The Cambrian Period was the first geological period of the Paleozoic Era, of the Phanerozoic Eon. The Cambrian lasted 55.6 million years from the end of the preceding Ediacaran Period 541 million years ago to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, its base, are somewhat in flux; the period was established by Adam Sedgwick, who named it after Cambria, the Latin name of Wales, where Britain's Cambrian rocks are best exposed. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells; as a result, our understanding of the Cambrian biology surpasses that of some periods. The Cambrian marked a profound change in life on Earth. Complex, multicellular organisms became more common in the millions of years preceding the Cambrian, but it was not until this period that mineralized—hence fossilized—organisms became common; the rapid diversification of life forms in the Cambrian, known as the Cambrian explosion, produced the first representatives of all modern animal phyla.
Phylogenetic analysis has supported the view that during the Cambrian radiation, metazoa evolved monophyletically from a single common ancestor: flagellated colonial protists similar to modern choanoflagellates. Although diverse life forms prospered in the oceans, the land is thought to have been comparatively barren—with nothing more complex than a microbial soil crust and a few molluscs that emerged to browse on the microbial biofilm. Most of the continents were dry and rocky due to a lack of vegetation. Shallow seas flanked the margins of several continents created during the breakup of the supercontinent Pannotia; the seas were warm, polar ice was absent for much of the period. Despite the long recognition of its distinction from younger Ordovician rocks and older Precambrian rocks, it was not until 1994 that the Cambrian system/period was internationally ratified; the base of the Cambrian lies atop a complex assemblage of trace fossils known as the Treptichnus pedum assemblage. The use of Treptichnus pedum, a reference ichnofossil to mark the lower boundary of the Cambrian, is difficult since the occurrence of similar trace fossils belonging to the Treptichnids group are found well below the T. pedum in Namibia and Newfoundland, in the western USA.
The stratigraphic range of T. pedum overlaps the range of the Ediacaran fossils in Namibia, in Spain. The Cambrian Period was followed by the Ordovician Period; the Cambrian is divided into ten ages. Only three series and six stages are named and have a GSSP; because the international stratigraphic subdivision is not yet complete, many local subdivisions are still used. In some of these subdivisions the Cambrian is divided into three series with locally differing names – the Early Cambrian, Middle Cambrian and Furongian. Rocks of these epochs are referred to as belonging to Upper Cambrian. Trilobite zones allow biostratigraphic correlation in the Cambrian; each of the local series is divided into several stages. The Cambrian is divided into several regional faunal stages of which the Russian-Kazakhian system is most used in international parlance: *Most Russian paleontologists define the lower boundary of the Cambrian at the base of the Tommotian Stage, characterized by diversification and global distribution of organisms with mineral skeletons and the appearance of the first Archaeocyath bioherms.
The International Commission on Stratigraphy list the Cambrian period as beginning at 541 million years ago and ending at 485.4 million years ago. The lower boundary of the Cambrian was held to represent the first appearance of complex life, represented by trilobites; the recognition of small shelly fossils before the first trilobites, Ediacara biota earlier, led to calls for a more defined base to the Cambrian period. After decades of careful consideration, a continuous sedimentary sequence at Fortune Head, Newfoundland was settled upon as a formal base of the Cambrian period, to be correlated worldwide by the earliest appearance of Treptichnus pedum. Discovery of this fossil a few metres below the GSSP led to the refinement of this statement, it is the T. pedum ichnofossil assemblage, now formally used to correlate the base of the Cambrian. This formal designation allowed radiometric dates to be obtained from samples across the globe that corresponded to the base of the Cambrian. Early dates of 570 million years ago gained favour, though the methods used to obtain this number are now considered to be unsuitable and inaccurate.
A more precise date using modern radiometric dating yield a date of 541 ± 0.3 million years ago. The ash horizon in Oman from which this date was recovered corresponds to a marked fall in the abundance of carbon-13 that correlates to equivalent excursions elsewhere in the world, to the disappearance of distinctive Ediacaran fossils. There are arguments that the dated horizon in Oman does not correspond to the Ediacaran-Cambrian boundary, but represents a facies change from marine to evaporite-dominated strata — which w
Fossilworks is a portal which provides query and analysis tools to facilitate access to the Paleobiology Database, a large relational database assembled by hundreds of paleontologists from around the world. Fossilworks is housed at Macquarie University, it includes many analysis and data visualization tools included in the Paleobiology Database. "Fossilworks". Retrieved 2010-04-08
The Ordovician is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.2 million years from the end of the Cambrian Period 485.4 million years ago to the start of the Silurian Period 443.8 Mya. The Ordovician, named after the Celtic tribe of the Ordovices, was defined by Charles Lapworth in 1879 to resolve a dispute between followers of Adam Sedgwick and Roderick Murchison, who were placing the same rock beds in northern Wales into the Cambrian and Silurian systems, respectively. Lapworth recognized that the fossil fauna in the disputed strata were different from those of either the Cambrian or the Silurian systems, placed them in a system of their own; the Ordovician received international approval in 1960, when it was adopted as an official period of the Paleozoic Era by the International Geological Congress. Life continued to flourish during the Ordovician as it did in the earlier Cambrian period, although the end of the period was marked by the Ordovician–Silurian extinction events.
Invertebrates, namely molluscs and arthropods, dominated the oceans. The Great Ordovician Biodiversification Event increased the diversity of life. Fish, the world's first true vertebrates, continued to evolve, those with jaws may have first appeared late in the period. Life had yet to diversify on land. About 100 times as many meteorites struck the Earth per year during the Ordovician compared with today; the Ordovician Period began with a major extinction called the Cambrian–Ordovician extinction event, about 485.4 Mya. It lasted for about 42 million years and ended with the Ordovician–Silurian extinction events, about 443.8 Mya which wiped out 60% of marine genera. The dates given are recent radiometric dates and vary from those found in other sources; this second period of the Paleozoic era created abundant fossils that became major petroleum and gas reservoirs. The boundary chosen for the beginning of both the Ordovician Period and the Tremadocian stage is significant, it correlates well with the occurrence of widespread graptolite and trilobite species.
The base of the Tremadocian allows scientists to relate these species not only to each other, but to species that occur with them in other areas. This makes it easier to place many more species in time relative to the beginning of the Ordovician Period. A number of regional terms have been used to subdivide the Ordovician Period. In 2008, the ICS erected a formal international system of subdivisions. There exist Baltoscandic, Siberian, North American, Chinese Mediterranean and North-Gondwanan regional stratigraphic schemes; the Ordovician Period in Britain was traditionally broken into Early and Late epochs. The corresponding rocks of the Ordovician System are referred to as coming from the Lower, Middle, or Upper part of the column; the faunal stages from youngest to oldest are: Late Ordovician Hirnantian/Gamach Rawtheyan/Richmond Cautleyan/Richmond Pusgillian/Maysville/Richmond Middle Ordovician Trenton Onnian/Maysville/Eden Actonian/Eden Marshbrookian/Sherman Longvillian/Sherman Soudleyan/Kirkfield Harnagian/Rockland Costonian/Black River Chazy Llandeilo Whiterock Llanvirn Early Ordovician Cassinian Arenig/Jefferson/Castleman Tremadoc/Deming/Gaconadian The Tremadoc corresponds to the Tremadocian.
The Floian corresponds to the lower Arenig. The Llanvirn occupies the rest of the Darriwilian, terminates with it at the base of the Late Ordovician; the Sandbian represents the first half of the Caradoc. During the Ordovician, the southern continents were collected into Gondwana. Gondwana started the period in equatorial latitudes and, as the period progressed, drifted toward the South Pole. Early in the Ordovician, the continents of Laurentia and Baltica were still independent continents, but Baltica began to move towards Laurentia in the period, causing the Iapetus Ocean between them to shrink; the small continent Avalonia separated from Gondwana and began to move north towards Baltica and Laurentia, opening the Rheic Ocean between Gondwana and Avalonia. The Taconic orogeny, a major mountain-building episode, was well under way in Cambrian times. In the early and middle Ordovician, temperatures were mild, but at the beginning of the Late Ordovician, from 460 to 450 Ma, volcanoes along the margin of the Iapetus Ocean spewed massive amounts of carbon dioxide, a greenhouse gas, into the atmosphere, turning the planet into a hothouse.
Sea levels were high, but as Gondwana moved south, ice accumulated into glaciers and sea levels dropped. At first, low-lying sea beds increased diversity, but glaciation led to mass extinctions as the seas drained and continental shelves became dry land. During the Ordovician, in fact during the Tremadocian, marine transgressions worldwide were the greatest for which evidence is preserved; these volcanic island arcs collided with proto North America to form the Appalachian mountains. By the end of the Late Ordovician the volcanic emissions had stopped. Gondwana had by that time neared the South Pole and was glaciated
In the vertebrate spinal column, each vertebra is an irregular bone with a complex structure composed of bone and some hyaline cartilage, the proportions of which vary according to the segment of the backbone and the species of vertebrate. The basic configuration of a vertebra varies; the upper and lower surfaces of the vertebra body give attachment to the intervertebral discs. The posterior part of a vertebra forms a vertebral arch, in eleven parts, consisting of two pedicles, two laminae, seven processes; the laminae give attachment to the ligamenta flava. There are vertebral notches formed from the shape of the pedicles, which form the intervertebral foramina when the vertebrae articulate; these foramina are the exit conducts for the spinal nerves. The body of the vertebra and the vertebral arch form the vertebral foramen, the larger, central opening that accommodates the spinal canal, which encloses and protects the spinal cord. Vertebrae articulate with each other to give strength and flexibility to the spinal column, the shape at their back and front aspects determines the range of movement.
Structurally, vertebrae are alike across the vertebrate species, with the greatest difference seen between an aquatic animal and other vertebrate animals. As such, vertebrates take their name from the vertebrae; each vertebra is an irregular bone. The size of the vertebrae varies according to placement in the vertebral column, spinal loading and pathology. Along the length of the spine the vertebrae change to accommodate different needs related to stress and mobility; every vertebra has a body, which consists of a large anterior middle portion called the centrum and a posterior vertebral arch called a neural arch. The body is composed of cancellous bone, the spongy type of osseous tissue, whose micro-anatomy has been studied within the pedicle bones; this cancellous bone is in turn, covered by a thin coating of cortical bone, the hard and dense type of osseous tissue. The vertebral arch and processes have thicker coverings of cortical bone; the upper and lower surfaces of the body of the vertebra are flattened and rough in order to give attachment to the intervertebral discs.
These surfaces are the vertebral endplates which are in direct contact with the intervertebral discs and form the joint. The endplates are formed from a thickened layer of the cancellous bone of the vertebral body, the top layer being more dense; the endplates function to contain the adjacent discs, to evenly spread the applied loads, to provide anchorage for the collagen fibers of the disc. They act as a semi-permeable interface for the exchange of water and solutes; the vertebral arch is formed by pedicles and laminae. Two pedicles extend from the sides of the vertebral body to join the body to the arch; the pedicles are short thick processes that extend, one from each side, from the junctions of the posteriolateral surfaces of the centrum, on its upper surface. From each pedicle a broad plate, a lamina, projects backwards and medialwards to join and complete the vertebral arch and form the posterior border of the vertebral foramen, which completes the triangle of the vertebral foramen; the upper surfaces of the laminae are rough to give attachment to the ligamenta flava.
These ligaments connect the laminae of adjacent vertebra along the length of the spine from the level of the second cervical vertebra. Above and below the pedicles are shallow depressions called vertebral notches; when the vertebrae articulate the notches align with those on adjacent vertebrae and these form the openings of the intervertebral foramina. The foramina allow the entry and exit of the spinal nerves from each vertebra, together with associated blood vessels; the articulating vertebrae provide a strong pillar of support for the body. There are seven processes projecting from the vertebra. A major part of a vertebra is a backward extending spinous process; this process points caudally from the junction of the laminae. The spinous process serves to attach ligaments; the two transverse processes, one on each side of the vertebral body, project from either side at the point where the lamina joins the pedicle, between the superior and inferior articular processes. They serve for the attachment of muscles and ligaments, in particular the intertransverse ligaments.
There is a facet on each of the transverse processes of thoracic vertebrae which articulates with the tubercle of the rib. A facet on each side of the thoracic vertebral body articulates with the head of the rib. There are superior and inferior articular facet joints on each side of the vertebra, which serve to restrict the range of movement possible; these facets are joined by a thin portion of the vertebral arch called the pars interarticularis. The transverse process of a lumbar vertebra is sometimes called the costal or costiform process because it corresponds to a rudimentary rib which, as opposed to the thorax, is not developed in the lumbar region. Vertebrae take their names from the regions of the vertebral column. There are thirty-three vertebrae in the human vertebral column—seven cervical vertebrae, twelve thoracic vertebrae, five lumbar vertebrae, five fused sacral vertebrae forming the sacrum and three to five coccygeal vertebrae, forming the coccyx; the regional vertebrae increase in size as they become smaller in the coccyx.
There are seven cervical vertebrae
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, although the exact origin and timing of the evolution of dinosaurs is the subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201 million years ago. Reverse genetic engineering and the fossil record both demonstrate that birds are modern feathered dinosaurs, having evolved from earlier theropods during the late Jurassic Period; as such, birds were the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event 66 million years ago. Dinosaurs can therefore be divided into birds; this article deals with non-avian dinosaurs. Dinosaurs are a varied group of animals from taxonomic and ecological standpoints. Birds, at over 10,000 living species, are the most diverse group of vertebrates besides perciform fish. Using fossil evidence, paleontologists have identified over 500 distinct genera and more than 1,000 different species of non-avian dinosaurs.
Dinosaurs are represented on every continent by fossil remains. Through the first half of the 20th century, before birds were recognized to be dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, has indicated that all dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction; some were herbivorous, others carnivorous. Evidence suggests that egg-laying and nest-building are additional traits shared by all dinosaurs and non-avian alike. While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, some extinct groups developed skeletal modifications such as bony armor and spines. While the dinosaurs' modern-day surviving avian lineage are small due to the constraints of flight, many prehistoric dinosaurs were large-bodied—the largest sauropod dinosaurs are estimated to have reached lengths of 39.7 meters and heights of 18 meters and were the largest land animals of all time.
Still, the idea that non-avian dinosaurs were uniformly gigantic is a misconception based in part on preservation bias, as large, sturdy bones are more to last until they are fossilized. Many dinosaurs were quite small: Xixianykus, for example, was only about 50 cm long. Since the first dinosaur fossils were recognized in the early 19th century, mounted fossil dinosaur skeletons have been major attractions at museums around the world, dinosaurs have become an enduring part of world culture; the large sizes of some dinosaur groups, as well as their monstrous and fantastic nature, have ensured dinosaurs' regular appearance in best-selling books and films, such as Jurassic Park. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, new discoveries are covered by the media; the taxon'Dinosauria' was formally named in 1841 by paleontologist Sir Richard Owen, who used it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were being recognized in England and around the world.
The term is derived from Ancient Greek δεινός, meaning'terrible, potent or fearfully great', σαῦρος, meaning'lizard or reptile'. Though the taxonomic name has been interpreted as a reference to dinosaurs' teeth and other fearsome characteristics, Owen intended it to evoke their size and majesty. Other prehistoric animals, including pterosaurs, ichthyosaurs and Dimetrodon, while popularly conceived of as dinosaurs, are not taxonomically classified as dinosaurs. Pterosaurs are distantly related to dinosaurs; the other groups mentioned are, like dinosaurs and pterosaurs, members of Sauropsida, except Dimetrodon. Under phylogenetic nomenclature, dinosaurs are defined as the group consisting of the most recent common ancestor of Triceratops and Neornithes, all its descendants, it has been suggested that Dinosauria be defined with respect to the MRCA of Megalosaurus and Iguanodon, because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria. Both definitions result in the same set of animals being defined as dinosaurs: "Dinosauria = Ornithischia + Saurischia", encompassing ankylosaurians, ceratopsians, ornithopods and sauropodomorphs.
Birds are now recognized as being the sole surviving lineage of theropod dinosaurs. In traditional taxonomy, birds were considered a separate class that had evolved from dinosaurs, a distinct superorder. However, a majority of contemporary paleontologists concerned with dinosaurs reject the traditional style of classification in favor of phylogenetic taxonomy. Birds are thus considered to be dinosaurs and dinosaurs are, not extinct. Birds are classified as belonging to the subgroup M
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype. Under the International Code of Zoological Nomenclature, a holotype is one of several kinds of name-bearing types. In the International Code of Nomenclature for algae and plants and ICZN the definitions of types are similar in intent but not identical in terminology or underlying concept. For example, the holotype for the butterfly Lycaeides idas longinus is a preserved specimen of that species, held by the Museum of Comparative Zoology at Harvard University. An isotype is a duplicate of the holotype and is made for plants, where holotype and isotypes are pieces from the same individual plant or samples from the same gathering. A holotype is not "typical" of that taxon, although ideally it should be. Sometimes just a fragment of an organism is the holotype in the case of a fossil.
For example, the holotype of Pelorosaurus humerocristatus, a large herbivorous dinosaur from the early Jurassic period, is a fossil leg bone stored at the Natural History Museum in London. If a better specimen is subsequently found, the holotype is not superseded. Under the ICN, an additional and clarifying type could be designated an epitype under Article 9.8, where the original material is demonstrably ambiguous or insufficient. A conserved type is sometimes used to correct a problem with a name, misapplied. In the absence of a holotype, another type may be selected, out of a range of different kinds of type, depending on the case, a lectotype or a neotype. For example, in both the ICN and the ICZN a neotype is a type, appointed in the absence of the original holotype. Additionally, under the ICZN the Commission is empowered to replace a holotype with a neotype, when the holotype turns out to lack important diagnostic features needed to distinguish the species from its close relatives. For example, the crocodile-like archosaurian reptile Parasuchus hislopi Lydekker, 1885 was described based on a premaxillary rostrum, but this is no longer sufficient to distinguish Parasuchus from its close relatives.
This made. Texan paleontologist Sankar Chatterjee proposed that a new type specimen, a complete skeleton, be designated; the International Commission on Zoological Nomenclature considered the case and agreed to replace the original type specimen with the proposed neotype. The procedures for the designation of a new type specimen when the original is lost come into play for some recent, high-profile species descriptions in which the specimen designated as the holotype was a living individual, allowed to remain in the wild. In such a case, there is no actual type specimen available for study, the possibility exists that—should there be any perceived ambiguity in the identity of the species—subsequent authors can invoke various clauses in the ICZN Code that allow for the designation of a neotype. Article 75.3.7 of the ICZN requires that the designation of a neotype must be accompanied by "a statement that the neotype is, or upon publication has become, the property of a recognized scientific or educational institution, cited by name, that maintains a research collection, with proper facilities for preserving name-bearing types, that makes them accessible for study", but there is no such requirement for a holotype.
Type Allotype Paratype Type species Genetypes- genetic sequence data from type specimens. BOA Photographs of type specimens of Neotropical Rhopalocera
The Devonian is a geologic period and system of the Paleozoic, spanning 60 million years from the end of the Silurian, 419.2 million years ago, to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, where rocks from this period were first studied; the first significant adaptive radiation of life on dry land occurred during the Devonian. Free-sporing vascular plants began to spread across dry land, forming extensive forests which covered the continents. By the middle of the Devonian, several groups of plants had evolved leaves and true roots, by the end of the period the first seed-bearing plants appeared. Various terrestrial arthropods became well-established. Fish reached substantial diversity during this time, leading the Devonian to be dubbed the "Age of Fishes." The first ray-finned and lobe-finned bony fish appeared, while the placoderms began dominating every known aquatic environment. The ancestors of all four-limbed vertebrates began adapting to walking on land, as their strong pectoral and pelvic fins evolved into legs.
In the oceans, primitive sharks became more numerous than in the Late Ordovician. The first ammonites, species of molluscs, appeared. Trilobites, the mollusc-like brachiopods and the great coral reefs, were still common; the Late Devonian extinction which started about 375 million years ago affected marine life, killing off all placodermi, all trilobites, save for a few species of the order Proetida. The palaeogeography was dominated by the supercontinent of Gondwana to the south, the continent of Siberia to the north, the early formation of the small continent of Euramerica in between; the period is named after Devon, a county in southwestern England, where a controversial argument in the 1830s over the age and structure of the rocks found distributed throughout the county was resolved by the definition of the Devonian period in the geological timescale. The Great Devonian Controversy was a long period of vigorous argument and counter-argument between the main protagonists of Roderick Murchison with Adam Sedgwick against Henry De la Beche supported by George Bellas Greenough.
Murchison and Sedgwick named the period they proposed as the Devonian System. While the rock beds that define the start and end of the Devonian period are well identified, the exact dates are uncertain. According to the International Commission on Stratigraphy, the Devonian extends from the end of the Silurian 419.2 Mya, to the beginning of the Carboniferous 358.9 Mya. In nineteenth-century texts the Devonian has been called the "Old Red Age", after the red and brown terrestrial deposits known in the United Kingdom as the Old Red Sandstone in which early fossil discoveries were found. Another common term is "Age of the Fishes", referring to the evolution of several major groups of fish that took place during the period. Older literature on the Anglo-Welsh basin divides it into the Downtonian, Dittonian and Farlovian stages, the latter three of which are placed in the Devonian; the Devonian has erroneously been characterised as a "greenhouse age", due to sampling bias: most of the early Devonian-age discoveries came from the strata of western Europe and eastern North America, which at the time straddled the Equator as part of the supercontinent of Euramerica where fossil signatures of widespread reefs indicate tropical climates that were warm and moderately humid but in fact the climate in the Devonian differed during its epochs and between geographic regions.
For example, during the Early Devonian, arid conditions were prevalent through much of the world including Siberia, North America, China, but Africa and South America had a warm temperate climate. In the Late Devonian, by contrast, arid conditions were less prevalent across the world and temperate climates were more common; the Devonian Period is formally broken into Early and Late subdivisions. The rocks corresponding to those epochs are referred to as belonging to the Lower and Upper parts of the Devonian System. Early DevonianThe Early Devonian lasted from 419.2 ± 2.8 to 393.3 ± 2.5 and began with the Lochkovian stage, which lasted until the Pragian. It spanned from 410.8 ± 2.8 to 407.6 ± 2.5, was followed by the Emsian, which lasted until the Middle Devonian began, 393.3± 2.7 million years ago. During this time, the first ammonoids appeared. Ammonoids during this time period differed little from their nautiloid counterparts; these ammonoids belong to the order Agoniatitida, which in epochs evolved to new ammonoid orders, for example Goniatitida and Clymeniida.
This class of cephalopod molluscs would dominate the marine fauna until the beginning of the Mesozoic era. Middle DevonianThe Middle Devonian comprised two subdivisions: first the Eifelian, which gave way to the Givetian 387.7± 2.7 million years ago. During this time the jawless agnathan fishes began to decline in diversity in freshwater and marine environments due to drastic environmental changes and due to the increasing competition and diversity of jawed fishes; the shallow, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop such essential characteristics as well developed lungs, the ability to crawl out of the water and onto the land for short periods of time. Late DevonianFinally, the Late Devonian started with the Frasnian, 382.7 ± 2.8 to 372.2 ± 2.5, during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing Famennian subdivisi