A stem is one of two main structural axes of a vascular plant, the other being the root. The stem is divided into nodes and internodes: The nodes hold one or more leaves, as well as buds which can grow into branches. Adventitious roots may be produced from the nodes; the internodes distance one node from another. The term "shoots" is confused with "stems". In most plants stems are located above the soil surface but some plants have underground stems. Stems have four main functions which are: Support for and the elevation of leaves and fruits; the stems keep the leaves in the light and provide a place for the plant to keep its flowers and fruits. Transport of fluids between the roots and the shoots in the xylem and phloem Storage of nutrients Production of new living tissue; the normal lifespan of plant cells is one to three years. Stems have cells called meristems. Stems are specialized for storage, asexual reproduction, protection or photosynthesis, including the following: Acaulescent – used to describe stems in plants that appear to be stemless.
These stems are just short, the leaves appearing to rise directly out of the ground, e.g. some Viola species. Arborescent – tree like with woody stems with a single trunk. Axillary bud – a bud which grows at the point of attachment of an older leaf with the stem, it gives rise to a shoot. Branched – aerial stems are described as being branched or unbranched Bud – an embryonic shoot with immature stem tip. Bulb – a short vertical underground stem with fleshy storage leaves attached, e.g. onion, tulip. Bulbs function in reproduction by splitting to form new bulbs or producing small new bulbs termed bulblets. Bulbs are a combination of stem and leaves so may better be considered as leaves because the leaves make up the greater part. Caespitose – when stems grow in a tangled mass or clump or in low growing mats. Cladode – a flattened stem that appears more-or-less leaf like and is specialized for photosynthesis, e.g. cactus pads. Climbing -- stems that wrap around other plants or structures. Corm – a short enlarged underground, storage stem, e.g. taro, gladiolus.
Decumbent -- stems that lie flat on the turn upwards at the ends. Fruticose -- stems. Herbaceous – non woody, they die at the end of the growing season. Internode – an interval between two successive nodes, it possesses the ability to elongate, either from its base or from its extremity depending on the species. Node – a point of attachment of a leaf or a twig on the stem in seed plants. A node is a small growth zone. Pedicel – stems that serve as the stalk of an individual flower in an inflorescence or infrutescence. Peduncle – a stem that supports an inflorescence Prickle – a sharpened extension of the stem's outer layers, e.g. roses. Pseudostem – a false stem made of the rolled bases of leaves, which may be 2 or 3 m tall as in banana Rhizome – a horizontal underground stem that functions in reproduction but in storage, e.g. most ferns, iris Runner – a type of stolon, horizontally growing on top of the ground and rooting at the nodes, aids in reproduction. E.g. garden strawberry, Chlorophytum comosum.
Scape – a stem that holds flowers that comes out of the ground and has no normal leaves. Hosta, Iris, Garlic. Stolon – a horizontal stem that produces rooted plantlets at its nodes and ends, forming near the surface of the ground. Thorn – a modified stem with a sharpened point. Tuber – a swollen, underground storage stem adapted for storage and reproduction, e.g. potato. Woody – hard textured stems with secondary xylem. Stem consist of three tissues, dermal tissue, ground tissue and vascular tissue; the dermal tissue covers the outer surface of the stem and functions to waterproof and control gas exchange. The ground tissue consists of parenchyma cells and fills in around the vascular tissue, it sometimes functions in photosynthesis. Vascular tissue provides structural support. Most or all ground tissue may be lost in woody stems; the dermal tissue of aquatic plants stems. The arrangement of the vascular tissues varies among plant species. Dicot stems with primary growth have pith in the center, with vascular bundles forming a distinct ring visible when the stem is viewed in cross section.
The outside of the stem is covered with an epidermis, covered by a waterproof cuticle. The epidermis may contain stomata for gas exchange and multicellular stem hairs called trichomes. A cortex consisting of hypodermis and endodermis is present above the pericycle and vascular bundles. Woody dicots and many nonwoody dicots have secondary growth originating from their lateral or secondary meristems: the vascular cambium and the cork cambium or phellogen; the vascular cambium forms between the xylem and phloem in the vascular bundles and connects to form a continuous cylinder. The vascular cambium cells divide to produce secondary xylem to the inside and secondary phloem to the outside; as the stem increases in diameter due to production of secondary xylem and secondary phloem, the cortex and epidermis are destroyed. Before the cortex is destroyed, a cork cambium develops there; the cork cambium divides to produce waterproof cork cells externally and sometimes phelloderm cells internally. Those three tissues form the periderm.
Areas of loosely pack
A leaf is an organ of a vascular plant and is the principal lateral appendage of the stem. The leaves and stem together form the shoot. Leaves are collectively referred to as foliage, as in "autumn foliage". A leaf is a thin, dorsiventrally flattened organ borne above ground and specialized for photosynthesis. In most leaves, the primary photosynthetic tissue, the palisade mesophyll, is located on the upper side of the blade or lamina of the leaf but in some species, including the mature foliage of Eucalyptus, palisade mesophyll is present on both sides and the leaves are said to be isobilateral. Most leaves have distinct upper surface and lower surface that differ in colour, the number of stomata, the amount and structure of epicuticular wax and other features. Leaves can have many different shapes and textures; the broad, flat leaves with complex venation of flowering plants are known as megaphylls and the species that bear them, the majority, as broad-leaved or megaphyllous plants. In the clubmosses, with different evolutionary origins, the leaves are simple and are known as microphylls.
Some leaves, such as bulb scales, are not above ground. In many aquatic species the leaves are submerged in water. Succulent plants have thick juicy leaves, but some leaves are without major photosynthetic function and may be dead at maturity, as in some cataphylls and spines. Furthermore, several kinds of leaf-like structures found in vascular plants are not homologous with them. Examples include flattened plant stems called phylloclades and cladodes, flattened leaf stems called phyllodes which differ from leaves both in their structure and origin; some structures of non-vascular plants function much like leaves. Examples include the phyllids of liverworts. Leaves are the most important organs of most vascular plants. Green plants are autotrophic, meaning that they do not obtain food from other living things but instead create their own food by photosynthesis, they capture the energy in sunlight and use it to make simple sugars, such as glucose and sucrose, from carbon dioxide and water. The sugars are stored as starch, further processed by chemical synthesis into more complex organic molecules such as proteins or cellulose, the basic structural material in plant cell walls, or metabolised by cellular respiration to provide chemical energy to run cellular processes.
The leaves draw water from the ground in the transpiration stream through a vascular conducting system known as xylem and obtain carbon dioxide from the atmosphere by diffusion through openings called stomata in the outer covering layer of the leaf, while leaves are orientated to maximise their exposure to sunlight. Once sugar has been synthesized, it needs to be transported to areas of active growth such as the plant shoots and roots. Vascular plants transport sucrose in a special tissue called the phloem; the phloem and xylem are parallel to each other but the transport of materials is in opposite directions. Within the leaf these vascular systems branch to form veins which supply as much of the leaf as possible, ensuring that cells carrying out photosynthesis are close to the transportation system. Leaves are broad and thin, thereby maximising the surface area directly exposed to light and enabling the light to penetrate the tissues and reach the chloroplasts, thus promoting photosynthesis.
They are arranged on the plant so as to expose their surfaces to light as efficiently as possible without shading each other, but there are many exceptions and complications. For instance plants adapted to windy conditions may have pendent leaves, such as in many willows and eucalyptss; the flat, or laminar, shape maximises thermal contact with the surrounding air, promoting cooling. Functionally, in addition to carrying out photosynthesis, the leaf is the principal site of transpiration, providing the energy required to draw the transpiration stream up from the roots, guttation. Many gymnosperms have thin needle-like or scale-like leaves that can be advantageous in cold climates with frequent snow and frost; these are interpreted as reduced from megaphyllous leaves of their Devonian ancestors. Some leaf forms are adapted to modulate the amount of light they absorb to avoid or mitigate excessive heat, ultraviolet damage, or desiccation, or to sacrifice light-absorption efficiency in favour of protection from herbivory.
For xerophytes the major constraint drought. Some window plants such as Fenestraria species and some Haworthia species such as Haworthia tesselata and Haworthia truncata are examples of xerophytes. and Bulbine mesembryanthemoides. Leaves function to store chemical energy and water and may become specialised organs serving other functions, such as tendrils of peas and other legumes, the protective spines of cacti and the insect traps in carnivorous plants such as Nepenthes and Sarracenia. Leaves are the fundamental structural units from which cones are constructed in gymnosperms and from which flowers are constructed in flowering plants; the internal organisation of most kinds of leaves has evolved to maximise exposure of the photosynthetic organelles, the chloroplasts, to light and to increase the absorption of carbon dioxide while at the same time controlling water loss. Their surfaces are waterproofed by the plant cuticle and gas exchange between the mesophyll cells and the atmosphere is controlled by minute openings called stomata which open or close to regulate the rate exchange of carbon dioxide and water vapour into
Cyanobacteria known as Cyanophyta, are a phylum of bacteria that obtain their energy through photosynthesis and are the only photosynthetic prokaryotes able to produce oxygen. The name cyanobacteria comes from the color of the bacteria. Cyanobacteria, which are prokaryotes, are called "blue-green algae", though the term algae in modern usage is restricted to eukaryotes. Unlike heterotrophic prokaryotes, cyanobacteria have internal membranes; these are flattened. Phototrophic eukaryotes perform photosynthesis by plastids that may have their ancestry in cyanobacteria, acquired long ago via a process called endosymbiosis; these endosymbiotic cyanobacteria in eukaryotes may have evolved or differentiated into specialized organelles such as chloroplasts and leucoplasts. By producing and releasing oxygen, cyanobacteria are thought to have converted the early oxygen-poor, reducing atmosphere into an oxidizing one, causing the Great Oxygenation Event and the "rusting of the Earth", which changed the composition of the Earth's life forms and led to the near-extinction of anaerobic organisms.
Cyanobacteria are a group of photosynthetic bacteria, some of which are nitrogen-fixing, that live in a wide variety of moist soils and water either or in a symbiotic relationship with plants or lichen-forming fungi. They include colonial species. Colonies may form filaments, sheets, or hollow spheres; some filamentous species can differentiate into several different cell types: vegetative cells – the normal, photosynthetic cells that are formed under favorable growing conditions. Some cyanobacteria can fix atmospheric nitrogen in anaerobic conditions by means of specialized cells called heterocysts. Heterocysts may form under the appropriate environmental conditions when fixed nitrogen is scarce. Heterocyst-forming species are specialized for nitrogen fixation and are able to fix nitrogen gas into ammonia, nitrites or nitrates, which can be absorbed by plants and converted to protein and nucleic acids. Free-living cyanobacteria are present in the water of rice paddies, cyanobacteria can be found growing as epiphytes on the surfaces of the green alga, where they may fix nitrogen.
Cyanobacteria such as Anabaena can provide rice plantations with biofertilizer. Many cyanobacteria form motile filaments of cells, called hormogonia, that travel away from the main biomass to bud and form new colonies elsewhere; the cells in a hormogonium are thinner than in the vegetative state, the cells on either end of the motile chain may be tapered. To break away from the parent colony, a hormogonium must tear apart a weaker cell in a filament, called a necridium; each individual cell has a thick, gelatinous cell wall. They lack flagella. Many of the multicellular filamentous. In water columns, some cyanobacteria float by forming gas vesicles, as in archaea; these vesicles are not organelles as such. They are not bounded by a protein sheath. Cyanobacteria can be found in every terrestrial and aquatic habitat—oceans, fresh water, damp soil, temporarily moistened rocks in deserts, bare rock and soil, Antarctic rocks, they can form phototrophic biofilms. They are found in endolithic ecosystem. A few are endosymbionts in lichens, various protists, or sponges and provide energy for the host.
Some live in the fur of sloths. Aquatic cyanobacteria are known for their extensive and visible blooms that can form in both freshwater and marine environments; the blooms can have the appearance of blue-green scum. These blooms can be toxic, lead to the closure of recreational waters when spotted. Marine bacteriophages are significant parasites of unicellular marine cyanobacteria. Cyanobacteria growth is favored in ponds and lakes where waters are calm and have less turbulent mixing, their life cycles are disrupted when the water or artificially mixes from churning currents caused by the flowing water of streams or the churning water of fountains. For this reason blooms of cyanobacteria occur in rivers unless the water is flowing slowly. Growth is favored at higher temperatures, making increasing water temperature as a result of global warming more problematic. At higher temperatures Microcystis species are able to outcompete green algae; this is a concern because of the production of toxins produced by Microcystis.
Based on environmental trends and observations suggest cyanobacteria will increase their dominance in aquatic environments. This can lead to serious consequences the contamination of sources of drinking water. Cyanobacteria can interfere with water treatment in various ways by plugging filters and by producing cyanotoxins, which have the potential to cause serious illness if consumed. Consequences may lie within
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 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 Carboniferous is a geologic period and system that spans 60 million years from the end of the Devonian Period 358.9 million years ago, to the beginning of the Permian Period, 298.9 Mya. The name Carboniferous means "coal-bearing" and derives from the Latin words carbō and ferō, was coined by geologists William Conybeare and William Phillips in 1822. Based on a study of the British rock succession, it was the first of the modern'system' names to be employed, reflects the fact that many coal beds were formed globally during that time; the Carboniferous is treated in North America as two geological periods, the earlier Mississippian and the Pennsylvanian. Terrestrial animal life was well established by the Carboniferous period. Amphibians were the dominant land vertebrates, of which one branch would evolve into amniotes, the first terrestrial vertebrates. Arthropods were very common, many were much larger than those of today. Vast swaths of forest covered the land, which would be laid down and become the coal beds characteristic of the Carboniferous stratigraphy evident today.
The atmospheric content of oxygen reached its highest levels in geological history during the period, 35% compared with 21% today, allowing terrestrial invertebrates to evolve to great size. The half of the period experienced glaciations, low sea level, mountain building as the continents collided to form Pangaea. A minor marine and terrestrial extinction event, the Carboniferous rainforest collapse, occurred at the end of the period, caused by climate change. In the United States the Carboniferous is broken into Mississippian and Pennsylvanian subperiods; the Mississippian is about twice as long as the Pennsylvanian, but due to the large thickness of coal-bearing deposits with Pennsylvanian ages in Europe and North America, the two subperiods were long thought to have been more or less equal in duration. In Europe the Lower Carboniferous sub-system is known as the Dinantian, comprising the Tournaisian and Visean Series, dated at 362.5-332.9 Ma, the Upper Carboniferous sub-system is known as the Silesian, comprising the Namurian and Stephanian Series, dated at 332.9-298.9 Ma.
The Silesian is contemporaneous with the late Mississippian Serpukhovian plus the Pennsylvanian. In Britain the Dinantian is traditionally known as the Carboniferous Limestone, the Namurian as the Millstone Grit, the Westphalian as the Coal Measures and Pennant Sandstone; the International Commission on Stratigraphy faunal stages from youngest to oldest, together with some of their regional subdivisions, are: A global drop in sea level at the end of the Devonian reversed early in the Carboniferous. There was a drop in south polar temperatures; these conditions had little effect in the deep tropics, where lush swamps to become coal, flourished to within 30 degrees of the northernmost glaciers. Mid-Carboniferous, a drop in sea level precipitated a major marine extinction, one that hit crinoids and ammonites hard; this sea level drop and the associated unconformity in North America separate the Mississippian subperiod from the Pennsylvanian subperiod. This happened about 323 million years ago, at the onset of the Permo-Carboniferous Glaciation.
The Carboniferous was a time of active mountain-building as the supercontinent Pangaea came together. The southern continents remained tied together in the supercontinent Gondwana, which collided with North America–Europe along the present line of eastern North America; this continental collision resulted in the Hercynian orogeny in Europe, the Alleghenian orogeny in North America. In the same time frame, much of present eastern Eurasian plate welded itself to Europe along the line of the Ural Mountains. Most of the Mesozoic supercontinent of Pangea was now assembled, although North China, South China continents were still separated from Laurasia; the Late Carboniferous Pangaea was shaped like an "O." There were two major oceans in the Carboniferous—Panthalassa and Paleo-Tethys, inside the "O" in the Carboniferous Pangaea. Other minor oceans were shrinking and closed - Rheic Ocean, the small, shallow Ural Ocean and Proto-Tethys Ocean. Average global temperatures in the Early Carboniferous Period were high: 20 °C.
However, cooling during the Middle Carboniferous reduced average global temperatures to about 12 °C. Lack of growth rings of fossilized trees suggest a lack of seasons of a tropical climate. Glaciations in Gondwana, triggered by Gondwana's southward movement, continued into the Permian and because of the lack of clear markers and breaks, the deposits of this glacial period are referred to as Permo-Carboniferous in age; the cooling and drying of the climate led to the Carboniferous Rainforest Collapse during the late Carboniferous. Tropical rainforests fragmented and were devastated by climate change. Carboniferous rocks in Europe and eastern North America consist of a repeated sequence of limestone, sandstone and coal beds. In North America, the early Carboniferous is marine
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