In phylogenetics, basal is the direction of the base of a rooted phylogenetic tree or cladogram. The term may be more applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to the root; each node in the tree corresponds to a clade. The terms deep-branching or early-branching are similar in meaning. While there must always be two or more basal clades sprouting from the root of every cladogram, those clades may differ in taxonomic rank and/or species diversity. If C is a basal clade within D that has the lowest rank of all basal clades within D, C may be described as the basal taxon of that rank within D. Greater diversification may be associated with more evolutionary innovation, but ancestral characters should not be imputed to the members of a less species-rich basal clade without additional evidence, as there can be no assurance such an assumption is valid. In general, clade A is more basal than clade B if B is a subgroup of the sister group of A.
Within large groups, "basal" may be used loosely to mean'closer to the root than the great majority of', in this context terminology such as "very basal" may arise. A'core clade' is a clade representing all but the basal clade of lowest rank within a larger clade. A basal group in the stricter sense forms a sister group to the rest of the larger clade, as in the following case: While it is easy to identify a basal clade in such a cladogram, the appropriateness of such an identification is dependent on the accuracy and completeness of the diagram, it is assumed in this example that the terminal branches of the cladogram depict all the extant taxa of a given rank within the clade. Additionally, this qualification does not ensure. In phylogenetics, the term basal can be objectively applied to clades of organisms, but tends to be applied selectively and more controversially to groups or lineages thought to possess ancestral characters, or to such presumed ancestral traits themselves. In describing characters, "ancestral" or "plesiomorphic" are preferred to "basal" or "primitive", the latter of which may carry false connotations of inferiority or a lack of complexity.
Despite the ubiquity of the usage of basal, some systematists believe its application to extant groups is unnecessary and misleading. The term is more applied when one branch is less diverse than another branch; the term may be equivocal in that it refers to the direction of the root of the tree, which represents a hypothetical ancestor. An extant basal group may or may not resemble the last common ancestor of a larger clade to a greater degree than other groups, is separated from that ancestor by the same amount of time as all other extant groups. However, there are cases where the unsually small size of a sister group does indeed correlate with an unusual number of ancestral traits, as in Amborella. Other famous examples of this phenomenon are the oviparous reproduction and nipple-less lactation of monotremes, a basal clade of mammals with just five species, the archaic anatomy of the tuatara, a basal clade of lepidosaurian with a single species; the flowering plant family Amborellaceae, restricted to New Caledonia in the southwestern Pacific, is a basal clade of extant angiosperms, consisting of the most basal species, genus and order within the group.
The traits of Amborella trichopoda are regarded as providing significant insight into the evolution of flowering plants. However, those traits are a mix of archaic and apomorphic features that have only been sorted out via comparison with other angiosperms and their positions within the phylogenetic tree. Within the primate family Hominidae, gorillas are a sister group to common chimpanzees and humans; these five species form the subfamily Homininae, of which Gorilla is the basal genus. However, if the analysis is not restricted to genera, the Homo plus Pan clade is basal. Moreover, orangutans are a sister group to Homininae and are the basal genus in the family as a whole. Subfamilies Homininae and Ponginae are both basal within Hominidae, but given that there are no nonbasal subfamilies in the cladogram it is unlikely the term would be applied to either. In general, a statement to the effect that one group is basal, or branches off first, within another group may not make sense unless the appropriate taxonomic level is specified.
If that level cannot be specified a more detailed description of the relevant sister groups may be needed. In this example, orangutans differ from the other genera in their Asian range; this fact plus their basal status provides a hint that the most recent common ancestor of extant great apes may have been Eurasian, a suggestion, consistent with other evidence. Orangutans differ from African apes in their more arboreal lifestyle, a
Binomial nomenclature called binominal nomenclature or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, although they can be based on words from other languages. Such a name is called a binomen, binominal name or a scientific name; the first part of the name – the generic name – identifies the genus to which the species belongs, while the second part – the specific name or specific epithet – identifies the species within the genus. For example, humans belong within this genus to the species Homo sapiens. Tyrannosaurus rex is the most known binomial; the formal introduction of this system of naming species is credited to Carl Linnaeus beginning with his work Species Plantarum in 1753. But Gaspard Bauhin, in as early as 1623, had introduced in his book Pinax theatri botanici many names of genera that were adopted by Linnaeus; the application of binomial nomenclature is now governed by various internationally agreed codes of rules, of which the two most important are the International Code of Zoological Nomenclature for animals and the International Code of Nomenclature for algae and plants.
Although the general principles underlying binomial nomenclature are common to these two codes, there are some differences, both in the terminology they use and in their precise rules. In modern usage, the first letter of the first part of the name, the genus, is always capitalized in writing, while that of the second part is not when derived from a proper noun such as the name of a person or place. Both parts are italicized when a binomial name occurs in normal text, thus the binomial name of the annual phlox is now written as Phlox drummondii. In scientific works, the authority for a binomial name is given, at least when it is first mentioned, the date of publication may be specified. In zoology "Patella vulgata Linnaeus, 1758"; the name "Linnaeus" tells the reader who it was that first published a description and name for this species of limpet. "Passer domesticus". The original name given by Linnaeus was Fringilla domestica; the ICZN does not require that the name of the person who changed the genus be given, nor the date on which the change was made, although nomenclatorial catalogs include such information.
In botany "Amaranthus retroflexus L." – "L." is the standard abbreviation used in botany for "Linnaeus". "Hyacinthoides italica Rothm. – Linnaeus first named this bluebell species Scilla italica. The name is composed of two word-forming elements: "bi", a Latin prefix for two, "-nomial", relating to a term or terms; the word "binomium" was used in Medieval Latin to mean a two-term expression in mathematics. Prior to the adoption of the modern binomial system of naming species, a scientific name consisted of a generic name combined with a specific name, from one to several words long. Together they formed a system of polynomial nomenclature; these names had two separate functions. First, to designate or label the species, second, to be a diagnosis or description. In a simple genus, containing only two species, it was easy to tell them apart with a one-word genus and a one-word specific name; such "polynomial names" may sometimes look like binomials, but are different. For example, Gerard's herbal describes various kinds of spiderwort: "The first is called Phalangium ramosum, Branched Spiderwort.
The other... is aptly termed Phalangium Ephemerum Virginianum, Soon-Fading Spiderwort of Virginia". The Latin phrases are short descriptions, rather than identifying labels; the Bauhins, in particular Caspar Bauhin, took some important steps towards the binomial system, by pruning the Latin descriptions, in many cases to two words. The adoption by biologists of a system of binomial nomenclature is due to Swedish botanist and physician Carl von Linné, more known by his Latinized name Carl Linnaeus, it was in his 1753 Species Plantarum that he first began using a one-word "trivial name" together with a generic name in a system of binomial nomenclature. This trivial name is what is now known as specific name; the Bauhins' genus names were retained in many of these, but the descriptive part was reduced to a single word. Linnaeus's trivial names introduced an important new idea, namely that the function of a name could be to give a species a unique label; this meant. Thus Gerard's Phalangium ephemerum virginianum became Tradescantia virgi
The Silurian is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago, to the beginning of the Devonian Period, 419.2 Mya. The Silurian is the shortest period of the Paleozoic Era; as with other geologic periods, the rock beds that define the period's start and end are well identified, but the exact dates are uncertain by several million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when 60% of marine species were wiped out. A significant evolutionary milestone during the Silurian was the diversification of jawed fish and bony fish. Multi-cellular life began to appear on land in the form of small, bryophyte-like and vascular plants that grew beside lakes and coastlines, terrestrial arthropods are first found on land during the Silurian. However, terrestrial life would not diversify and affect the landscape until the Devonian; the Silurian system was first identified by British geologist Roderick Murchison, examining fossil-bearing sedimentary rock strata in south Wales in the early 1830s.
He named the sequences for a Celtic tribe of Wales, the Silures, inspired by his friend Adam Sedgwick, who had named the period of his study the Cambrian, from the Latin name for Wales. This naming does not indicate any correlation between the occurrence of the Silurian rocks and the land inhabited by the Silures. In 1835 the two men presented a joint paper, under the title On the Silurian and Cambrian Systems, Exhibiting the Order in which the Older Sedimentary Strata Succeed each other in England and Wales, the germ of the modern geological time scale; as it was first identified, the "Silurian" series when traced farther afield came to overlap Sedgwick's "Cambrian" sequence, provoking furious disagreements that ended the friendship. Charles Lapworth resolved the conflict by defining a new Ordovician system including the contested beds. An early alternative name for the Silurian was "Gotlandian" after the strata of the Baltic island of Gotland; the French geologist Joachim Barrande, building on Murchison's work, used the term Silurian in a more comprehensive sense than was justified by subsequent knowledge.
He divided the Silurian rocks of Bohemia into eight stages. His interpretation was questioned in 1854 by Edward Forbes, the stages of Barrande, F, G and H, have since been shown to be Devonian. Despite these modifications in the original groupings of the strata, it is recognized that Barrande established Bohemia as a classic ground for the study of the earliest fossils; the Llandovery Epoch lasted from 443.8 ± 1.5 to 433.4 ± 2.8 mya, is subdivided into three stages: the Rhuddanian, lasting until 440.8 million years ago, the Aeronian, lasting to 438.5 million years ago, the Telychian. The epoch is named for the town of Llandovery in Wales; the Wenlock, which lasted from 433.4 ± 1.5 to 427.4 ± 2.8 mya, is subdivided into the Sheinwoodian and Homerian ages. It is named after Wenlock Edge in England. During the Wenlock, the oldest-known tracheophytes of the genus Cooksonia, appear; the complexity of later Gondwana plants like Baragwanathia, which resembled a modern clubmoss, indicates a much longer history for vascular plants, extending into the early Silurian or Ordovician.
The first terrestrial animals appear in the Wenlock, represented by air-breathing millipedes from Scotland. The Ludlow, lasting from 427.4 ± 1.5 to 423 ± 2.8 mya, comprises the Gorstian stage, lasting until 425.6 million years ago, the Ludfordian stage. It is named for the town of Ludlow in England; the Přídolí, lasting from 423 ± 1.5 to 419.2 ± 2.8 mya, is the final and shortest epoch of the Silurian. It is named after one locality at the Homolka a Přídolí nature reserve near the Prague suburb Slivenec in the Czech Republic. Přídolí is the old name of a cadastral field area. In North America a different suite of regional stages is sometimes used: Cayugan Lockportian Tonawandan Ontarian Alexandrian In Estonia the following suite of regional stages is used: Ohessaare stage Kaugatuma stage Kuressaare stage Paadla stage Rootsiküla stage Jaagarahu stage Jaani stage Adavere stage Raikküla stage Juuru stage With the supercontinent Gondwana covering the equator and much of the southern hemisphere, a large ocean occupied most of the northern half of the globe.
The high sea levels of the Silurian and the flat land resulted in a number of island chains, thus a rich diversity of environmental settings. During the Silurian, Gondwana continued a slow southward drift to high southern latitudes, but there is evidence that the Silurian icecaps were less extensive than those of the late-Ordovician glaciation; the southern continents remained united during this period. The melting of icecaps and glaciers contributed to a rise in sea level, recognizable from the fact that Silurian sediments overlie eroded Ordovician sediments, forming an unconformity; the continents of Avalonia and Laurentia drifted together near the equator, starting the formation of a second supercontinent known as Euramerica. When the proto-Europe coll
The Jurassic period was a geologic period and system that spanned 56 million years from the end of the Triassic Period 201.3 million years ago to the beginning of the Cretaceous Period 145 Mya. The Jurassic constitutes the middle period of the Mesozoic Era known as the Age of Reptiles; the start of the period was marked by the major Triassic–Jurassic extinction event. Two other extinction events occurred during the period: the Pliensbachian-Toarcian extinction in the Early Jurassic, the Tithonian event at the end; the Jurassic period is divided into three epochs: Early and Late. In stratigraphy, the Jurassic is divided into the Lower Jurassic, Middle Jurassic, Upper Jurassic series of rock formations; the Jurassic is named after the Jura Mountains within the European Alps, where limestone strata from the period were first identified. By the beginning of the Jurassic, the supercontinent Pangaea had begun rifting into two landmasses: Laurasia to the north, Gondwana to the south; this created more coastlines and shifted the continental climate from dry to humid, many of the arid deserts of the Triassic were replaced by lush rainforests.
On land, the fauna transitioned from the Triassic fauna, dominated by both dinosauromorph and crocodylomorph archosaurs, to one dominated by dinosaurs alone. The first birds appeared during the Jurassic, having evolved from a branch of theropod dinosaurs. Other major events include the appearance of the earliest lizards, the evolution of therian mammals, including primitive placentals. Crocodilians made the transition from a terrestrial to an aquatic mode of life; the oceans were inhabited by marine reptiles such as ichthyosaurs and plesiosaurs, while pterosaurs were the dominant flying vertebrates. The chronostratigraphic term "Jurassic" is directly linked to the Jura Mountains, a mountain range following the course of the France–Switzerland border. During a tour of the region in 1795, Alexander von Humboldt recognized the limestone dominated mountain range of the Jura Mountains as a separate formation that had not been included in the established stratigraphic system defined by Abraham Gottlob Werner, he named it "Jura-Kalkstein" in 1799.
The name "Jura" is derived from the Celtic root *jor via Gaulish *iuris "wooded mountain", borrowed into Latin as a place name, evolved into Juria and Jura. The Jurassic period is divided into three epochs: Early and Late. In stratigraphy, the Jurassic is divided into the Lower Jurassic, Middle Jurassic, Upper Jurassic series of rock formations known as Lias and Malm in Europe; the separation of the term Jurassic into three sections originated with Leopold von Buch. The faunal stages from youngest to oldest are: During the early Jurassic period, the supercontinent Pangaea broke up into the northern supercontinent Laurasia and the southern supercontinent Gondwana; the Jurassic North Atlantic Ocean was narrow, while the South Atlantic did not open until the following Cretaceous period, when Gondwana itself rifted apart. The Tethys Sea closed, the Neotethys basin appeared. Climates were warm, with no evidence of a glacier having appeared; as in the Triassic, there was no land over either pole, no extensive ice caps existed.
The Jurassic geological record is good in western Europe, where extensive marine sequences indicate a time when much of that future landmass was submerged under shallow tropical seas. In contrast, the North American Jurassic record is the poorest of the Mesozoic, with few outcrops at the surface. Though the epicontinental Sundance Sea left marine deposits in parts of the northern plains of the United States and Canada during the late Jurassic, most exposed sediments from this period are continental, such as the alluvial deposits of the Morrison Formation; the Jurassic was a time of calcite sea geochemistry in which low-magnesium calcite was the primary inorganic marine precipitate of calcium carbonate. Carbonate hardgrounds were thus common, along with calcitic ooids, calcitic cements, invertebrate faunas with dominantly calcitic skeletons; the first of several massive batholiths were emplaced in the northern American cordillera beginning in the mid-Jurassic, marking the Nevadan orogeny. Important Jurassic exposures are found in Russia, South America, Japan and the United Kingdom.
In Africa, Early Jurassic strata are distributed in a similar fashion to Late Triassic beds, with more common outcrops in the south and less common fossil beds which are predominated by tracks to the north. As the Jurassic proceeded and more iconic groups of dinosaurs like sauropods and ornithopods proliferated in Africa. Middle Jurassic strata are neither well studied in Africa. Late Jurassic strata are poorly represented apart from the spectacular Tendaguru fauna in Tanzania; the Late Jurassic life of Tendaguru is similar to that found in western North America's Morrison Formation. During the Jurassic period, the primary vertebrates living in the sea were marine reptiles; the latter include ichthyosaurs, which were at the peak of their diversity, plesiosaurs and marine crocodiles of the families Teleosauridae and Metriorhynchidae. Numerous turtles could be found in rivers. In the invertebrate world, several new groups appeared, including rudists (a reef-formi
The vertebral column known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord found in all chordates has been replaced by a segmented series of bone: vertebrae separated by intervertebral discs; the vertebral column houses a cavity that encloses and protects the spinal cord. There are about 50,000 species of animals; the human vertebral column is one of the most-studied examples. In a human's vertebral column there are thirty-three vertebrae; the articulating vertebrae are named according to their region of the spine. There are twelve thoracic vertebrae and five lumbar vertebrae; the number of vertebrae in a region overall the number remains the same. The number of those in the cervical region however is only changed. There are ligaments extending the length of the column at the front and the back, in between the vertebrae joining the spinous processes, the transverse processes and the vertebral laminae.
The vertebrae in the human vertebral column are divided into different regions, which correspond to the curves of the spinal column. The articulating vertebrae are named according to their region of the spine. Vertebrae in these regions are alike, with minor variation; these regions are called the cervical spine, thoracic spine, lumbar spine and coccyx. There are twelve thoracic vertebrae and five lumbar vertebrae; the number of vertebrae in a region overall the number remains the same. The number of those in the cervical region however is only changed; the vertebrae of the cervical and lumbar spines are independent bones, quite similar. The vertebrae of the sacrum and coccyx are fused and unable to move independently. Two special vertebrae are the axis, on which the head rests. A typical vertebra consists of two parts: the vertebral arch; the vertebral arch is posterior. Together, these enclose the vertebral foramen; because the spinal cord ends in the lumbar spine, the sacrum and coccyx are fused, they do not contain a central foramen.
The vertebral arch is formed by a pair of pedicles and a pair of laminae, supports seven processes, four articular, two transverse, one spinous, the latter being known as the neural spine. Two transverse processes and one spinous process are posterior to the vertebral body; the spinous process comes out the back, one transverse process comes out the left, one on the right. The spinous processes of the cervical and lumbar regions can be felt through the skin. Above and below each vertebra are joints called facet joints; these restrict the range of movement possible, are joined by a thin portion of the neural arch called the pars interarticularis. In between each pair of vertebrae are two small holes called intervertebral foramina; the spinal nerves leave the spinal cord through these holes. Individual vertebrae are named according to their position. From top to bottom, the vertebrae are: Cervical spine: 7 vertebrae Thoracic spine: 12 vertebrae Lumbar spine: 5 vertebrae Sacrum: 5 vertebrae Coccyx: 4 vertebrae The upper cervical spine has a curve, convex forward, that begins at the axis at the apex of the odontoid process or dens, ends at the middle of the second thoracic vertebra.
This inward curve is known as a lordotic curve. The thoracic curve, concave forward, begins at the middle of the second and ends at the middle of the twelfth thoracic vertebra, its most prominent point behind corresponds to the spinous process of the seventh thoracic vertebra. This curve is known as a kyphotic curve; the lumbar curve is more marked in the female than in the male. It is convex anteriorly, the convexity of the lower three vertebrae being much greater than that of the upper two; this curve is described as a lordotic curve. The sacral curve begins at the sacrovertebral articulation, ends at the point of the coccyx; the thoracic and sacral kyphotic curves are termed primary curves, because they are present in the fetus. The cervical and lumbar curves are compensatory or secondary, are developed after birth; the cervical curve forms when the infant is able to sit upright. The lumbar curve forms from twelve to eighteen months, when the child begins to walk. Anterior surfaceWhen viewed from in front, the width of the bodies of the vertebrae is seen to increase from the second cervical to the first thoracic.
From this point there is a rapid diminution, to the apex of the coccyx. Posterior surfaceFrom behind, the vertebral column presents in the median line the spinous processes. In the cervical region these are short and bifid. In the upper part of the thoracic region they are directed obliquely downward.
The Cretaceous is a geologic period and system that spans 79 million years from the end of the Jurassic Period 145 million years ago to the beginning of the Paleogene Period 66 mya. It is the last period of the Mesozoic Era, the longest period of the Phanerozoic Eon; the Cretaceous Period is abbreviated K, for its German translation Kreide. The Cretaceous was a period with a warm climate, resulting in high eustatic sea levels that created numerous shallow inland seas; these oceans and seas were populated with now-extinct marine reptiles and rudists, while dinosaurs continued to dominate on land. During this time, new groups of mammals and birds, as well as flowering plants, appeared; the Cretaceous ended with the Cretaceous–Paleogene extinction event, a large mass extinction in which many groups, including non-avian dinosaurs and large marine reptiles died out. The end of the Cretaceous is defined by the abrupt Cretaceous–Paleogene boundary, a geologic signature associated with the mass extinction which lies between the Mesozoic and Cenozoic eras.
The Cretaceous as a separate period was first defined by Belgian geologist Jean d'Omalius d'Halloy in 1822, using strata in the Paris Basin and named for the extensive beds of chalk, found in the upper Cretaceous of Western Europe. The name Cretaceous was derived from Latin creta; the Cretaceous is divided into Early and Late Cretaceous epochs, or Lower and Upper Cretaceous series. In older literature the Cretaceous is sometimes divided into three series: Neocomian and Senonian. A subdivision in eleven stages, all originating from European stratigraphy, is now used worldwide. In many parts of the world, alternative local subdivisions are still in use; as with other older geologic periods, the rock beds of the Cretaceous are well identified but the exact age of the system's base is uncertain by a few million years. No great extinction or burst of diversity separates the Cretaceous from the Jurassic. However, the top of the system is defined, being placed at an iridium-rich layer found worldwide, believed to be associated with the Chicxulub impact crater, with its boundaries circumscribing parts of the Yucatán Peninsula and into the Gulf of Mexico.
This layer has been dated at 66.043 Ma. A 140 Ma age for the Jurassic-Cretaceous boundary instead of the accepted 145 Ma was proposed in 2014 based on a stratigraphic study of Vaca Muerta Formation in Neuquén Basin, Argentina. Víctor Ramos, one of the authors of the study proposing the 140 Ma boundary age sees the study as a "first step" toward formally changing the age in the International Union of Geological Sciences. From youngest to oldest, the subdivisions of the Cretaceous period are: Late Cretaceous Maastrichtian – Campanian – Santonian – Coniacian – Turonian – Cenomanian – Early Cretaceous Albian – Aptian – Barremian – Hauterivian – Valanginian – Berriasian – The high sea level and warm climate of the Cretaceous meant large areas of the continents were covered by warm, shallow seas, providing habitat for many marine organisms; the Cretaceous was named for the extensive chalk deposits of this age in Europe, but in many parts of the world, the deposits from the Cretaceous are of marine limestone, a rock type, formed under warm, shallow marine circumstances.
Due to the high sea level, there was extensive space for such sedimentation. Because of the young age and great thickness of the system, Cretaceous rocks are evident in many areas worldwide. Chalk is a rock type characteristic for the Cretaceous, it consists of coccoliths, microscopically small calcite skeletons of coccolithophores, a type of algae that prospered in the Cretaceous seas. In northwestern Europe, chalk deposits from the Upper Cretaceous are characteristic for the Chalk Group, which forms the white cliffs of Dover on the south coast of England and similar cliffs on the French Normandian coast; the group is found in England, northern France, the low countries, northern Germany, Denmark and in the subsurface of the southern part of the North Sea. Chalk is not consolidated and the Chalk Group still consists of loose sediments in many places; the group has other limestones and arenites. Among the fossils it contains are sea urchins, belemnites and sea reptiles such as Mosasaurus. In southern Europe, the Cretaceous is a marine system consisting of competent limestone beds or incompetent marls.
Because the Alpine mountain chains did not yet exist in the Cretaceous, these deposits formed on the southern edge of the European continental shelf, at the margin of the Tethys Ocean. Stagnation of deep sea currents in middle Cretaceous times caused anoxic conditions in the sea water leaving the deposited organic matter undecomposed. Half the worlds petroleum reserves were laid down at this time in the anoxic conditions of what would become the Persian Gulf and the Gulf of Mexico. In many places around the world, dark anoxic shales were formed during this interval; these shales are an important source rock for oil and gas, for example in the subsurface of the North Sea. During th
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