The Permian is a geologic period and system which spans 47 million years from the end of the Carboniferous Period 298.9 million years ago, to the beginning of the Triassic period 251.902 Mya. It is the last period of the Paleozoic era; the concept of the Permian was introduced in 1841 by geologist Sir Roderick Murchison, who named it after the city of Perm. The Permian witnessed the diversification of the early amniotes into the ancestral groups of the mammals, turtles and archosaurs; the world at the time was dominated by two continents known as Pangaea and Siberia, surrounded by a global ocean called Panthalassa. The Carboniferous rainforest collapse left behind vast regions of desert within the continental interior. Amniotes, who could better cope with these drier conditions, rose to dominance in place of their amphibian ancestors; the Permian ended with the Permian–Triassic extinction event, the largest mass extinction in Earth's history, in which nearly 96% of marine species and 70% of terrestrial species died out.
It would take well into the Triassic for life to recover from this catastrophe. Recovery from the Permian–Triassic extinction event was protracted; the term "Permian" was introduced into geology in 1841 by Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Édouard de Verneuil; the region now lies in the Perm Krai of Russia. Official ICS 2017 subdivisions of the Permian System from most recent to most ancient rock layers are: Lopingian epoch Changhsingian Wuchiapingian Others: Waiitian Makabewan Ochoan Guadalupian epoch Capitanian stage Wordian stage Roadian stage Others: Kazanian or Maokovian Braxtonian stage Cisuralian epoch Kungurian stage Artinskian stage Sakmarian stage Asselian stage Others: Telfordian Mangapirian Sea levels in the Permian remained low, near-shore environments were reduced as all major landmasses collected into a single continent—Pangaea; this could have in part caused the widespread extinctions of marine species at the end of the period by reducing shallow coastal areas preferred by many marine organisms.
During the Permian, all the Earth's major landmasses were collected into a single supercontinent known as Pangaea. Pangaea straddled the equator and extended toward the poles, with a corresponding effect on ocean currents in the single great ocean, the Paleo-Tethys Ocean, a large ocean that existed between Asia and Gondwana; the Cimmeria continent rifted away from Gondwana and drifted north to Laurasia, causing the Paleo-Tethys Ocean to shrink. A new ocean was growing on its southern end, the Tethys Ocean, an ocean that would dominate much of the Mesozoic era. Large continental landmass interiors experience climates with extreme variations of heat and cold and monsoon conditions with seasonal rainfall patterns. Deserts seem to have been widespread on Pangaea; such dry conditions favored gymnosperms, plants with seeds enclosed in a protective cover, over plants such as ferns that disperse spores in a wetter environment. The first modern trees appeared in the Permian. Three general areas are noted for their extensive Permian deposits—the Ural Mountains and the southwest of North America, including the Texas red beds.
The Permian Basin in the U. S. states of Texas and New Mexico is so named because it has one of the thickest deposits of Permian rocks in the world. The climate in the Permian was quite varied. At the start of the Permian, the Earth was still in an ice age. Glaciers receded around the mid-Permian period as the climate warmed, drying the continent's interiors. In the late Permian period, the drying continued although the temperature cycled between warm and cool cycles. Permian marine deposits are rich in fossil mollusks and brachiopods. Fossilized shells of two kinds of invertebrates are used to identify Permian strata and correlate them between sites: fusulinids, a kind of shelled amoeba-like protist, one of the foraminiferans, ammonoids, shelled cephalopods that are distant relatives of the modern nautilus. By the close of the Permian, trilobites and a host of other marine groups became extinct. Terrestrial life in the Permian included diverse plants, fungi and various types of tetrapods; the period saw a massive desert covering the interior of Pangaea.
The warm zone spread in the northern hemisphere. The rocks formed at that time were stained red by iron oxides, the result of intense heating by the sun of a surface devoid of vegetation cover. A number of older types of plants and animals became marginal elements; the Permian began with the Carboniferous flora still flourishing. About the middle of the Permian a major transition in vegetation began; the swamp-loving
A chordate is an animal constituting the phylum Chordata. During some period of their life cycle, chordates possess a notochord, a dorsal nerve cord, pharyngeal slits, an endostyle, a post-anal tail: these five anatomical features define this phylum. Chordates are bilaterally symmetric; the Chordata and Ambulacraria together form the superphylum Deuterostomia. Chordates are divided into three subphyla: Vertebrata. There are extinct taxa such as the Vetulicolia. Hemichordata has been presented as a fourth chordate subphylum, but now is treated as a separate phylum: hemichordates and Echinodermata form the Ambulacraria, the sister phylum of the Chordates. Of the more than 65,000 living species of chordates, about half are bony fish that are members of the superclass Osteichthyes. Chordate fossils have been found from as early as the Cambrian explosion, 541 million years ago. Cladistically, vertebrates - chordates with the notochord replaced by a vertebral column during development - are considered to be a subgroup of the clade Craniata, which consists of chordates with a skull.
The Craniata and Tunicata compose the clade Olfactores. Chordates form a phylum of animals that are defined by having at some stage in their lives all of the following anatomical features: A notochord, a stiff rod of cartilage that extends along the inside of the body. Among the vertebrate sub-group of chordates the notochord develops into the spine, in wholly aquatic species this helps the animal to swim by flexing its tail. A dorsal neural tube. In fish and other vertebrates, this develops into the spinal cord, the main communications trunk of the nervous system. Pharyngeal slits; the pharynx is the part of the throat behind the mouth. In fish, the slits are modified to form gills, but in some other chordates they are part of a filter-feeding system that extracts particles of food from the water in which the animals live. Post-anal tail. A muscular tail that extends backwards behind the anus. An endostyle; this is a groove in the ventral wall of the pharynx. In filter-feeding species it produces mucus to gather food particles, which helps in transporting food to the esophagus.
It stores iodine, may be a precursor of the vertebrate thyroid gland. There are soft constraints that separate chordates from certain other biological lineages, but are not part of the formal definition: All chordates are deuterostomes; this means. All chordates are based on a bilateral body plan. All chordates are coelomates, have a fluid filled body cavity called a coelom with a complete lining called peritoneum derived from mesoderm; the following schema is from the third edition of Vertebrate Palaeontology. The invertebrate chordate classes are from Fishes of the World. While it is structured so as to reflect evolutionary relationships, it retains the traditional ranks used in Linnaean taxonomy. Phylum Chordata †Vetulicolia? Subphylum Cephalochordata – Class Leptocardii Clade Olfactores Subphylum Tunicata – Class Ascidiacea Class Thaliacea Class Appendicularia Class Sorberacea Subphylum Vertebrata Infraphylum incertae sedis Cyclostomata Superclass'Agnatha' paraphyletic Class Myxini Class Petromyzontida or Hyperoartia Class †Conodonta Class †Myllokunmingiida Class †Pteraspidomorphi Class †Thelodonti Class †Anaspida Class †Cephalaspidomorphi Infraphylum Gnathostomata Class †Placodermi Class Chondrichthyes Class †Acanthodii Superclass Osteichthyes Class Actinopterygii Class Sarcopterygii Superclass Tetrapoda Class Amphibia Class Sauropsida Class Synapsida Craniates, one of the three subdivisions of chordates, all have distinct skulls.
They include the hagfish. Michael J. Benton commented that "craniates are characterized by their heads, just as chordates, or all deuterostomes, are by their tails". Most craniates are vertebrates; these consist of a series of bony or cartilaginous cylindrical vertebrae with neural arches that protect the spinal cord, with projections that link the vertebrae. However hagfish have incomplete braincases and no vertebrae, are therefore not regarded as vertebrates, but as members of the craniates, the group from which vertebrates are thought to have evolved; however the cladistic exclusion of hagfish from the vertebrates is controversial, as they ma
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
Cartilage is a resilient and smooth elastic tissue, a rubber-like padding that covers and protects the ends of long bones at the joints, is a structural component of the rib cage, the ear, the nose, the bronchial tubes, the intervertebral discs, many other body components. It is not as hard and rigid as bone; the matrix of cartilage is made up of glycosaminoglycans, collagen fibers and, elastin. Because of its rigidity, cartilage serves the purpose of holding tubes open in the body. Examples include the rings such as the cricoid cartilage and carina. Cartilage is composed of specialized cells called chondrocytes that produce a large amount of collagenous extracellular matrix, abundant ground substance, rich in proteoglycan and elastin fibers. Cartilage is classified in three types, elastic cartilage, hyaline cartilage and fibrocartilage, which differ in relative amounts of collagen and proteoglycan. Cartilage does not contain blood nerves. Nutrition is supplied to the chondrocytes by diffusion.
The compression of the articular cartilage or flexion of the elastic cartilage generates fluid flow, which assists diffusion of nutrients to the chondrocytes. Compared to other connective tissues, cartilage has a slow turnover of its extracellular matrix and does not repair. In embryogenesis, the skeletal system is derived from the mesoderm germ layer. Chondrification is the process by which cartilage is formed from condensed mesenchyme tissue, which differentiates into chondroblasts and begins secreting the molecules that form the extracellular matrix. Following the initial chondrification that occurs during embryogenesis, cartilage growth consists of the maturing of immature cartilage to a more mature state; the division of cells within cartilage occurs slowly, thus growth in cartilage is not based on an increase in size or mass of the cartilage itself. The articular cartilage function is dependent on the molecular composition of the extracellular matrix; the ECM consists of proteoglycan and collagens.
The main proteoglycan in cartilage is aggrecan, which, as its name suggests, forms large aggregates with hyaluronan. These aggregates hold water in the tissue; the collagen collagen type II, constrains the proteoglycans. The ECM responds to compressive forces that are experienced by the cartilage. Cartilage growth thus refers to the matrix deposition, but can refer to both the growth and remodeling of the extracellular matrix. Due to the great stress on the patellofemoral joint during resisted knee extension, the articular cartilage of the patella is among the thickest in the human body; the mechanical properties of articular cartilage in load-bearing joints such as the knee and hip have been studied extensively at macro and nano-scales. These mechanical properties include the response of cartilage in frictional, compressive and tensile loading. Cartilage displays viscoelastic properties. Lubricin, a glycoprotein abundant in cartilage and synovial fluid, plays a major role in bio-lubrication and wear protection of cartilage.
Cartilage has limited repair capabilities: Because chondrocytes are bound in lacunae, they cannot migrate to damaged areas. Therefore, cartilage damage is difficult to heal; because hyaline cartilage does not have a blood supply, the deposition of new matrix is slow. Damaged hyaline cartilage is replaced by fibrocartilage scar tissue. Over the last years and scientists have elaborated a series of cartilage repair procedures that help to postpone the need for joint replacement. Bioengineering techniques are being developed to generate new cartilage, using a cellular "scaffolding" material and cultured cells to grow artificial cartilage. Several diseases can affect cartilage. Chondrodystrophies are a group of diseases, characterized by the disturbance of growth and subsequent ossification of cartilage; some common diseases that affect the cartilage are listed below. Osteoarthritis: Osteoarthritis is a disease of the whole joint, however one of the most affected tissues is the articular cartilage.
The cartilage covering bones is thinned completely wearing away, resulting in a "bone against bone" within the joint, leading to reduced motion, pain. Osteoarthritis affects the joints exposed to high stress and is therefore considered the result of "wear and tear" rather than a true disease, it is treated by arthroplasty, the replacement of the joint by a synthetic joint made of a stainless steel alloy and ultra high molecular weight polyethylene. Chondroitin sulfate or glucosamine sulfate supplements, have been claimed to reduce the symptoms of osteoarthritis but there is little good evidence to support this claim. Traumatic rupture or detachment: The cartilage in the knee is damaged but can be repaired through knee cartilage replacement therapy; when athletes talk of damaged "cartilage" in their knee, they are referring to a damaged meniscus and not the articular cartilage. Achondroplasia: Reduced proliferation of chondrocytes in the epiphyseal plate of long bones during infancy and childhood, resulting in dwarfism.
Costochondritis: Inflammation of cartilage in the ribs, causing chest pain. Spinal disc herniation: Asymmetrical compression of an intervertebral disc ruptures the sac-like disc, causing a herniation of its soft content; the hernia compresses the adjacent nerves and causes back pain. Relapsing polychondritis: a destruction aut
The Lamniformes are an order of sharks known as mackerel sharks. It includes some of the most familiar species of sharks, such as the great white and extinct megalodon, as well as more unusual representatives, such as the goblin shark and megamouth shark. Members of the order are distinguished by possessing two dorsal fins, an anal fin, five gill slits, eyes without nictitating membranes, a mouth extending behind the eyes. Unlike other sharks, they maintain a higher body temperature than the surrounding water; the order Lamniformes includes 10 families with 22 species, with a total of seven living families and 17 living species: Order Lamniformes Family Alopiidae Bonaparte, 1838 Genus Alopias Rafinesque, 1810 Alopias pelagicus Nakamura, 1935 Alopias superciliosus R. T. Lowe, 1841 Alopias vulpinus Alopias sp. not yet described Family Anacoracidae Capetta, 1987 † Genus Squalicorax Genus Telodontaspis Genus Pseudocorax Genus Galeocorax Genus Scindocorax Genus Nanocorax Genus Ptychocorax Family Cetorhinidae Gill, 1862 Genus Cetorhinus Blainville, 1816 Cetorhinus maximus Family Eoptolamnidae*† Genus Leptostyrax*†Leptostyrax macrorhiza Family Lamnidae J. P. Müller and Henle, 1838 Genus Carcharodon A. Smith, 1838 Carcharodon carcharias Carcharodon hubbelli † Genus Isurus Rafinesque, 1810 Isurus oxyrinchus Rafinesque, 1810 Isurus paucus Guitart-Manday, 1966 Genus Lamna Cuvier, 1816 Lamna ditropis Hubbs & Follett, 1947 Lamna nasus Family Megachasmidae Taylor, Compagno & Struhsaker, 1983 Genus Megachasma Taylor, Compagno & Struhsaker, 1983 Megachasma pelagios Taylor, Compagno & Struhsaker, 1983 Family Mitsukurinidae D. S. Jordan, 1898 Genus Mitsukurina D. S. Jordan, 1898 Mitsukurina owstoni D. S. Jordan, 1898 Family Odontaspididae Müller & Henle, 1839 Genus Carcharias Rafinesque, 1810 Carcharias taurus Rafinesque, 1810 Genus Odontaspis Agassiz, 1838 Odontaspis ferox Odontaspis noronhai Family Pseudocarchariidae Compagno, 1973 Genus Pseudocarcharias Cadenat, 1963 Pseudocarcharias kamoharai Family Cardabiodontidae † Genus Cardabiodon Siverson, 1999 Cardabiodon ricki Siverson, 1999 † Cardabiodon venator Siverson and Lindgren, 2005 † Family Cretoxyrhinidae † Genus Cretoxyrhina Agassiz, 1843 Cretoxyrhina vraconensis Zhelezko, 2000 † Cretoxyrhina denticulata Glückman, 1957 † Cretoxyrhina agassizensis Underwood and Cumbaa, 2010 † Cretoxyrhina mantelli Agassiz, 1843 † Family Otodontidae † Genus Carcharocles Carcharocles auriculatus Carcharocles angustidens Carcharocles chubutensis Carcharocles megalodon † In 2010, Greenpeace International added the shortfin mako shark to its seafood red list.
Compagno, Leonard Sharks of the World: Bullhead and carpet sharks Volume 2, FAO Species Catalogue, Rome. ISBN 92-5-104543-7. Joseph S. Nelson. "Order Lamniformes". Fishes of the World. John Wiley and Sons. Pp. 57–60. ISBN 978-0-471-25031-9. Elasmo-research
Crested bullhead shark
The crested bullhead shark is an uncommon species of bullhead shark, in the family Heterodontidae. It lives off the coast of eastern Australia from the coast to a depth of 93 m; this shark can be distinguished from other members of its family by the large size of the ridges above its eyes and by its color pattern of large dark blotches. It attains a length of 1.2 m. Nocturnal and bottom-dwelling, the crested bullhead shark favors rocky reefs and vegetated areas, where it hunts for sea urchins and other small organisms, it is oviparous, with females producing auger-shaped egg capsules that are secured to seaweed or sponges with long tendrils. Sexual maturation is slow, with one female in captivity not laying eggs until 12 years of age; the International Union for Conservation of Nature has assessed this harmless shark as of Least Concern. British zoologist Albert Günther described the crested bullhead shark as Cestracion galeatus in the 1870 eighth volume of Catalogue of the Fishes in the British Museum.
He chose the specific epithet galeatus from the Latin for "helmeted", referring to the prominent ridges above the shark's eyes that give it its common name. Subsequent authors moved this species to the genera Gyropleurodus and Molochophrys before placing it in Heterodontus; the type specimen is a 68-cm-long female caught off Australia. This shark may be referred to as crested shark, crested bull shark, crested horn shark, crested Port Jackson shark; the head of the crested bullhead shark is wide, with a blunt, pig-like snout. The eyes are placed high on the lack nictitating membranes; the supraorbital ridges above the eyes of this species are larger than any other member of its family. The nostrils are separated into incurrent and excurrent openings by a long flap of skin that reaches the mouth. A furrow encircles the incurrent opening and another furrow runs from the excurrent opening to the mouth, located nearly at the tip of the snout; the teeth at the front of the jaws are small and pointed with a central cusp and two lateral cusplets, while those at the back of the jaws are wide and molar-like.
The deep furrows at the corners of the mouth extend onto both jaws. The pectoral fins are large and rounded, while the pelvic and anal fins are smaller and more angular; the first dorsal fin is moderately tall with a rounded to angular apex and a stout spine on the leading margin, originating behind the pectoral fins. The second dorsal fin resembles the first and is as large, is located between the pelvic and anal fins; the caudal fin is broad, with a strong ventral notch near the tip of the upper lobe. The dermal denticles are large and rough on the flanks; the coloration consists of five brown to diffusely edged saddles on a light tan background. There are dark marks below each eye. Most crested bullhead sharks measure no more than 1.2 m long. The range of the crested bullhead shark is restricted to the warm temperate waters along the eastern coast of Australia, from Cape Moreton, Queensland, to Batemans Bay, New South Wales. Dubious records exist of this species from off Cape York Peninsula in the north and Tasmania in the south.
This species co-occurs with the related Port Jackson shark across much of its range, but is much rarer except off southern Queensland and northern New South Wales, where it tends to replace the other species. Bottom-dwelling in nature, the crested bullhead shark is found over the continental shelf from the intertidal zone to a depth of 93 m, being more common in deeper waters, it prefers rocky reefs, mats of seaweed, seagrass beds. The crested bullhead shark is a slow-moving, nocturnal species seen wedging its head between rocks in search of food, it feeds on the sea urchins Centrostephanus rodgersii and Heliocidaris erythrogramma, but has been known to take a variety of other invertebrates and small fishes. A steady diet of sea urchins may stain the teeth of this shark pinkish purple; the crested bullhead shark is a major predator of the eggs of the Port Jackson shark, which are seasonally available and rich in nutrients. Individual sharks have been observed taking the egg capsules in their mouths and chewing on the tough casing, rupturing it and allowing the contents to be sucked out.
Unlike the Port Jackson shark, the crested bullhead shark is not known to form large aggregations. Crested bullhead sharks are oviparous with a annual reproductive cycle. Females produce 10–16 eggs per year during late winter in July and August, though Michael noted that egg-laying may continue year-round; the egg cases measure around 11 cm in length, with a pair of thin flanges spiraling 6–7 times around the outside and two slender tendrils up to 2 m long at one end, used to attach the capsule to seaweed or sponges. The capsules are deposited at a depth of 20–30 m, much deeper than the Port Jackson shark, though there is a single record of an egg being found only 8.6 m down. The time to hatching has been variously reported as 8 -- 9 months. Last and Stevens gave the lengths at maturity for males and females at 60 cm and 70 cm though mature males as small as 53.5 cm long were found off Queensland. Growth and aging has been documented for one captive female at the Taronga Park Aquarium, which grew an average of 5 cm per year and did n
The Neogene is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period 23.03 million years ago to the beginning of the present Quaternary Period 2.58 Mya. The Neogene is sub-divided into two epochs, the earlier Miocene and the Pliocene; some geologists assert that the Neogene cannot be delineated from the modern geological period, the Quaternary. The term "Neogene" was coined in 1853 by the Austrian palaeontologist Moritz Hörnes. During this period and birds continued to evolve into modern forms, while other groups of life remained unchanged. Early hominids, the ancestors of humans, appeared in Africa near the end of the period; some continental movement took place, the most significant event being the connection of North and South America at the Isthmus of Panama, late in the Pliocene. This cut off the warm ocean currents from the Pacific to the Atlantic Ocean, leaving only the Gulf Stream to transfer heat to the Arctic Ocean; the global climate cooled over the course of the Neogene, culminating in a series of continental glaciations in the Quaternary Period that follows.
In ICS terminology, from upper to lower: The Pliocene Epoch is subdivided into 2 ages: Piacenzian Age, preceded by Zanclean AgeThe Miocene Epoch is subdivided into 6 ages: Messinian Age, preceded by Tortonian Age Serravallian Age Langhian Age Burdigalian Age Aquitanian AgeIn different geophysical regions of the world, other regional names are used for the same or overlapping ages and other timeline subdivisions. The terms Neogene System and upper Tertiary System describe the rocks deposited during the Neogene Period; the continents in the Neogene were close to their current positions. The Isthmus of Panama formed, connecting South America; the Indian subcontinent continued forming the Himalayas. Sea levels fell, creating land bridges between Africa and Eurasia and between Eurasia and North America; the global climate became seasonal and continued an overall drying and cooling trend which began at the start of the Paleogene. The ice caps on both poles began to grow and thicken, by the end of the period the first of a series of glaciations of the current Ice Age began.
Marine and continental flora and fauna have a modern appearance. The reptile group Choristodera became extinct in the early part of the period, while the amphibians known as Allocaudata disappeared at the end. Mammals and birds continued to be the dominant terrestrial vertebrates, took many forms as they adapted to various habitats; the first hominins, the ancestors of humans, may have appeared in southern Europe and migrated into Africa. In response to the cooler, seasonal climate, tropical plant species gave way to deciduous ones and grasslands replaced many forests. Grasses therefore diversified, herbivorous mammals evolved alongside it, creating the many grazing animals of today such as horses and bison. Eucalyptus fossil leaves occur in the Miocene of New Zealand, where the genus is not native today, but have been introduced from Australia; the Neogene traditionally ended at the end of the Pliocene Epoch, just before the older definition of the beginning of the Quaternary Period. However, there was a movement amongst geologists to include ongoing geological time in the Neogene, while others insist the Quaternary to be a separate period of distinctly different record.
The somewhat confusing terminology and disagreement amongst geologists on where to draw what hierarchical boundaries is due to the comparatively fine divisibility of time units as time approaches the present, due to geological preservation that causes the youngest sedimentary geological record to be preserved over a much larger area and to reflect many more environments than the older geological record. By dividing the Cenozoic Era into three periods instead of seven epochs, the periods are more comparable to the duration of periods in the Mesozoic and Paleozoic eras; the International Commission on Stratigraphy once proposed that the Quaternary be considered a sub-era of the Neogene, with a beginning date of 2.58 Ma, namely the start of the Gelasian Stage. In the 2004 proposal of the ICS, the Neogene would have consisted of the Miocene and Pliocene epochs; the International Union for Quaternary Research counterproposed that the Neogene and the Pliocene end at 2.58 Ma, that the Gelasian be transferred to the Pleistocene, the Quaternary be recognized as the third period in the Cenozoic, citing key changes in Earth's climate and biota that occurred 2.58 Ma and its correspondence to the Gauss-Matuyama magnetostratigraphic boundary.
In 2006 ICS and INQUA reached a compromise that made Quaternary a subera, subdividing Cenozoic into the old classical Tertiary and Quaternary, a compromise, rejected by International Union of Geological Sciences because it split both Neogene and Pliocene in two. Following formal discussions at the 2008 International Geological Congress in Oslo, the ICS decided in May 2009 to make the Quaternary the youngest period of the Cenozoic Era with its base at 2.58 Mya and including the Gelasian age, considered part of the Neogene Period and Pliocene Epoch. Thus the Neogene Period ends bounding the succeeding Quaternary Period at 2.58 Mya. "Digital Atlas of Neogene Life for the Southeastern United States". San Jose State University. Archived from the original on 2013-04-23. Retrieved 21 September 2018