Vertebrate paleontology is the subfield of

paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...

that seeks to discover, through the study of

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

ized remains, the behavior, reproduction and appearance of extinct animals with

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...

e or a

notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consi ...

. It also tries to connect, by using the

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

ary

timeline A timeline is a display of a list of events in chronological order. It is typically a graphic design showing a long bar labelled with dates paralleling it, and usually contemporaneous events. Timelines can use any suitable scale represen ...

, the animals of the past and their modern-day relatives. The fossil record shows aspects of the meandering evolutionary path from early aquatic

vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...

s to

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...

s, with a host of transitional fossils, though there are still large blank areas. The earliest known fossil vertebrates were heavily armored

fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...

discovered in rocks from the

Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...

Period about 500 to 430 Ma (

megaannum A year or annus is the orbital period of a planetary body, for example, the Earth, moving in its orbit around the Sun. Due to the Earth's axial tilt, the course of a year sees the passing of the seasons, marked by change in weather, the hour ...

, million years ago). The

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...

Period (395 to 345 Ma) brought in the changes that allowed primitive air-breathing fish to remain on land as long as they wished, thus becoming the first terrestrial vertebrates, the

amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...

s. Amphibians developed forms of reproduction and locomotion and a

metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run ...

better suited for life exclusively on land, becoming more reptilian. Full-fledged reptiles appeared in the

Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...

Period (345 to 280 Ma). The reptilian changes and adaptations to diet and geography are chronicled in the fossil record of the varying forms of therapsida. True mammals showed up in the

Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...

Period (225 to 190 Ma) around the same time as the

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...

s, which also sprouted from the reptilian line.

Bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

s first diverged from dinosaurs between 100 Ma and 60 Ma. One specimen of ''

Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...

'' from around 150 Ma subjected to x-ray microtomography revealed significant bird-like alterations.

History

One of the people who helped figure out the vertebrate progression was French zoologist

Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...

(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

, his work proved the (at the time) contested theory of

extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...

of species.

Thomas Jefferson Thomas Jefferson (April 13, 1743 – July 4, 1826) was an American statesman, diplomat, lawyer, architect, philosopher, and Founding Fathers of the United States, Founding Father who served as the third president of the United States from 18 ...

is credited with initiating the science of vertebrate paleontology in the United States with the reading of a paper to the

American Philosophical Society The American Philosophical Society (APS), founded in 1743 in Philadelphia, is a scholarly organization that promotes knowledge in the sciences and humanities through research, professional meetings, publications, library resources, and communit ...

Philadelphia Philadelphia, often called Philly, is the largest city in the Commonwealth of Pennsylvania, the sixth-largest city in the U.S., the second-largest city in both the Northeast megalopolis and Mid-Atlantic regions after New York City. Since ...

in 1797. Jefferson presented fossil bones of a ground sloth found in a cave in western Virginia and named the genus (''

Megalonyx ''Megalonyx'' ( Greek, "large claw") is an extinct genus of ground sloths of the family Megalonychidae, native to North America during the Pliocene and Pleistocene epochs. It became extinct during the Quaternary extinction event at the end ...

''). The species 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 The woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus s ...

. Paleontology really got started though, with the publication of '' Recherches sur les poissons fossiles'' (1833–1843) by Swiss naturalist

Louis Agassiz Jean Louis Rodolphe Agassiz ( ; ) FRS (For) FRSE (May 28, 1807 – December 14, 1873) was a Swiss-born American biologist and geologist who is recognized as a scholar of Earth's natural history. Spending his early life in Switzerland, he rec ...

(1807–1873). He studied, described and listed hundreds of species of fossil fish, beginning the serious study into the lives of extinct animals. With the publication of the Origin of Species by

Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...

in 1859, the field got a theoretical framework. Much of the subsequent work has been to map the relationship between fossil and

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

organisms, as well as their history through time. In modern times, Alfred Romer (1894–1973) wrote what has been termed the definitive textbook on 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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

. Romer became the first president of the

Society of Vertebrate Paleontology The Society of Vertebrate Paleontology (SVP) is a professional organization that was founded in the United States in 1940 to advance the science of vertebrate paleontology around the world. Mission and Activities SVP has about 2,300 members inte ...

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 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''.

Classification

The "traditional" vertebrate classification scheme employ

evolutionary taxonomy Evolutionary taxonomy, evolutionary systematics or Darwinian classification is a branch of biological classification that seeks to classify organisms using a combination of phylogenetic relationship (shared descent), progenitor-descendant relat ...

where several of the taxa listed are

paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...

, i.e. have given rise to another taxa that have been given the same rank. For instance,

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

s are generally considered to be the descendants of

reptiles Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchoceph ...

(Saurischian

s to be precise), but in this system both are listed as separate classes. Under

phylogenetic nomenclature Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...

, such an arrangement is unacceptable, though it offers excellent overview. This classical scheme 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).Fortuny, J., Bolet, A., Sellés, A.G., J. Cartanyà, J. & À. Galobart, À. (2011)
New insights on the Permian and Triassic vertebrates from the Iberian Peninsula with emphasis on the Pyrenean and Catalonian basins
. ''

Journal of Iberian Geology ''Journal of Iberian Geology'' (formerly ''Cuadernos de Geología Ibérica'') is a triannual peer-reviewed scientific journal published by the Universidad Complutense de Madrid. The journal covers the field of geology and related earth sciences ...

'' no 37 (1): pp 65-86 Kingdom

Animalia Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...

* Phylum Chordata (vertebrates) ** Class Agnatha (jawless fish) *** Subclass

Cyclostomata Cyclostomi, often referred to as Cyclostomata , is a group of vertebrates that comprises the living jawless fishes: the lampreys and hagfishes. Both groups have jawless mouths with horny epidermal structures that function as teeth called cer ...

(

hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, ...

and

lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...

s) *** Subclass Ostracodermi (armoured jawless fish) † ** Class

Chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...

(cartilaginous fish) *** Subclass Elasmobranchii (

shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...

s and rayes) *** Subclass Holocephali (

chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...

s and extinct relatives) ** Class

Placodermi Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally ' plate-skinned') is a class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devonian period. Their head and thorax were co ...

(armoured fish) † ** Class

Acanthodii Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, ...

("spiny sharks", sometimes classified under bony fishes) † ** Class

Osteichthyes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...

(bony fish) *** Subclass

Actinopterygii Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...

*** Subclass Sarcopterygii ** Class

Amphibia Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...

*** Subclass

Labyrinthodontia "Labyrinthodontia" ( Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally con ...

† *** Subclass Lepospondyli † *** Subclass

Lissamphibia The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relativ ...

** Class Reptilia *** Subclass

Anapsida An anapsid is an amniote whose skull lacks one or more skull openings (fenestra, or fossae) near the temples. Traditionally, the Anapsida are the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Diapsida evolved ...

**** Order Cotylosauria † **** Order

Testudines Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...

*** Subclass Synapsida **** Order

Pelycosauria Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is no ...

† **** Order Therapsida † *** Subclass Euryapsida **** Order Sauropterygia † **** Order Ichthyosauria † *** Subclass Diapsida (lizards & snakes too) **** Order

Crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...

(crocodiles, alligators etc.) **** Order Sphenodontia (Tuatara and relatives) **** Order

Squamata Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest order of reptiles, comprising lizards, snakes, and amphisbaenians (worm lizards), which are collectively known as squamates or scaled reptiles. With over 10,900 species, ...

(Lizards and snakes) **** Order Thecodonts † **** Order Pterosauria † **** Order Saurischia (dinosaurs) † **** Order

Ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...

(dinosaurs) † ** Class

Aves Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

*** Subclass

Archaeornithes The Archaeornithes, classically Archæornithes, is an extinct group of the first primitive, reptile-like birds. It is an evolutionary grade of transitional fossils, the primitive birds halfway between non avian dinosaur ancestors and the derived m ...

(primitive dinosaur-like birds like ''

'') † *** Subclass Neornithes (modern birds and some advanced Cretaceous forms) **** Superorder

Odontognathae 230px, Ichthyornis skeletons Odontognathae is a disused name for a paraphyletic group of toothed prehistoric birds. The group was originally proposed by Alexander Wetmore, who attempted to link fossil birds with the presence of teeth, specifical ...

(Cretaceous toothed birds) † **** Superorder

Palaeognathae Palaeognathae (; ) is a infraclass of birds, called paleognaths, within the class Aves of the clade Archosauria. It is one of the two extant infraclasses of birds, the other being Neognathae, both of which form Neornithes. Palaeognathae conta ...

( ratites) **** Superorder

Neognathae Neognathae (; ) is a infraclass of birds, called neognaths, within the class Aves of the clade Archosauria. Neognathae includes the majority of living birds; the exceptions being the tinamous and the flightless ratites, which belong instead to ...

(All other extant birds) ** Class

Mammalia Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...

*** Subclass Prototheria **** Order Monotremata (

platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or mono ...

and the

echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...

s) *** Subclass

Theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...

**** Infraclass

Metatheria Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as w ...

***** Order

Marsupialia Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...

(kangaroos, dunnarts, opossums, wombats etc.) **** Infraclass Eutheria (placentals) ***** Order Insectivora ***** Order Chiroptera (bats) ***** Order Creodonta ***** Order

Carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...

(dogs/cats) ***** Order Perissodactyla (horses) ***** Order Artiodactyla (cattle and other ungulates) ***** Order

Proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family ( Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Fr ...

(elephants) ***** Order Edentata ***** Order

Cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...

(whales and dolphins) ***** Order

Rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...

ia (mice, rats etc.) ***** Order Lagomorpha (rabbits) ***** Order

Primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter including ...

(monkeys, apes and primates)

History

Classification

See also

References

Further reading