The Carboniferous is a geologic period and system of the Paleozoic era that spans 60 million years from the end of the Devonian Period 358.9 Ma to the beginning of the Permian Period, 298.9 Ma. It is the fifth and penultimate period of the Paleozoic era and the fifth period of the Phanerozoic eon. In North America, the Carboniferous is often treated as two separate geological periods, the earlier Mississippian and the later Pennsylvanian.
The PaleozoicEra is the first of three geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic and ends 251.9 Ma at the start of the Mesozoic Era. The Paleozoic is subdivided into six geologic periods, Cambrian, Ordovician, Silurian, Devonian, Carboniferous and Permian. Some geological timescales divide the Paleozoic informally into early and late sub-eras: the Early Paleozoic consisting of the Cambrian, Ordovician and Silurian; the Late Paleozoic consisting of the Devonian, Carboniferous and Permian.
Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in their adult form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are members of the largest crinoid order, Comatulida. Crinoids are echinoderms in the phylum Echinodermata, which also includes the starfish, brittle stars, sea urchins and sea cucumbers. They live in both shallow water and in depths of over 9,000 metres (30,000 ft).
Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or female. Cycads vary in size from having trunks only a few centimeters to several meters tall. They typically grow slowly and have long lifespans. Because of their superficial resemblance to palms or ferns, they are sometimes mistaken for them, but they are not closely related to either group. Cycads are gymnosperms (naked-seeded), meaning their unfertilized seeds are open to the air to be directly fertilized by pollination, as contrasted with angiosperms, which have enclosed seeds with more complex fertilization arrangements. Cycads have very specialized pollinators, usually a specific species of beetle. Both male and female cycads bear cones (strobili), somewhat similar to conifer cones.
Meganeura is a genus of extinct insects from the Late Carboniferous. It is a member of the extinct order Meganisoptera, which are closely related to and resemble dragonflies and damselflies. Like other odonoapterans, they were predatory, with their diet mainly consisting of other insects. The genus belongs to the Meganeuridae, a family including other similarly giant dragonfly-like insects ranging from the Late Carboniferous to Middle Permian. With single wing length reaching 32 centimetres (13 in) and a wingspan about 65–75 cm (2.13–2.46 ft), M. monyi is one of the largest-known flying insect species.
Holocephali, sometimes given the name Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. Little is known about these primitive forms, and the only surviving group in the subclass is the order Chimaeriformes.
Endocerida is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. Endocerida was a diverse group of cephalopods that lived from the Early Ordovician possibly to the Late Silurian. Their shells were variable in form. Some were straight (orthoconic), others curved (cyrtoconic); some were long (longiconic), others short (breviconic). Some long-shelled forms like Endoceras attained shell lengths close to 6 metres (20 ft). The related Cameroceras is anecdotally reported to have reached lengths approaching 9 metres (30 ft), but these claims are problematic. The overwhelming majority of endocerids and nautiloids in general are much smaller, usually less than a meter long when fully grown.
Helicoprion is an extinct genus of shark-like eugeneodont fish. Almost all fossil specimens are of spirally arranged clusters of the individuals' teeth, called "tooth whorls", which in life were embedded in the lower jaw. As with most extinct cartilaginous fish, the skeleton is mostly unknown. Fossils of Helicoprion are known from a 20 million year timespan during the Permian period from the Artinskian stage of the Cisuralian to the Roadian stage of the Guadalupian. The closest living relatives of Helicoprion are the chimaeras, though their relationship is very distant. The unusual tooth arrangement is thought to have been an adaption for feeding on soft bodied prey, and may have functioned as a deshelling mechanism for hard bodied cephalopods such as nautiloids and ammonoids. In 2013, systematic revision of Helicoprion via morphometric analysis of the tooth whorls found only H. davisii, H. bessonowi and H. ergassaminon to be valid, with some of the larger tooth whorls being outliers.
In the geologic time scale, the Changhsingian or Changxingian is the latest age or uppermost stage of the Permian. It is also the upper or latest of two subdivisions of the Lopingian Epoch or Series. The Changhsingian lasted from 254.14 to 251.9 Ma ago. It is preceded by the Wuchiapingian age/stage and is followed by the Induan age/stage.
Parahelicoprion is an extinct genus of shark-like cartilaginous fish that lived during the Early Permian. Two species are known ; P. clerci from Arta Beds of the Ural Mountains of Russia, and P. mariosuarezi from the Copacabana Formation of Bolivia. Members of the genus possessed a row of large tooth crowns on the midline of the lower jaw, known as a tooth whorl. The characteristics of this whorl are unique to fishes of the order Eugeneodontida, and more specifically the family Helicoprionidae to which Parahelicoprion belongs. The genus name refers to Helicoprion, another eugeneodont from the Ural Mountains that bore a similar midline tooth arrangement.
The Eugeneodontida, sometimes also called Eugeneodontiformes, is an extinct and poorly known order of cartilaginous fishes. They possessed "tooth-whorls" on the symphysis of either the lower or both jaws and pectoral fins supported by long radials. They probably lacked pelvic fins and anal fins. The palatoquadrate was either fused to the skull or reduced. Now determined to be within the Holocephali, their closest living relatives are chimaeras. The eugeneodonts are named after paleontologist Eugene S. Richardson, Jr. The group first appeared in the fossil record during the late Mississippian (Serpukhovian). The youngest eugeneodonts are known from the Early Triassic. The geologically youngest fossils of the group are known from the Sulphur Mountain Formation, Vardebukta Formation and Wordie Creek Formation (Greenland).
Helicoprionidae is an extinct family of holocephalans within the order Eugeneodontida. Members of the Helicoprionidae possessed a "whorl" of tooth crowns connected by a single root along the midline of the lower jaw. While historically considered elasmobranchs related sharks and rays, the closest living relatives of the Helicoprionidae and all other eugeneodonts are now thought to be the ratfishes. The anatomy of the tooth-whorls vary between taxa, with some possessing highly specialized, coiling spirals, while others such as Sarcoprion and Parahelicoprion possessed shorter whorls.
Trihecaton is an extinct genus of microsaur from the Late Pennsylvanian of Colorado. Known from a single species, Trihecaton howardinus, this genus is distinctive compared to other microsaurs due to possessing a number of plesiomorphic ("primitive") features relative to the rest of the group. These include large intercentra, folded enamel, and a large coronoid process of the jaw. Its classification is controversial due to combining a long body with strong limbs, features which typically are not present at the same time in other microsaurs. Due to its distinctiveness, Trihecaton has been given its own monospecific family, Trihecatontidae.
Cambrian Stage 4 is the still unnamed fourth stage of the Cambrian and the upper stage of Cambrian Series 2. It follows Cambrian Stage 3 and lies below the Wuliuan. The lower boundary has not been formally defined by the International Commission on Stratigraphy. One proposal is the first appearance of two trilobite genera, Olenellus or Redlichia. Another proposal is the first appearance of the trilobite species Arthricocephalus chauveaui. Both proposals will set the lower boundary close to 514 million years ago. The upper boundary corresponds to the beginning of the Wuliuan.
Toxoprion is an extinct genus of eugeneodont holocephalans whose fossils are found in marine strata from the Early Carboniferous until the Late Permian near Eureka, Nevada.
Ornithoprion is a genus of extinct cartilaginous fish in the family Caseodontidae. The only species, O. hertwigi, lived during the Moscovian stage of the Pennsylvanian, between 315.2 to 307 million years ago, and is known from black shale deposits in what is now the Midwestern United States. The discovery and description of Ornithoprion, performed primarily via radiography, helped clarify the skull anatomy of eugeneodonts; a group which includes O. hertwigi and which were previously known primarily from isolated teeth. The genus name, which is derived from the ancient Greek órnith- meaning 'bird' and príōn meaning 'saw', was inspired by the animal's vaguely bird-like skull and the saw-like appearance of the lower teeth. The species name honors Oscar Hertwig.
Ceratodontiformes is the only extant order of lungfish, containing the families Neoceratodontidae, Lepidosirenidae, and Protopteridae as well as many other extinct groups. Members of this group are the only lungfish known to have survived the Permian-Triassic extinction event. Although lungfish originated in marine environments, the Ceratodontiformes have been an exclusively freshwater group since the Carboniferous. This order was formerly considered the suborder Ceratodontoidei.
Romerodus is an extinct genus of cartilaginous fish in the family Caseodontidae. While it and the rest of its family were historically considered elasmobranchs related to sharks and rays, they are now regarded as holocephalans, a diverse subclass which is today only represented by chimaeras. Romerodus is known from the Carboniferous and possibly Permian periods of North America, and the only named species, R. orodontus, was discovered in organic shale deposits in the U.S. state of Nebraska. It is one of few members of its order, the Eugeneodontida, that is known from multiple complete, well preserved body fossils, and is thus an important taxon for understanding the anatomy and ecology of less well preserved eugeneodonts such as Helicoprion. The genus name honors paleontologist Alfred Romer.
The Caseodontidae is an extinct family of eugeneodont holocephalans known from the late Paleozoic to earliest Mesozoic of Greenland, Canada and the United States. Members of the group are characterized by a reduced or absent palatoquadrate, elongate upper and mandibular rostra, and bulbous, crushing dentition, including a small symphyseal whorl of teeth on the lower jaw and batteries of teeth fused directly to the neurocranium. Several genera are known from partial or complete body fossils.
Paredestus is a monotypic genus of extinct eugeneodont holocephalan from the Early Triassic of Canada. The type and only species, P. bricircum represents the last known member of the superfamily Edestoidea and among the last known eugeneodonts. It was named in 2008 based on tooth and jaw material, with the holotype representing the only known specimen.
