Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct vertebrates (animals with vertebrae and their descendants). It also tries to connect, by using the evolutionary timeline , the animals of the past and their modern-day relatives.
120-527: The fossil record shows aspects of the meandering evolutionary path from early aquatic vertebrates to modern fish as well as mammals , birds , reptiles and amphibians , with a host of transitional fossils , though there are still large blank areas. The earliest known fossil vertebrates were heavily armored fish discovered in rocks from the Ordovician period about 485 to 444 Ma ( megaannum , million years ago), with jawed vertebrates emerging in
240-549: 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
360-438: A food chain who lose their prey. "Species coextinction is a manifestation of one of the interconnectednesses of organisms in complex ecosystems ... While coextinction may not be the most important cause of species extinctions, it is certainly an insidious one." Coextinction is especially common when a keystone species goes extinct. Models suggest that coextinction is the most common form of biodiversity loss . There may be
480-427: A jaw , hyoid and/or the middle ear ossicles . An iodine-concentrating organ called the endostyle , which functions as a filter feeding organ in aquatic animals 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
600-669: A nautilus to the Royal Society that was more than two feet in diameter, and morphologically distinct from any known living species. Hooke theorized that this was simply because the species lived in the deep ocean and no one had discovered them yet. While he contended that it was possible a species could be "lost", he thought this highly unlikely. Similarly, in 1695, Sir Thomas Molyneux published an account of enormous antlers found in Ireland that did not belong to any extant taxa in that area. Molyneux reasoned that they came from
720-415: A species or a population is the variety of genetic information in its living members. A large gene pool (extensive genetic diversity ) is associated with robust populations that can survive bouts of intense selection . Meanwhile, low genetic diversity (see inbreeding and population bottlenecks ) reduces the range of adaptions possible. Replacing native with alien genes narrows genetic diversity within
840-436: A viable population for species preservation and possible future reintroduction to the wild, through use of carefully planned breeding programs . The extinction of one species' wild population can have knock-on effects, causing further extinctions. These are also called "chains of extinction". This is especially common with extinction of keystone species . A 2018 study indicated that the sixth mass extinction started in
960-479: A Lazarus species when extant individuals were described in 2019. Attenborough's long-beaked echidna ( Zaglossus attenboroughi ) is an example of a Lazarus species from Papua New Guinea that had last been sighted in 1962 and believed to be possibly extinct, until it was recorded again in November 2023. Some species currently thought to be extinct have had continued speculation that they may still exist, and in
1080-503: A cascade of coextinction across the trophic levels . Such effects are most severe in mutualistic and parasitic relationships. An example of coextinction is the Haast's eagle and the moa : the Haast's eagle was a predator that became extinct because its food source became extinct. The moa were several species of flightless birds that were a food source for the Haast's eagle. Extinction as
1200-417: 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
1320-429: A common ancestor with modern horses. Pseudoextinction is much easier to demonstrate for larger taxonomic groups. A Lazarus taxon or Lazarus species refers to instances where a species or taxon was thought to be extinct, but was later rediscovered. It can also refer to instances where large gaps in the fossil record of a taxon result in fossils reappearing much later, although the taxon may have ultimately become extinct at
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#17327943031491440-399: A fact that was accepted by most scientists. The primary debate focused on whether this turnover caused by extinction was gradual or abrupt in nature. Cuvier understood extinction to be the result of cataclysmic events that wipe out huge numbers of species, as opposed to the gradual decline of a species over time. His catastrophic view of the nature of extinction garnered him many opponents in
1560-708: A higher risk of extinction and die out faster than less sexually dimorphic species, the least sexually dimorphic species surviving for millions of years while the most sexually dimorphic species die out within mere thousands of years. Earlier studies based on counting the number of currently living species in modern taxa have shown a higher number of species in more sexually dimorphic taxa which have been interpreted as higher survival in taxa with more sexual selection, but such studies of modern species only measure indirect effects of extinction and are subject to error sources such as dying and doomed taxa speciating more due to splitting of habitat ranges into more small isolated groups during
1680-525: 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
1800-405: A large range, a lack of individuals of both sexes (in sexually reproducing species), or other reasons. Pinpointing the extinction (or pseudoextinction ) of a species requires a clear definition of that species . If it is to be declared extinct, the species in question must be uniquely distinguishable from any ancestor or daughter species, and from any other closely related species. Extinction of
1920-607: A later point. The coelacanth , a fish related to lungfish and tetrapods , is an example of a Lazarus taxon that was known only from the fossil record and was considered to have been extinct since the end of the Cretaceous Period . In 1938, however, a living specimen was found off the Chalumna River (now Tyolomnqa) on the east coast of South Africa. Calliostoma bullatum , a species of deepwater sea snail originally described from fossils in 1844 proved to be
2040-594: A mathematical model that falls in all positions. By contrast, conservation biology uses the extinction vortex model to classify extinctions by cause. When concerns about human extinction have been raised, for example in Sir Martin Rees ' 2003 book Our Final Hour , those concerns lie with the effects of climate change or technological disaster. Human-driven extinction started as humans migrated out of Africa more than 60,000 years ago. Currently, environmental groups and some governments are concerned with
2160-456: A natural part of the evolutionary process. Only recently have extinctions been recorded and scientists have become alarmed at the current high rate of extinctions . Most species that become extinct are never scientifically documented. Some scientists estimate that up to half of presently existing plant and animal species may become extinct by 2100. A 2018 report indicated that the phylogenetic diversity of 300 mammalian species erased during
2280-441: A new mega-predator or by transporting animals and plants from one part of the world to another. Such introductions have been occurring for thousands of years, sometimes intentionally (e.g. livestock released by sailors on islands as a future source of food) and sometimes accidentally (e.g. rats escaping from boats). In most cases, the introductions are unsuccessful, but when an invasive alien species does become established,
2400-610: A population a higher chance in the short term of surviving an adverse change in conditions. Effects that cause or reward a loss in genetic diversity can increase the chances of extinction of a species. Population bottlenecks can dramatically reduce genetic diversity by severely limiting the number of reproducing individuals and make inbreeding more frequent. Extinction sometimes results for species evolved to specific ecologies that are subjected to genetic pollution —i.e., uncontrolled hybridization , introgression and genetic swamping that lead to homogenization or out-competition from
2520-589: 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
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#17327943031492640-407: A race of animals to become extinct. A series of fossils were discovered in the late 17th century that appeared unlike any living species. As a result, the scientific community embarked on a voyage of creative rationalization, seeking to understand what had happened to these species within a framework that did not account for total extinction. In October 1686, Robert Hooke presented an impression of
2760-502: A reduction in agricultural productivity. Furthermore, increased erosion contributes to poorer water quality by elevating the levels of sediment and pollutants in rivers and streams. Habitat degradation through toxicity can kill off a species very rapidly, by killing all living members through contamination or sterilizing them. It can also occur over longer periods at lower toxicity levels by affecting life span, reproductive capacity, or competitiveness. Habitat degradation can also take
2880-499: A result of climate change has been confirmed by fossil studies. Particularly, the extinction of amphibians during the Carboniferous Rainforest Collapse , 305 million years ago. A 2003 review across 14 biodiversity research centers predicted that, because of climate change, 15–37% of land species would be "committed to extinction" by 2050. The ecologically rich areas that would potentially suffer
3000-606: 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 ,
3120-479: A species (or replacement by a daughter species) plays a key role in the punctuated equilibrium hypothesis of Stephen Jay Gould and Niles Eldredge . In ecology , extinction is sometimes used informally to refer to local extinction , in which a species ceases to exist in the chosen area of study, despite still existing elsewhere. Local extinctions may be made good by the reintroduction of individuals of that species taken from other locations; wolf reintroduction
3240-439: A species may come suddenly when an otherwise healthy species is wiped out completely, as when toxic pollution renders its entire habitat unliveable; or may occur gradually over thousands or millions of years, such as when a species gradually loses out in competition for food to better adapted competitors. Extinction may occur a long time after the events that set it in motion, a phenomenon known as extinction debt . Assessing
3360-404: A species or group of species. "Just as each species is unique", write Beverly and Stephen C. Stearns , "so is each extinction ... the causes for each are varied—some subtle and complex, others obvious and simple". Most simply, any species that cannot survive and reproduce in its environment and cannot move to a new environment where it can do so, dies out and becomes extinct. Extinction of
3480-518: A subsequent report, IPBES listed unsustainable fishing, hunting and logging as being some of the primary drivers of the global extinction crisis. In June 2019, one million species of plants and animals were at risk of extinction. At least 571 plant species have been lost since 1750, but likely many more. The main cause of the extinctions is the destruction of natural habitats by human activities, such as cutting down forests and converting land into fields for farming. A dagger symbol (†) placed next to
3600-523: Is also evidence to suggest that this event was preceded by another mass extinction, known as Olson's Extinction . The Cretaceous–Paleogene extinction event (K–Pg) occurred 66 million years ago, at the end of the Cretaceous period; it is best known for having wiped out non-avian dinosaurs , among many other species. According to a 1998 survey of 400 biologists conducted by New York 's American Museum of Natural History , nearly 70% believed that
3720-444: Is an example of this. Species that are not globally extinct are termed extant . Those species that are extant, yet are threatened with extinction, are referred to as threatened or endangered species . Currently, an important aspect of extinction is human attempts to preserve critically endangered species. These are reflected by the creation of the conservation status "extinct in the wild" (EW) . Species listed under this status by
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3840-585: Is difficult to demonstrate unless one has a strong chain of evidence linking a living species to members of a pre-existing species. For example, it is sometimes claimed that the extinct Hyracotherium , which was an early horse that shares a common ancestor with the modern horse , is pseudoextinct, rather than extinct, because there are several extant species of Equus , including zebra and donkey ; however, as fossil species typically leave no genetic material behind, one cannot say whether Hyracotherium evolved into more modern horse species or merely evolved from
3960-417: Is estimated as 100 to 1,000 times "background" rates (the average extinction rates in the evolutionary time scale of planet Earth), faster than at any other time in human history, while future rates are likely 10,000 times higher. However, some groups are going extinct much faster. Biologists Paul R. Ehrlich and Stuart Pimm , among others, contend that human population growth and overconsumption are
4080-440: 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 :
4200-583: Is still used in works where systematic overview is essential, e.g. Benton (1998), Hildebrand and Goslow (2001) and Knobill and Neill (2006). While mostly seen in general works, it is also still used in some specialist works like Fortuny & al. (2011). Kingdom Animalia The oVert (openVertebrate) Thematic Collection Network (TCN) is a project that aims to generate and distribute high-resolution digital three-dimensional data for internal anatomy across vertebrate diversity. The project will CT-scan over 20,000 fluid-preserved specimens, representing more than 80% of
4320-592: Is the destruction of ocean floors by bottom trawling . Diminished resources or introduction of new competitor species also often accompany habitat degradation. Global warming has allowed some species to expand their range, bringing competition to other species that previously occupied that area. Sometimes these new competitors are predators and directly affect prey species, while at other times they may merely outcompete vulnerable species for limited resources. Vital resources including water and food can also be limited during habitat degradation, leading to extinction. In
4440-525: Is the most important determinant of genus extinction at background rates but becomes increasingly irrelevant as mass extinction arises. Limited geographic range is a cause both of small population size and of greater vulnerability to local environmental catastrophes. Extinction rates can be affected not just by population size, but by any factor that affects evolvability , including balancing selection , cryptic genetic variation , phenotypic plasticity , and robustness . A diverse or deep gene pool gives
4560-457: Is the termination of a taxon by the death of its last member . A taxon may become functionally extinct before the death of its last member if it loses the capacity to reproduce and recover. Because a species' potential range may be very large, determining this moment is difficult, and is usually done retrospectively. This difficulty leads to phenomena such as Lazarus taxa , where a species presumed extinct abruptly "reappears" (typically in
4680-635: 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
4800-559: 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
4920-483: The International Union for Conservation of Nature (IUCN) are not known to have any living specimens in the wild and are maintained only in zoos or other artificial environments. Some of these species are functionally extinct, as they are no longer part of their natural habitat and it is unlikely the species will ever be restored to the wild. When possible, modern zoological institutions try to maintain
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5040-474: 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
5160-595: The Jurassic , with crown-group birds ( Neornithes ) emerging in the Cretaceous between 100 Ma and 60 Ma. The K-Pg mass extinction wiped out many vertebrate clades, including the pterosaurs , plesiosaurs , mosasaurs and nearly all dinosaurs , leaving many ecological niches open. While therian mammals had already evolved in the Late Jurassic, they would rise to prominence in the Paleogene following
5280-578: 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
5400-485: The Late Pleistocene could take up to 5 to 7 million years to restore mammal diversity to what it was before the human era. Extinction of a parent species where daughter species or subspecies are still extant is called pseudoextinction or phyletic extinction. Effectively, the old taxon vanishes, transformed ( anagenesis ) into a successor, or split into more than one ( cladogenesis ). Pseudoextinction
5520-464: 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 synapomorphies . These are a rigid axial skeleton that includes a vertebral column developed around an elastic notochord . The notochord becomes the intervertebral discs , and runs dorsally to the gut tube along
5640-574: 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
5760-842: 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] Extinction Extinction
5880-604: The Triassic period (252 to 201 Ma). Lissamphibians , modern amphibians, likely arose around that time from temnospondyls. True mammals , derived from cynodont therapsids, showed up in the Middle Triassic around the same time as the dinosaurs , who emerged from a clade of archosaurs. At the same time, ray-finned fish diversified, leading to teleost fish dominating the seas. Ancestral birds ( Avialae ) like Archaeopteryx first evolved from dinosaurs during
6000-399: The blue whale , at up to 33 m (108 ft). With only one exception, the defining characteristic of all vertebrates is the vertebral column , in which the embryonic notochord found in all chordates is replaced by a segmented series of mineralized elements called vertebrae separated by fibrocartilaginous intervertebral discs , which are embryonic and evolutionary remnants of
6120-616: The fossil record ) after a period of apparent absence. More than 99% of all species that ever lived on Earth , amounting to over five billion species, are estimated to have died out. It is estimated that there are currently around 8.7 million species of eukaryotes globally, and possibly many times more if microorganisms , like bacteria , are included. Notable extinct animal species include non-avian dinosaurs , saber-toothed cats , dodos , mammoths , ground sloths , thylacines , trilobites , golden toads , and passenger pigeons . Through evolution , species arise through
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#17327943031496240-565: 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,
6360-410: 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
6480-410: The slender-billed curlew ( Numenius tenuirostris ), not seen since 2007. As long as species have been evolving, species have been going extinct. It is estimated that over 99.9% of all species that ever lived are extinct. The average lifespan of a species is 1–10 million years, although this varies widely between taxa. A variety of causes can contribute directly or indirectly to the extinction of
6600-488: The strata of the Paris basin. They saw alternating saltwater and freshwater deposits, as well as patterns of the appearance and disappearance of fossils throughout the record. From these patterns, Cuvier inferred historic cycles of catastrophic flooding, extinction, and repopulation of the earth with new species. Cuvier's fossil evidence showed that very different life forms existed in the past than those that exist today,
6720-432: 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
6840-609: The (at the time) contested theory of extinction of species. Thomas Jefferson is credited with initiating the science of vertebrate paleontology in the United States with the reading of a paper to the American Philosophical Society in Philadelphia in 1797. Jefferson presented fossil bones of a ground sloth found in a cave in western Virginia and named the genus ( Megalonyx ). The species
6960-639: The 20 biodiversity goals laid out by the Aichi Biodiversity Targets in 2010, only 6 were "partially achieved" by the deadline of 2020. The report warned that biodiversity will continue to decline if the status quo is not changed, in particular the "currently unsustainable patterns of production and consumption, population growth and technological developments". In a 2021 report published in the journal Frontiers in Conservation Science , some top scientists asserted that even if
7080-451: The Aichi Biodiversity Targets set for 2020 had been achieved, it would not have resulted in a significant mitigation of biodiversity loss. They added that failure of the global community to reach these targets is hardly surprising given that biodiversity loss is "nowhere close to the top of any country's priorities, trailing far behind other concerns such as employment, healthcare, economic growth, or currency stability." For much of history,
7200-618: The Earth is currently in the early stages of a human-caused mass extinction, known as the Holocene extinction . In that survey, the same proportion of respondents agreed with the prediction that up to 20% of all living populations could become extinct within 30 years (by 2028). A 2014 special edition of Science declared there is widespread consensus on the issue of human-driven mass species extinctions. A 2020 study published in PNAS stated that
7320-584: The North American moose and that the animal had once been common on the British Isles . Rather than suggest that this indicated the possibility of species going extinct, he argued that although organisms could become locally extinct, they could never be entirely lost and would continue to exist in some unknown region of the globe. The antlers were later confirmed to be from the extinct deer Megaloceros . Hooke and Molyneux's line of thinking
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#17327943031497440-522: The Paris basin, could be formed by a slow rise and fall of sea levels . The concept of extinction was integral to Charles Darwin 's On the Origin of Species , with less fit lineages disappearing over time. For Darwin, extinction was a constant side effect of competition . Because of the wide reach of On the Origin of Species , it was widely accepted that extinction occurred gradually and evenly (a concept now referred to as background extinction ). It
7560-429: The accumulation of slightly deleterious mutations , then a population will go extinct. Smaller populations have fewer beneficial mutations entering the population each generation, slowing adaptation. It is also easier for slightly deleterious mutations to fix in small populations; the resulting positive feedback loop between small population size and low fitness can cause mutational meltdown . Limited geographic range
7680-418: The axial endoskeleton (enclosed by protective skeletal extensions known as neural arches ), with a fore-end enlargement that is contained within a distinct skeletonized braincase (hence the alternative name for vertebrates, the craniates ). All vertebrate embryos develop transient pharyngeal arches , which in fish develop into the branchial arches that support the gills . Other vertebrate features are
7800-427: 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
7920-409: 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
8040-600: The consequences can be catastrophic. Invasive alien species can affect native species directly by eating them, competing with them, and introducing pathogens or parasites that sicken or kill them; or indirectly by destroying or degrading their habitat. Human populations may themselves act as invasive predators. According to the "overkill hypothesis", the swift extinction of the megafauna in areas such as Australia (40,000 years before present), North and South America (12,000 years before present), Madagascar , Hawaii (AD 300–1000), and New Zealand (AD 1300–1500), resulted from
8160-418: The contemporary extinction crisis "may be the most serious environmental threat to the persistence of civilization, because it is irreversible." Biologist E. O. Wilson estimated in 2002 that if current rates of human destruction of the biosphere continue, one-half of all plant and animal species of life on earth will be extinct in 100 years. More significantly, the current rate of global species extinctions
8280-476: The deliberate destruction of some species, such as dangerous viruses , and the total destruction of other problematic species has been suggested. Other species were deliberately driven to extinction, or nearly so, due to poaching or because they were "undesirable", or to push for other human agendas. One example was the near extinction of the American bison , which was nearly wiped out by mass hunts sanctioned by
8400-556: 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
8520-408: 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
8640-432: The early Carboniferous, with temnospondyls dominating the ecosystem and becoming the first land vertebrate megafauna. A lineage of reptiliomorphs developed a metabolism better suited for life exclusively on land, as well as a novel form of reproduction freeing them from the water: the amniotic egg , with full-fledged amniotes appearing in the mid-Carboniferous. Sharks and their holocephalian relatives flourished in
8760-403: The endangered wild water buffalo is most threatened with extinction by genetic pollution from the abundant domestic water buffalo ). Such extinctions are not always apparent from morphological (non-genetic) observations. Some degree of gene flow is a normal evolutionary process; nevertheless, hybridization (with or without introgression) threatens rare species' existence. The gene pool of
8880-591: The event of rediscovery would be considered Lazarus species. Examples include the thylacine , or Tasmanian tiger ( Thylacinus cynocephalus ), the last known example of which died in Hobart Zoo in Tasmania in 1936; the Japanese wolf ( Canis lupus hodophilax ), last sighted over 100 years ago; the American ivory-billed woodpecker ( Campephilus principalis ), with the last universally accepted sighting in 1944; and
9000-443: The extinction crisis. According to the International Union for Conservation of Nature (IUCN), 784 extinctions have been recorded since the year 1500, the arbitrary date selected to define "recent" extinctions, up to the year 2004; with many more likely to have gone unnoticed. Several species have also been listed as extinct since 2004. If adaptation increasing population fitness is slower than environmental degradation plus
9120-475: The extinction of species caused by humanity, and they try to prevent further extinctions through a variety of conservation programs. Humans can cause extinction of a species through overharvesting , pollution , habitat destruction , introduction of invasive species (such as new predators and food competitors ), overhunting, and other influences. Explosive, unsustainable human population growth and increasing per capita consumption are essential drivers of
9240-540: The field of zoology , and biology in general, and has also become an area of concern outside the scientific community. A number of organizations, such as the Worldwide Fund for Nature , have been created with the goal of preserving species from extinction. Governments have attempted, through enacting laws, to avoid habitat destruction, agricultural over-harvesting, and pollution . While many human-caused extinctions have been accidental, humans have also engaged in
9360-483: 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
9480-523: The following Silurian period (444 to 419 Ma) with the placoderms and acanthodians . The Devonian period (419 to 359 Ma) saw primitive air-breathing fish to develop limbs allowing them to walk on land, thus becoming the first terrestrial vertebrates, the stegocephalians . Romer's gap in the early Carboniferous period (359 to 299 Ma) left little of the early stegocephalians, but allowed vertebrates more adapted to life on land to flourish in their wake. Crown-group tetrapods appeared in
9600-435: The form of a physical destruction of niche habitats. The widespread destruction of tropical rainforests and replacement with open pastureland is widely cited as an example of this; elimination of the dense forest eliminated the infrastructure needed by many species to survive. For example, a fern that depends on dense shade for protection from direct sunlight can no longer survive without forest to shelter it. Another example
9720-421: 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
9840-400: 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
9960-580: The habitat retreat of taxa approaching extinction. Possible causes of the higher extinction risk in species with more sexual selection shown by the comprehensive fossil studies that rule out such error sources include expensive sexually selected ornaments having negative effects on the ability to survive natural selection , as well as sexual selection removing a diversity of genes that under current ecological conditions are neutral for natural selection but some of which may be important for surviving climate change. There have been at least five mass extinctions in
10080-676: The heaviest losses include the Cape Floristic Region and the Caribbean Basin . These areas might see a doubling of present carbon dioxide levels and rising temperatures that could eliminate 56,000 plant and 3,700 animal species. Climate change has also been found to be a factor in habitat loss and desertification . Studies of fossils following species from the time they evolved to their extinction show that species with high sexual dimorphism , especially characteristics in males that are used to compete for mating, are at
10200-447: The history of life on earth, and four in the last 350 million years in which many species have disappeared in a relatively short period of geological time. A massive eruptive event that released large quantities of tephra particles into the atmosphere is considered to be one likely cause of the " Permian–Triassic extinction event " about 250 million years ago, which is estimated to have killed 90% of species then existing. There
10320-475: The human era since the Late Pleistocene would require 5 to 7 million years to recover. According to the 2019 Global Assessment Report on Biodiversity and Ecosystem Services by IPBES , the biomass of wild mammals has fallen by 82%, natural ecosystems have lost about half their area and a million species are at risk of extinction—all largely as a result of human actions. Twenty-five percent of plant and animal species are threatened with extinction. In
10440-443: 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,
10560-431: The introduced ( or hybrid ) species. Endemic populations can face such extinctions when new populations are imported or selectively bred by people, or when habitat modification brings previously isolated species into contact. Extinction is likeliest for rare species coming into contact with more abundant ones; interbreeding can swamp the rarer gene pool and create hybrids, depleting the purebred gene pool (for example,
10680-407: 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
10800-432: The length of an animal, hence the common name of backbone. The axial endoskeleton typically continues beyond the anus / cloaca to form an elongated tail . Some vertebrates evolved to become tailless with only 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
10920-567: The living genera of vertebrates, at a network of digitization centers across the US. This collection of digital imagery and three-dimensional volumes will be open for exploration, download, and use to address questions related to the discovery of new species, documenting patterns of anatomical diversity and growth, and testing hypotheses of function and evolution. Vertebrate Ossea Batsch, 1788 Vertebrates ( / ˈ v ɜːr t ə b r ɪ t s , - ˌ b r eɪ t s / ) are animals with
11040-488: The main drivers of the modern extinction crisis. In January 2020, the UN's Convention on Biological Diversity drafted a plan to mitigate the contemporary extinction crisis by establishing a deadline of 2030 to protect 30% of the Earth's land and oceans and reduce pollution by 50%, with the goal of allowing for the restoration of ecosystems by 2050. The 2020 United Nations ' Global Biodiversity Outlook report stated that of
11160-460: 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
11280-434: The mass extinction and remain to this day, although squamates and birds still lead in diversity. One of the people who helped figure out the vertebrate progression was French zoologist Georges Cuvier (1769–1832), who realized that fossils found in older rock strata differed greatly from more recent fossils or modern animals. He published his findings in 1812 and, although he steadfastly refuted evolution , his work proved
11400-421: The modern understanding of extinction as the end of a species was incompatible with the prevailing worldview. Prior to the 19th century, much of Western society adhered to the belief that the world was created by God and as such was complete and perfect. This concept reached its heyday in the 1700s with the peak popularity of a theological concept called the great chain of being , in which all life on earth, from
11520-490: The name of a species or other taxon normally indicates its status as extinct. Examples of species and subspecies that are extinct include: A species is extinct when the last existing member dies. Extinction therefore becomes a certainty when there are no surviving individuals that can reproduce and create a new generation. A species may become functionally extinct when only a handful of individuals survive, which cannot reproduce due to poor health, age, sparse distribution over
11640-469: The natural course of events, species become extinct for a number of reasons, including but not limited to: extinction of a necessary host, prey or pollinator, interspecific competition , inability to deal with evolving diseases and changing environmental conditions (particularly sudden changes) which can act to introduce novel predators, or to remove prey. Recently in geological time, humans have become an additional cause of extinction of some species, either as
11760-410: The newly emerging school of uniformitarianism . Jean-Baptiste Lamarck , a gradualist and colleague of Cuvier, saw the fossils of different life forms as evidence of the mutable character of species. While Lamarck did not deny the possibility of extinction, he believed that it was exceptional and rare and that most of the change in species over time was due to gradual change. Unlike Cuvier, Lamarck
11880-411: The notochord. Hagfish are the only extant vertebrate whose notochord persists and is not integrated/ replaced by the vertebral column. A few vertebrates have secondarily lost this feature and retain the notochord into adulthood, such as the sturgeon . Most vertebrates are aquatic and carry out gas exchange via gills . The gills are carried right behind the head, bordering the posterior margins of
12000-399: The original population, thereby increasing the chance of extinction. Habitat degradation is currently the main anthropogenic cause of species extinctions. The main cause of habitat degradation worldwide is agriculture, with urban sprawl , logging, mining, and some fishing practices close behind. The degradation of a species' habitat may alter the fitness landscape to such an extent that
12120-702: The process of speciation —where new varieties of organisms arise and thrive when they are able to find and exploit an ecological niche —and species become extinct when they are no longer able to survive in changing conditions or against superior competition . The relationship between animals and their ecological niches has been firmly established. A typical species becomes extinct within 10 million years of its first appearance, although some species, called living fossils , survive with little to no morphological change for hundreds of millions of years. Mass extinctions are relatively rare events; however, isolated extinctions of species and clades are quite common, and are
12240-609: 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
12360-410: The relative importance of genetic factors compared to environmental ones as the causes of extinction has been compared to the debate on nature and nurture . The question of whether more extinctions in the fossil record have been caused by evolution or by competition or by predation or by disease or by catastrophe is a subject of discussion; Mark Newman, the author of Modeling Extinction , argues for
12480-661: 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
12600-474: The seas, while rivers were dominated by lobe-finned fish like rhizodonts . During the Permian period (299 to 252 Ma), one of the two major branches of amniotes, the synapsids , flourished, with derived therapsids taking over in the middle of the period . The Great Dying wiped out most of the synapsid diversity, with archosaurs , emerging from the other sauropsid branch, replacing many of them in
12720-459: The serious study into the lives of extinct animals. With the publication of the Origin of Species by Charles Darwin in 1859, the field got a theoretical framework. Much of the subsequent work has been to map the relationship between fossil and extant organisms, as well as their history through time. In modern times, Alfred Romer (1894–1973) wrote what has been termed the definitive textbook on
12840-460: The species is no longer able to survive and becomes extinct. This may occur by direct effects, such as the environment becoming toxic , or indirectly, by limiting a species' ability to compete effectively for diminished resources or against new competitor species. Habitat destruction, particularly the removal of vegetation that stabilizes soil, enhances erosion and diminishes nutrient availability in terrestrial ecosystems. This degradation can lead to
12960-537: The subject was written by Robert L. Carroll of McGill University, the 1988 text Vertebrate Paleontology and Evolution . Carroll was president of the Society of Vertebrate Paleontology in 1983. The Society keeps its members informed on the latest discoveries through newsletters and the Journal of Vertebrate Paleontology . The "traditional" vertebrate classification scheme employ evolutionary taxonomy where several of
13080-510: The subject, called Vertebrate Paleontology . It shows the progression of evolution in fossil fish, and amphibians and reptiles through comparative anatomy, including a list of all the (then) known fossil vertebrate genera . Romer became the first president of the Society of Vertebrate Paleontology in 1940, alongside co-founder Howard Chiu. An updated work that largely carried on the tradition from Romer, and by many considered definitive book on
13200-596: 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
13320-430: The sudden introduction of human beings to environments full of animals that had never seen them before and were therefore completely unadapted to their predation techniques. Coextinction refers to the loss of a species due to the extinction of another; for example, the extinction of parasitic insects following the loss of their hosts. Coextinction can also occur when a species loses its pollinator , or to predators in
13440-421: The taxa listed are paraphyletic , i.e. have given rise to another taxa that have been given the same rank. For instance, birds are generally considered to be the descendants of reptiles (Saurischian dinosaurs to be precise), but in this system both are listed as separate classes. Under phylogenetic nomenclature , such an arrangement is unacceptable, though it offers excellent overview. This classical scheme
13560-412: The tiniest microorganism to God, is linked in a continuous chain. The extinction of a species was impossible under this model, as it would create gaps or missing links in the chain and destroy the natural order. Thomas Jefferson was a firm supporter of the great chain of being and an opponent of extinction, famously denying the extinction of the woolly mammoth on the grounds that nature never allows
13680-567: The total extinction of the dodo and the extirpation of indigenous horses to the British Isles. He similarly argued against mass extinctions , believing that any extinction must be a gradual process. Lyell also showed that Cuvier's original interpretation of the Parisian strata was incorrect. Instead of the catastrophic floods inferred by Cuvier, Lyell demonstrated that patterns of saltwater and freshwater deposits , like those seen in
13800-402: 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,
13920-581: The wider scientific community of his theory. Cuvier was a well-regarded geologist, lauded for his ability to reconstruct the anatomy of an unknown species from a few fragments of bone. His primary evidence for extinction came from mammoth skulls found in the Paris basin . Cuvier recognized them as distinct from any known living species of elephant, and argued that it was highly unlikely such an enormous animal would go undiscovered. In 1812, Cuvier, along with Alexandre Brongniart and Geoffroy Saint-Hilaire , mapped
14040-491: Was difficult to disprove. When parts of the world had not been thoroughly examined and charted, scientists could not rule out that animals found only in the fossil record were not simply "hiding" in unexplored regions of the Earth. Georges Cuvier is credited with establishing the modern conception of extinction in a 1796 lecture to the French Institute , though he would spend most of his career trying to convince
14160-430: Was not until 1982, when David Raup and Jack Sepkoski published their seminal paper on mass extinctions, that Cuvier was vindicated and catastrophic extinction was accepted as an important mechanism . The current understanding of extinction is a synthesis of the cataclysmic extinction events proposed by Cuvier, and the background extinction events proposed by Lyell and Darwin. Extinction is an important research topic in
14280-571: Was skeptical that catastrophic events of a scale large enough to cause total extinction were possible. In his geological history of the earth titled Hydrogeologie, Lamarck instead argued that the surface of the earth was shaped by gradual erosion and deposition by water, and that species changed over time in response to the changing environment. Charles Lyell , a noted geologist and founder of uniformitarianism , believed that past processes should be understood using present day processes. Like Lamarck, Lyell acknowledged that extinction could occur, noting
14400-455: Was ultimately named Megalonyx jeffersonii in his honor. Jefferson corresponded with Cuvier, including sending him a shipment of highly desirable bones of the American mastodon and the woolly mammoth . Paleontology really got started though, with the publication of Recherches sur les poissons fossiles (1833–1843) by Swiss naturalist Louis Agassiz (1807–1873). He studied, described and listed hundreds of species of fossil fish , beginning
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