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45-630: The formation preserves some fossils, including a diverse assemblage of microvertebrates (tiny fragments of vertebrate fossils) from a quarry near Moncure, NC . Palynology and general geological characteristics suggest that the Cumnock Formation is equivalent in age to the Cow Branch Formation of the Dan River / Danville basin on the Virginia -North Carolina border. It has also been compared to

90-491: 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

135-417: 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

180-525: 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

225-589: 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

270-407: A single orifice for excreting both solids and liquids and, in females, a separate vagina for reproduction. Female placental mammals have completely separate orifices for defecation , urination , and reproduction; males have one opening for defecation and another for both urination and reproduction , although the channels flowing to that orifice are almost completely separate. The development of

315-594: A specific stratigraphic formation in North Carolina is a stub . You can help Misplaced Pages by expanding it . This article related to the Triassic period is a stub . You can help Misplaced Pages by expanding it . Vertebrate Ossea Batsch, 1788 Vertebrates ( / ˈ v ɜːr t ə b r ɪ t s , - ˌ b r eɪ t s / ) are animals with a backbone or spine, consisting of vertebrae and intervertebral discs . The vertebrae are irregular bones , and

360-440: 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 :

405-617: 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

450-548: 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

495-474: 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

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540-623: 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

585-458: 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

630-562: 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

675-669: The Stockton and Lockatong formations of the Newark Basin in the northeastern United States. Some authors even reduce the Cumnock Formation to a member of the Lockatong Formation, in a broad interpretation of the latter. The Cumnock Formation is probably early Norian in age based on comparisons to similar strata in the eastern United States, as well as early sections of the Chinle Formation further west. This article about

720-858: The Tree of Life Web Project and Delsuc et al., and complemented (based on, and ). A dagger (†) denotes an extinct clade , whereas all other clades have living descendants . Hyperoartia ( lampreys ) [REDACTED] Myxini ( hagfish ) [REDACTED] † Euconodonta [REDACTED] † Myllokunmingiida [REDACTED]   † Pteraspidomorphi [REDACTED] † Thelodonti [REDACTED] † Anaspida [REDACTED] † Galeaspida [REDACTED] † Pituriaspida [REDACTED] † Osteostraci [REDACTED]   † Antiarchi [REDACTED] † Petalichthyida [REDACTED] † Arthrodira [REDACTED] † Ptyctodontida [REDACTED] † Entelognathus   [REDACTED] Anus In mammals , invertebrates and most fish ,

765-406: The anus ( pl. : anuses or ani ; from Latin , 'ring' or 'circle') is the external body orifice at the exit end of the digestive tract ( bowel ), i.e. the opposite end from the mouth . Its function is to facilitate the expulsion of wastes that remain after digestion . Bowel contents that pass through the anus include the gaseous flatus and the semi-solid feces , which (depending on

810-410: The archenteron , the first phase in the growth of the gut . In deuterostomes, the original dent becomes the anus while the gut eventually tunnels through to make another opening, which forms the mouth. The protostomes were so named because it was thought that in their embryos the dent formed the mouth first ( proto– meaning "first") and the anus was formed later at the opening made by the other end of

855-400: 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

900-462: 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

945-565: 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,

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990-410: 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

1035-432: 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

1080-421: The anus is typically controlled by muscular sphincters , and failure to stop unwanted passages results in fecal incontinence . Amphibians , reptiles and birds use a similar orifice (known as the cloaca ) for excreting liquid and solid wastes, for copulation and egg-laying . Monotreme mammals also have a cloaca, which is thought to be a feature inherited from the earliest amniotes . Marsupials have

1125-405: The anus was an important stage in the evolution of multicellular animals. It appears to have happened at least twice, following different paths in protostomes and deuterostomes . This accompanied or facilitated other important evolutionary developments: the bilaterian body plan , the coelom , and metamerism , in which the body was built of repeated "modules" which could later specialize, such as

1170-473: 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

1215-427: 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

1260-409: 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

1305-551: 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

1350-408: 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

1395-483: 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

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1440-421: 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

1485-400: 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

1530-651: 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 ,

1575-401: The heads of most arthropods , which are composed of fused, specialized segments. In comb jellies , there are species with one and sometimes two permanent anuses, species like the warty comb jelly grows an anus, which then disappear when it is no longer needed. In animals at least as complex as an earthworm , the embryo forms a dent on one side, the blastopore , which deepens to become

1620-443: 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,

1665-430: 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

1710-407: 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

1755-460: 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

1800-409: 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

1845-591: 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

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1890-644: 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

1935-633: 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

1980-402: 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,

2025-493: The type of animal) include: indigestible matter such as bones , hair pellets , endozoochorous seeds and digestive rocks ; residual food material after the digestible nutrients have been extracted, for example cellulose or lignin ; ingested matter which would be toxic if it remained in the digestive tract; excreted metabolites like bilirubin -containing bile ; and dead mucosal epithelia or excess gut bacteria and other endosymbionts . Passage of feces through

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