the age of the most recent common ancestor of all of the species in the clade. The stem age of a clade refers to the time that the ancestral lineage of the clade diverged from its sister clade. A clade's stem age is either the same as or older than its crown age. Ages of clades cannot be directly observed. They are inferred, either from stratigraphy of fossils , or from molecular clock estimates. Viruses , and particularly RNA viruses form clades. These are useful in tracking
350-525: A hollow neural tube ) running along the dorsal aspect of the notochord . Of particular importance and unique to vertebrates is the presence of neural crest cells, which are progenitor cells critical to coordinating the functions of cellular components. Neural crest cells migrate through the body from the dorsal nerve cord during development, initiate the formation of neuronal ganglia and various special sense organs. The peripheral nervous system forms when neural crest cells branch out laterally from
400-589: A protein related to ubiquitin carboxyl-terminal hydrolase are exclusively shared by all vertebrates and reliably distinguish them from all other animals. A specific relationship between vertebrates and tunicates is strongly supported by two CSIs found in the proteins Rrp44 (associated with the exosome complex ) and serine palmitoyltransferase . These are exclusively shared by species from these two subphyla, but not cephalochordates . This indicates hat vertebrates are more closely related to tunicates than cephalochordates. The "Notochordata hypothesis" suggested that
450-422: A revised taxonomy based on a concept strongly resembling clades, although the term clade itself would not be coined until 1957 by his grandson, Julian Huxley . German biologist Emil Hans Willi Hennig (1913–1976) is considered to be the founder of cladistics . He proposed a classification system that represented repeated branchings of the family tree, as opposed to the previous systems, which put organisms on
500-429: A suffix added should be e.g. "dracohortian". A clade is by definition monophyletic , meaning that it contains one ancestor which can be an organism, a population, or a species and all its descendants. The ancestor can be known or unknown; any and all members of a clade can be extant or extinct. The science that tries to reconstruct phylogenetic trees and thus discover clades is called phylogenetics or cladistics ,
550-438: Is a stub . You can help Misplaced Pages by expanding it . Clade In biological phylogenetics , a clade (from Ancient Greek κλάδος (kládos) 'branch'), also known as a monophyletic group or natural group , is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree . In the taxonomical literature, sometimes
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#1732852014565600-499: Is also used with a similar meaning in other fields besides biology, such as historical linguistics ; see Cladistics § In disciplines other than biology . The term "clade" was coined in 1957 by the biologist Julian Huxley to refer to the result of cladogenesis , the evolutionary splitting of a parent species into two distinct species, a concept Huxley borrowed from Bernhard Rensch . Many commonly named groups – rodents and insects , for example – are clades because, in each case,
650-440: Is found in invertebrate chordates such as lancelets (a sister subphylum known as the cephalochordates ), though it lacks eyes and other complex special sense organs comparable to those of vertebrates. Other chordates do not show any trends towards cephalization. The rostral end of the neural tube is expanded by a thickening of the walls and expansion of the central canal of spinal cord into three primary brain vesicles :
700-471: Is in turn included in the mammal, vertebrate and animal clades. The idea of a clade did not exist in pre- Darwinian Linnaean taxonomy , which was based by necessity only on internal or external morphological similarities between organisms. Many of the better known animal groups in Linnaeus's original Systema Naturae (mostly vertebrate groups) do represent clades. The phenomenon of convergent evolution
750-515: Is responsible for many cases of misleading similarities in the morphology of groups that evolved from different lineages. With the increasing realization in the first half of the 19th century that species had changed and split through the ages, classification increasingly came to be seen as branches on the evolutionary tree of life . The publication of Darwin's theory of evolution in 1859 gave this view increasing weight. In 1876 Thomas Henry Huxley , an early advocate of evolutionary theory, proposed
800-489: Is still controversial. As an example, see the full current classification of Anas platyrhynchos (the mallard duck) with 40 clades from Eukaryota down by following this Wikispecies link and clicking on "Expand". The name of a clade is conventionally a plural, where the singular refers to each member individually. A unique exception is the reptile clade Dracohors , which was made by haplology from Latin "draco" and "cohors", i.e. "the dragon cohort "; its form with
850-617: The Cephalochordata is the sister taxon to Craniata (Vertebrata). This group, called the Notochordata, was placed as sister group to the Tunicata (Urochordata). Studies since 2006 analyzing large sequencing datasets however strongly support Olfactores (tunicates + vertebrates) as a clade, and hence the placement of Cephalochordata as sister-group to Olfactores (known as the " Olfactores hypothesis "). The following cladogram summarizes
900-548: The Chengjiang biota and lived about 518 million years ago. These include Haikouichthys , Myllokunmingia , Zhongjianichthys , and probably Haikouella . 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. All of these early vertebrates lacked jaws in the common sense and relied on filter feeding close to
950-474: The Izu–Ogasawara Trench at a depth of 8,336 metres (27,349 feet). Many fish varieties are the main predators in most of the world's freshwater and marine water bodies . The rest of the vertebrate species are tetrapods, a single lineage that includes amphibians (with roughly 7,000 species); mammals (with approximately 5,500 species); and reptiles and birds (with about 20,000 species divided evenly between
1000-623: The Jurassic . The Cenozoic world saw great diversification of bony fishes, amphibians, reptiles, birds and mammals. Over half of all living vertebrate species (about 32,000 species) are fish (non-tetrapod craniates), a diverse set of lineages that inhabit all the world's aquatic ecosystems, from the Tibetan stone loach ( Triplophysa stolickai ) in western Tibetan hot springs near Longmu Lake at an elevation of 5,200 metres (17,100 feet) to an unknown species of snailfish (genus Pseudoliparis ) in
1050-458: The Latin vertebratus ("jointed"), from vertebra meaning "joint", in turn from Latin vertere to turn. All vertebrates are built along the basic chordate body plan of five shared characteristics . These are a rigid axial skeleton that includes a vertebral column developed around an elastic or flexible notochord , and a skull. The notochord stretches out for almost the whole length of
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#17328520145651100-594: The Latin form cladus (plural cladi ) is used rather than the English form. Clades are the fundamental unit of cladistics , a modern approach to taxonomy adopted by most biological fields. The common ancestor may be an individual, a population , or a species ( extinct or extant ). Clades are nested, one in another, as each branch in turn splits into smaller branches. These splits reflect evolutionary history as populations diverged and evolved independently. Clades are termed monophyletic (Greek: "one clan") groups. Over
1150-562: The Placodermi and the Acanthodii , both considered paraphyletic . Other ways of classifying the vertebrates have been devised, particularly with emphasis on the phylogeny of early amphibians and reptiles. An example based on Janvier (1981, 1997), Shu et al. (2003), and Benton (2004) is given here († = extinct ): While this traditional classification is orderly, most of the groups are paraphyletic , i.e. do not contain all descendants of
1200-400: The blue whale , at up to 33 m (108 ft). With only one exception, the defining characteristics of a vertebrate are the vertebral column , and the skull . The embryonic notochord is replaced by a segmented series of irregular bones known as vertebrae . The vertebrae are separated by intervertebral discs of fibrocartilage , which are remnants of the notochord. Hagfish are
1250-462: The branchial arches that support the gills . An organ in the pharynx , the endostyle , which assists in filter feeding in adult cephalochordates and urochordates , and in the larvae of lampreys , has evolved into the thyroid in most vertebrates. Vertebrates vary in body length ranging from the frog species Brachycephalus pulex , a Brazilian flea toad, with a minimum adult snout–vent length of 6.45 millimetres (0.254 in) to
1300-565: The genetics of organisms. Phylogenetic classification is based solely on phylogeny . Evolutionary systematics gives an overview; phylogenetic systematics gives detail. The two systems are thus complementary rather than opposed. Conventional classification has extant vertebrates grouped into seven classes based on traditional interpretations of gross anatomical and physiological traits. The commonly held classification lists three classes of fish and four of tetrapods . In addition to these, there are two classes of extinct armoured fishes,
1350-410: The prosencephalon ( forebrain ), mesencephalon ( midbrain ) and rhombencephalon ( hindbrain ), which are further differentiated in the various vertebrate groups. Two laterally placed retinas and optical nerves form around outgrowths from the midbrain, except in hagfish which may have secondarily lost the structures. The forebrain is more well-developed in most tetrapods and subdivided into
1400-432: The telencephalon and diencephalon , while the midbrain dominates in fish and some salamanders . In vertebrates with paired appendages, especially tetrapods, a pair of secondary enlargements of the hindbrain become the cerebella , which modulate complex motor coordinations . The brain vesicles are usually bilaterally symmetrical , giving rise to the paired cerebral hemispheres in mammals . The resultant anatomy of
1450-473: The body of a chordate , it lies dorsal to the gastrointestinal tract and provides flexible support to the body. The notochord remains in the jawless fish and cephalochordates (non-vertebrate chordates). In vertebrates it becomes replaced either wholly or in part by the vertebral column. The vertebral column typically continues beyond the rear orifice ( anus , or cloaca ) to form an elongated tail . Some vertebrates evolved to become tailless with only
1500-427: The brain (which itself is a fused cluster of segmental ganglia from the rostral metameres ). Molecular markers known as conserved signature indels (CSIs) in protein sequences have been identified and provide distinguishing criteria for the vertebrate subphylum. Specifically, 5 CSIs in the following proteins: protein synthesis elongation factor-2 , eukaryotic translation initiation factor 3 , adenosine kinase and
1550-409: The class's common ancestor. For instance, descendants of the first reptiles include modern reptiles, mammals and birds; 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
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1600-551: The demise of virtually all jawless fishes save for lampreys and hagfishes, as well as the Placodermi , a group of armoured fish that dominated the entirety of that period since the late Silurian as well as the eurypterids , dominant animals of the preceding Silurian , and the anomalocarids . By the middle of the Devonian, several droughts, anoxic events and oceanic competition led a lineage of sarcopterygii to leave water, eventually establishing themselves as terrestrial tetrapods in
1650-408: The dorsal nerve cord and migrate together with the mesodermal somites to innervate the various different structures that develop in the body. The vertebrates are the only chordate group with neural cephalization , and their neural functions are centralized towards a series of enlarged clusters in the head , which give rise to a brain . A slight swelling of the anterior end of the nerve cord
1700-483: The first gill arch pair evolved into the jointed jaws and form an additional oral cavity ahead of the pharynx . Research also suggests that the sixth branchial arch contributed to the formation of the vertebrate shoulder, which separated the head as a distinct part of the body. In amphibians and some primitive bony fishes, the larvae bear external gills , branching off from the gill arches. These are reduced in adulthood, their respiratory function taken over by
1750-421: The gill arches form during fetal development , and form 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 . The central nervous system of vertebrates is based on the embryonic dorsal nerve cord (which then flattens into a neural plate before folding and fusing over into
1800-546: The group consists of a common ancestor with all its descendant branches. Rodents, for example, are a branch of mammals that split off after the end of the period when the clade Dinosauria stopped being the dominant terrestrial vertebrates 66 million years ago. The original population and all its descendants are a clade. The rodent clade corresponds to the order Rodentia, and insects to the class Insecta. These clades include smaller clades, such as chipmunk or ant , each of which consists of even smaller clades. The clade "rodent"
1850-400: The gut tube, with a split brain stem circumventing the foregut around each side to form a brain on the dorsal side of the mouth . The higher functions of the vertebrate CNS are highly centralized towards the brain (particularly the forebrain), while the invertebrate CNS is significantly more decentralized with the segmental ganglia having substantial neural autonomy independent of
1900-651: The head, bordering the posterior margins of a series of crescentic openings from the pharynx to the outside. Each gill is supported by a cartilaginous or bony gill arch , which develop embryonically from pharyngeal arches . Bony fish have three pairs of gill arches, cartilaginous fish have five to seven pairs, while the primitive jawless fish have seven pairs. The ancestral vertebrates likely had more arches than seven, as some of their chordate relatives have more than 50 pairs of gill opens, although most, if not all, of these openings are actually involved in filter feeding rather than respiration . In jawed vertebrates ,
1950-443: The internal gills proper in fishes and by cutaneous respiration in most amphibians. While some amphibians such as axolotl retain the external gills into adulthood, the complex internal gill system as seen in fish apparently being irrevocably lost very early in the evolution of tetrapods , who evolved lungs (which are homologous to swim bladders ) to breathe air. While the more specialized terrestrial vertebrates lack gills,
2000-407: The jawless Agnatha , and the jawed Gnathostomata . The jawed fish include both the cartilaginous fish and the bony fish . Bony fish include the lobe-finned fish , which gave rise to the tetrapods , the animals with four limbs. Vertebrates make up less than five percent of all described animal species ; the rest are all invertebrates , that lack a backbone. The word 'vertebrate' derives from
2050-590: The last few decades, the cladistic approach has revolutionized biological classification and revealed surprising evolutionary relationships among organisms. Increasingly, taxonomists try to avoid naming taxa that are not clades; that is, taxa that are not monophyletic . Some of the relationships between organisms that the molecular biology arm of cladistics has revealed include that fungi are closer relatives to animals than they are to plants, archaea are now considered different from bacteria , and multicellular organisms may have evolved from archaea. The term "clade"
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2100-518: The latter term coined by Ernst Mayr (1965), derived from "clade". The results of phylogenetic/cladistic analyses are tree-shaped diagrams called cladograms ; they, and all their branches, are phylogenetic hypotheses. Three methods of defining clades are featured in phylogenetic nomenclature : node-, stem-, and apomorphy-based (see Phylogenetic nomenclature§Phylogenetic definitions of clade names for detailed definitions). The relationship between clades can be described in several ways: The age of
2150-460: 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. In phylogenetics , the relationships between animals are illustrated as a hierarchy known as a phylogenetic tree . The cladogram below is based on studies compiled by Philippe Janvier and others for
2200-409: The only extant vertebrate whose notochord persists and is not integrated or replaced by the vertebral column. In the sturgeon the endoskeleton is of cartilage and the vertebral column consists of mostly vertebrae of cartilage, without vertebral centrums , supported by the retained notochord. Most vertebrates are aquatic and carry out gas exchange via gills . The gills are carried right behind
2250-591: The relationships between the Olfactores (vertebrates and tunicates) and the Cephalochordata. Amphioxiformes (lancelets) [REDACTED] Tunicata /Urochordata ( sea squirts , salps , larvaceans ) [REDACTED] Vertebrata [REDACTED] Vertebrates originated during the Cambrian explosion , which saw a rise in organism diversity. The earliest known vertebrates belongs to
2300-644: The seabed. A vertebrate group of uncertain phylogeny, small eel-like conodonts , are known from microfossils of their paired tooth segments from the late Cambrian to the end of the Triassic. The first jawed vertebrates may have appeared in the late Ordovician (~445 mya) and became common in the Devonian period , often known as the "Age of Fishes". The two groups of bony fishes , the Actinopterygii and Sarcopterygii , evolved and became common. The Devonian also saw
2350-503: The spread of viral infections . HIV , for example, has clades called subtypes, which vary in geographical prevalence. HIV subtype (clade) B, for example is predominant in Europe, the Americas and Japan, whereas subtype A is more common in east Africa. Vertebrate Ossea Batsch, 1788 Vertebrates ( / ˈ v ɜːr t ə b r ɪ t s , - ˌ b r eɪ t s / ) are animals with
2400-633: The succeeding Carboniferous . Amniotes branched from amphibious tetrapods early in the Carboniferous period. The synapsid amniotes were dominant during the late Paleozoic, the Permian , while diapsid amniotes became dominant during the Mesozoic . In the sea, the teleosts and sharks became dominant. Mesothermic synapsids called cynodonts gave rise to endothermic mammals and diapsids called dinosaurs eventually gave rise to endothermic birds , both in
2450-578: The superfamily Miacoidea and family Miacidae are paraphyletic , with "miacids" being more closely related to carnivorans than to viverravids. In 2010 Flynn, Finarelli & Spaulding named a new clade Carnivoraformes within Carnivoramorpha, containing carnivorans and "miacids" but not viverravids. The authors defined Carnivoraformes as the clade containing Carnivora and all taxa that are more closely related to Carnivora (represented by Canis lupus ) than to viverravids (represented by Viverravus gracilis ). This prehistoric mammal -related article
2500-402: The two classes). Tetrapods comprise the dominant megafauna of most terrestrial environments and also include many partially or fully aquatic groups (e.g., sea snakes , penguins , cetaceans). 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,
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