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108-545: The Protolepidodendrales are an extinct order of lycopsids that flourished from the Devonian to the lower Carboniferous ( Mississippian ) periods. Leclercqia is one of the best-known genera. Protolepidodendropsis is another genus included in this order. This article about a prehistoric lycophyte is a stub . You can help Misplaced Pages by expanding it . Devonian The Devonian ( / d ə ˈ v oʊ n i . ən , d ɛ -/ də- VOH -nee-ən, deh- )

216-749: A Darwin Point is reached. One such point exists at the northwestern end of the Hawaiian Archipelago; see Evolution of Hawaiian volcanoes#Coral atoll stage . However, reefs in the Persian Gulf have adapted to temperatures of 13 °C (55 °F) in winter and 38 °C (100 °F) in summer. 37 species of scleractinian corals inhabit such an environment around Larak Island . Deep-water coral inhabits greater depths and colder temperatures at much higher latitudes, as far north as Norway. Although deep water corals can form reefs, little

324-644: A continental shelf, and not around a volcanic island, Darwin's principles apply. Development stopped at the barrier reef stage, since Australia is not about to submerge. It formed the world's largest barrier reef, 300–1,000 m (980–3,280 ft) from shore, stretching for 2,000 km (1,200 mi). Healthy tropical coral reefs grow horizontally from 1 to 3 cm (0.39 to 1.18 in) per year, and grow vertically anywhere from 1 to 25 cm (0.39 to 9.84 in) per year; however, they grow only at depths shallower than 150 m (490 ft) because of their need for sunlight, and cannot grow above sea level. As

432-417: A few species of the order Proetida . The subsequent end-Devonian extinction , which occurred at around 359 Ma, further impacted the ecosystems and completed the extinction of all calcite sponge reefs and placoderms. Devonian palaeogeography was dominated by the supercontinent Gondwana to the south, the small continent of Siberia to the north, and the medium-sized continent of Laurussia to

540-763: A low-latitude archipelago to the north of Gondwana. They were separated from the southern continent by an oceanic basin: the Paleo-Tethys . Although the western Paleo-Tethys Ocean had existed since the Cambrian, the eastern part only began to rift apart as late as the Silurian. This process accelerated in the Devonian. The eastern branch of the Paleo-Tethys was fully opened when South China and Annamia (a terrane equivalent to most of Indochina ), together as

648-529: A massive extinction event . ( See Late Devonian extinction ). Primitive arthropods co-evolved with this diversified terrestrial vegetation structure. The evolving co-dependence of insects and seed plants that characterized a recognizably modern world had its genesis in the Late Devonian Epoch. The development of soils and plant root systems probably led to changes in the speed and pattern of erosion and sediment deposition. The rapid evolution of

756-412: A pinhead to 12 inches (30 cm) across. Reef-building or hermatypic corals live only in the photic zone (above 70 m), the depth to which sufficient sunlight penetrates the water. Coral polyps do not photosynthesize, but have a symbiotic relationship with microscopic algae ( dinoflagellates ) of the genus Symbiodinium , commonly referred to as zooxanthellae . These organisms live within

864-496: A pseudo-atoll. These can be distinguished from real atolls only by detailed investigation, possibly including core drilling. Some platform reefs of the Laccadives are U-shaped, due to wind and water flow. Atolls or atoll reefs are a more or less circular or continuous barrier reef that extends all the way around a lagoon without a central island. They are usually formed from fringing reefs around volcanic islands. Over time,

972-420: A range of evidence, such as plant distribution, points to a Late Devonian warming. The climate would have affected the dominant organisms in reefs ; microbes would have been the main reef-forming organisms in warm periods, with corals and stromatoporoid sponges taking the dominant role in cooler times. The warming at the end of the Devonian may even have contributed to the extinction of the stromatoporoids. At

1080-433: A rapidly subsiding bottom can overwhelm coral growth, killing the coral and the reef, due to what is called coral drowning . Corals that rely on zooxanthellae can die when the water becomes too deep for their symbionts to adequately photosynthesize , due to decreased light exposure. The two main variables determining the geomorphology , or shape, of coral reefs are the nature of the substrate on which they rest, and

1188-664: A terrestrial ecosystem that contained copious animals opened the way for the first vertebrates to seek terrestrial living. By the end of the Devonian, arthropods were solidly established on the land. The Late Devonian extinction is not a single event, but rather is a series of pulsed extinctions at the Givetian-Frasnian boundary, the Frasnian-Famennian boundary, and the Devonian-Carboniferous boundary. Together, these are considered one of

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1296-541: A unified continent, detached from the northeastern sector of Gondwana. Nevertheless, they remained close enough to Gondwana that their Devonian fossils were more closely related to Australian species than to north Asian species. Other Asian terranes remained attached to Gondwana, including Sibumasu (western Indochina), Tibet, and the rest of the Cimmerian blocks. While the South China-Annamia continent

1404-602: A volcanically active region during the Devonian, as it continued to assimilate smaller island arcs. The island arcs of the region, such as the Balkhash-West Junggar Arc, exhibited biological endemism as a consequence of their location. Siberia was located just north of the equator as the largest landmass in the Northern Hemisphere. At the beginning of the Devonian, Siberia was inverted (upside down) relative to its modern orientation. Later in

1512-622: 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, Middle and Late subdivisions. The rocks corresponding to those epochs are referred to as belonging to the Lower, Middle and Upper parts of the Devonian System. The Early Devonian lasted from 419.2 to 393.3 Ma. It began with

1620-423: A zone extending from approximately 30° N to 30° S of the equator. Tropical corals do not grow at depths of over 50 meters (160 ft). The optimum temperature for most coral reefs is 26–27 °C (79–81 °F), and few reefs exist in waters below 18 °C (64 °F). When the net production by reef building corals no longer keeps pace with relative sea level and the reef structure permanently drowns

1728-762: Is a geologic period and system of the Paleozoic era during the Phanerozoic eon , spanning 60.3 million years from the end of the preceding Silurian period at 419.2 million years ago ( Ma ), to the beginning of the succeeding Carboniferous period at 358.9 Ma. It is the fourth period of both the Paleozoic and the Phanerozoic. It is named after Devon , South West England , where rocks from this period were first studied. The first significant evolutionary radiation of life on land occurred during

1836-578: Is constantly changing. Each reef is made up of irregular patches of algae, sessile invertebrates, and bare rock and sand. The size, shape and relative abundance of these patches change from year to year in response to the various factors that favor one type of patch over another. Growing coral, for example, produces constant change in the fine structure of reefs. On a larger scale, tropical storms may knock out large sections of reef and cause boulders on sandy areas to move. Coral reefs are estimated to cover 284,300 km (109,800 sq mi), just under 0.1% of

1944-445: Is directly attached to a shore, or borders it with an intervening narrow, shallow channel or lagoon. It is the most common reef type. Fringing reefs follow coastlines and can extend for many kilometres. They are usually less than 100 metres wide, but some are hundreds of metres wide. Fringing reefs are initially formed on the shore at the low water level and expand seawards as they grow in size. The final width depends on where

2052-578: Is good evidence that Rheic oceanic crust experienced intense subduction and metamorphism under Mexico and Central America. The closure of the eastern part of the Rheic Ocean is associated with the assemblage of central and southern Europe. In the early Paleozoic, much of Europe was still attached to Gondwana, including the terranes of Iberia , Armorica (France), Palaeo-Adria (the western Mediterranean area), Bohemia , Franconia , and Saxothuringia . These continental blocks, collectively known as

2160-523: Is known about them. The northernmost coral reef on Earth is located near Eilat , Israel . Coral reefs are rare along the west coasts of the Americas and Africa , due primarily to upwelling and strong cold coastal currents that reduce water temperatures in these areas (the Humboldt , Benguela , and Canary Currents , respectively). Corals are seldom found along the coastline of South Asia —from

2268-430: Is observed in many of those plants. Some of the early land plants such as Drepanophycus likely spread by vegetative growth and spores. The earliest land plants such as Cooksonia consisted of leafless, dichotomous axes with terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall. Fossils of Armoricaphyton chateaupannense , about 400 million years old, represent

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2376-418: Is often the reef's most diverse area. Coral and calcareous algae provide complex habitats and areas that offer protection, such as cracks and crevices. Invertebrates and epiphytic algae provide much of the food for other organisms. A common feature on this forereef zone is spur and groove formations that serve to transport sediment downslope. The reef flat is the sandy-bottomed flat, which can be behind

2484-641: The Antler orogeny , which extended into the Carboniferous. Mountain building could also be found in the far northeastern extent of the continent, as minor tropical island arcs and detached Baltic terranes re-join the continent. Deformed remnants of these mountains can still be found on Ellesmere Island and Svalbard . Many of the Devonian collisions in Laurussia produce both mountain chains and foreland basins , which are frequently fossiliferous. Gondwana

2592-1437: The Cambrian . Sometimes called rainforests of the sea , shallow coral reefs form some of Earth's most diverse ecosystems. They occupy less than 0.1% of the world's ocean area, about half the area of France, yet they provide a home for at least 25% of all marine species , including fish , mollusks , worms , crustaceans , echinoderms , sponges , tunicates and other cnidarians . Coral reefs flourish in ocean waters that provide few nutrients. They are most commonly found at shallow depths in tropical waters, but deep water and cold water coral reefs exist on smaller scales in other areas. Shallow tropical coral reefs have declined by 50% since 1950, partly because they are sensitive to water conditions. They are under threat from excess nutrients (nitrogen and phosphorus), rising ocean heat content and acidification , overfishing (e.g., from blast fishing , cyanide fishing , spearfishing on scuba ), sunscreen use, and harmful land-use practices, including runoff and seeps (e.g., from injection wells and cesspools). Coral reefs deliver ecosystem services for tourism, fisheries and shoreline protection . The annual global economic value of coral reefs has been estimated at anywhere from US$ 30–375 billion (1997 and 2003 estimates) to US$ 2.7 trillion (a 2020 estimate) to US$ 9.9 trillion (a 2014 estimate). Though

2700-565: The Frasnian , 382.7 to 372.2 Ma, during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing Famennian subdivision, the beginning and end of which are marked with extinction events. This lasted until the end of the Devonian, 358.9 Ma. The Devonian was a relatively warm period, although significant glaciers may have existed during the Early and Middle Devonian. The temperature gradient from

2808-483: The Late Carboniferous . The first ammonites , a subclass of cephalopod molluscs , appeared. Trilobites , brachiopods and the great coral reefs were still common during the Devonian. The Late Devonian extinction , which started about 375 Ma, severely affected marine life, killing off most of the reef systems, most of the jawless fish, half of all placoderms, and nearly all trilobites save for

2916-475: The Late Cretaceous (100–66 Ma), when reefs formed by a group of bivalves called rudists existed; one of the valves formed the main conical structure and the other, much smaller valve acted as a cap. Measurements of the oxygen isotopic composition of the aragonitic skeleton of coral reefs, such as Porites , can indicate changes in sea surface temperature and sea surface salinity conditions during

3024-640: The Lochkovian Stage 419.2 to 410.8 Ma, which was followed by the Pragian from 410.8 to 407.6 Ma and then by the Emsian , which lasted until the Middle Devonian began, 393.3 Ma. During this time, the first ammonoids appeared, descending from bactritoid nautiloids . Ammonoids during this time period were simple and differed little from their nautiloid counterparts. These ammonoids belong to

3132-739: The Mesoamerican Barrier Reef System and the New Caledonian Barrier Reef . Barrier reefs are also found on the coasts of Providencia , Mayotte , the Gambier Islands , on the southeast coast of Kalimantan , on parts of the coast of Sulawesi , southeastern New Guinea and the south coast of the Louisiade Archipelago . Platform reefs, variously called bank or table reefs, can form on the continental shelf , as well as in

3240-592: The Middle Cambrian (513–501 Ma ), Devonian (416–359 Ma) and Carboniferous (359–299 Ma), owing to extinct order Rugosa corals, and Late Cretaceous (100–66 Ma) and Neogene (23 Ma–present), owing to order Scleractinia corals. Not all reefs in the past were formed by corals: those in the Early Cambrian (542–513 Ma) resulted from calcareous algae and archaeocyathids (small animals with conical shape, probably related to sponges ) and in

3348-475: The cladoxylopsids and progymnosperm Archaeopteris . These tracheophytes were able to grow to large size on dry land because they had evolved the ability to biosynthesize lignin , which gave them physical rigidity and improved the effectiveness of their vascular system while giving them resistance to pathogens and herbivores. In Eifelian age, cladoxylopsid trees formed the first forests in Earth history. By

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3456-480: The class Anthozoa in the animal phylum Cnidaria , which includes sea anemones and jellyfish . Unlike sea anemones, corals secrete hard carbonate exoskeletons that support and protect the coral. Most reefs grow best in warm, shallow, clear, sunny and agitated water. Coral reefs first appeared 485 million years ago, at the dawn of the Early Ordovician , displacing the microbial and sponge reefs of

3564-417: The tropics where the water is agitated, but can only live within a limited depth range, starting just below low tide . Where the level of the underlying earth allows, the corals grow around the coast to form fringing reefs, and can eventually grow to become a barrier reef. Where the bottom is rising, fringing reefs can grow around the coast, but coral raised above sea level dies. If the land subsides slowly,

3672-452: The " Big Five " mass extinctions in Earth's history. The Devonian extinction crisis primarily affected the marine community, and selectively affected shallow warm-water organisms rather than cool-water organisms. The most important group to be affected by this extinction event were the reef-builders of the great Devonian reef systems. Amongst the severely affected marine groups were the brachiopods, trilobites, ammonites, and acritarchs , and

3780-710: The Armorican Terrane Assemblage, split away from Gondwana in the Silurian and drifted towards Laurussia through the Devonian. Their collision with Laurussia leads to the beginning of the Variscan Orogeny , a major mountain-building event which would escalate further in the Late Paleozoic. Franconia and Saxothuringia collided with Laurussia near the end of the Early Devonian, pinching out the easternmost Rheic Ocean. The rest of

3888-536: The Armorican terranes followed, and by the end of the Devonian they were fully connected with Laurussia. This sequence of rifting and collision events led to the successive creation and destruction of several small seaways, including the Rheno-Hercynian, Saxo-Thuringian, and Galicia-Moldanubian oceans. Their sediments were eventually compressed and completely buried as Gondwana fully collided with Laurussia in

3996-511: The Carboniferous. Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by conodonts, bryozoans , diverse and abundant brachiopods , the enigmatic hederellids , microconchids , and corals . Lily-like crinoids (animals, their resemblance to flowers notwithstanding) were abundant, and trilobites were still fairly common. Bivalves became commonplace in deep water and outer shelf environments. The first ammonites also appeared during or slightly before

4104-533: The Devonian Period. The newly evolved forests drew carbon out of the atmosphere, which were then buried into sediments. This may be reflected by a Mid-Devonian cooling of around 5 °C (9 °F). The Late Devonian warmed to levels equivalent to the Early Devonian; while there is no corresponding increase in CO 2 concentrations, continental weathering increases (as predicted by warmer temperatures); further,

4212-577: The Devonian extinction events were caused by an asteroid impact. However, while there were Late Devonian collision events (see the Alamo bolide impact ), little evidence supports the existence of a large enough Devonian crater. Coral reef A coral reef is an underwater ecosystem characterized by reef-building corals . Reefs are formed of colonies of coral polyps held together by calcium carbonate . Most coral reefs are built from stony corals , whose polyps cluster in groups. Coral belongs to

4320-408: The Devonian seas. The first abundant genus of cartilaginous fish, Cladoselache , appeared in the oceans during the Devonian Period. The great diversity of fish around at the time has led to the Devonian being given the name "The Age of Fishes" in popular culture. The Devonian saw significant expansion in the diversity of nektonic marine life driven by the abundance of planktonic microorganisms in

4428-645: The Devonian to often be dubbed the Age of Fishes . The armored placoderms began dominating almost every known aquatic environment. In the oceans, cartilaginous fishes such as primitive sharks became more numerous than in the Silurian and Late Ordovician . Tetrapodomorphs , which include the ancestors of all four- limbed vertebrates (i.e. tetrapods ), began diverging from freshwater lobe-finned fish as their more robust and muscled pectoral and pelvic fins gradually evolved into forelimbs and hindlimbs , though they were not fully established for life on land until

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4536-434: The Devonian, as free- sporing land plants ( pteridophytes ) began to spread across dry land , forming extensive coal forests which covered the continents. By the middle of the Devonian, several groups of vascular plants had evolved leaves and true roots , and by the end of the period the first seed -bearing plants ( pteridospermatophytes ) appeared. This rapid evolution and colonization process, which had begun during

4644-451: The Early Devonian and the existence of fossils such as Protichnites suggest that amphibious arthropods may have appeared as early as the Cambrian ). By far the largest land organism at the beginning of this period was the enigmatic Prototaxites , which was possibly the fruiting body of an enormous fungus, rolled liverwort mat, or another organism of uncertain affinities that stood more than 8 metres (26 ft) tall, and towered over

4752-471: The Early and Middle Devonian, the west coast of Laurussia was a passive margin with broad coastal waters, deep silty embayments, river deltas and estuaries, found today in Idaho and Nevada . In the Late Devonian, an approaching volcanic island arc reached the steep slope of the continental shelf and began to uplift deep water deposits. This minor collision sparked the start of a mountain-building episode called

4860-738: The Indian Ocean, for example, in the Maldives , the Chagos Islands , the Seychelles and around Cocos Island . The entire Maldives consist of 26 atolls. Coral reef ecosystems contain distinct zones that host different kinds of habitats. Usually, three major zones are recognized: the fore reef, reef crest, and the back reef (frequently referred to as the reef lagoon). The three zones are physically and ecologically interconnected. Reef life and oceanic processes create opportunities for

4968-637: The Late Devonian the tectonic situation had relaxed and much of South America was covered by shallow seas. These south polar seas hosted a distinctive brachiopod fauna, the Malvinokaffric Realm, which extended eastward to marginal areas now equivalent to South Africa and Antarctica. Malvinokaffric faunas even managed to approach the South Pole via a tongue of Panthalassa which extended into the Paraná Basin . The northern rim of Gondwana

5076-682: The Late Devonian. The Altai-Sayan region was shaken by volcanism in the Early and Middle Devonian, while Late Devonian magmatism was magnified further to produce the Vilyuy Traps , flood basalts which may have contributed to the Late Devonian Mass Extinction. The last major round of volcanism, the Yakutsk Large Igneous Province, continued into the Carboniferous to produce extensive kimberlite deposits. Similar volcanic activity also affected

5184-640: The Silurian and Devonian, it decreased across the span of the Devonian, particularly during the Pragian, and that the overall diversity of nektonic taxa did not increase significantly during the Devonian compared to during other geologic periods, and was in fact higher during the intervals spanning from the Wenlock to the Lochkovian and from the Carboniferous to the Permian. The study's authors instead attribute

5292-560: The Silurian, is known as the Silurian-Devonian Terrestrial Revolution . The earliest land animals , predominantly arthropods such as myriapods , arachnids and hexapods , also became well-established early in this period, after beginning their colonization of land at least from the Ordovician period. Fishes , especially jawed fish , reached substantial diversity during this time, leading

5400-599: The UV radiation and allowing them to better adapt to warmer water temperatures. In the event of UV or thermal damage, if and when repair occurs, it will increase the likelihood of survival of the host and symbiont. This leads to the idea that, evolutionarily, clade A is more UV resistant and thermally resistant than the other clades. Clades B and C are found more frequently in deeper water, which may explain their higher vulnerability to increased temperatures. Terrestrial plants that receive less sunlight because they are found in

5508-661: The beginning of the Mississippian subperiod of the Carboniferous. In 19th-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

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5616-502: The coast. The fringing reefs of the Red Sea are "some of the best developed in the world" and occur along all its shores except off sandy bays. Barrier reefs are separated from a mainland or island shore by a deep channel or lagoon . They resemble the later stages of a fringing reef with its lagoon but differ from the latter mainly in size and origin. Their lagoons can be several kilometres wide and 30 to 70 metres deep. Above all,

5724-485: The coastal plains had become continental islands . As sea level rise continued, water topped most of the continental islands. The corals could then overgrow the hills, forming cays and reefs. Sea level on the Great Barrier Reef has not changed significantly in the last 6,000 years. The age of living reef structure is estimated to be between 6,000 and 8,000 years. Although the Great Barrier Reef formed along

5832-684: The continent (such as Greenland and Ellesmere Island ) established tropical conditions, most of the continent was located within the natural dry zone along the Tropic of Capricorn , which (as nowadays) is a result of the convergence of two great air-masses, the Hadley cell and the Ferrel cell . In these near-deserts, the Old Red Sandstone sedimentary beds formed, made red by the oxidised iron ( hematite ) characteristic of drought conditions. The abundance of red sandstone on continental land also lends Laurussia

5940-548: The continents Laurentia (modern day North America) and Baltica (modern day northern and eastern Europe). The tectonic effects of this collision continued into the Devonian, producing a string of mountain ranges along the southeastern coast of the continent. In present-day eastern North America, the Acadian Orogeny continued to raise the Appalachian Mountains . Further east, the collision also extended

6048-451: The coral animals. Coral that loses a large fraction of its zooxanthellae becomes white (or sometimes pastel shades in corals that are pigmented with their own proteins) and is said to be bleached , a condition which, unless corrected, can kill the coral. There are eight clades of Symbiodinium phylotypes . Most research has been conducted on clades A–D. Each clade contributes their own benefits as well as less compatible attributes to

6156-415: The corals shelter the zooxanthellae, averaging one million for every cubic centimetre of coral, and provide a constant supply of the carbon dioxide they need for photosynthesis. The varying pigments in different species of zooxanthellae give them an overall brown or golden-brown appearance and give brown corals their colors. Other pigments such as reds, blues, greens, etc. come from colored proteins made by

6264-529: The deep sea away from continental shelves , around oceanic islands and atolls . The majority of these islands are volcanic in origin. Others have tectonic origins where plate movements lifted the deep ocean floor. In The Structure and Distribution of Coral Reefs , Charles Darwin set out his theory of the formation of atoll reefs, an idea he conceived during the voyage of the Beagle . He theorized that uplift and subsidence of Earth's crust under

6372-405: The early Devonian Period around 400  Ma. Bactritoids make their first appearance in the Early Devonian as well; their radiation, along with that of ammonoids, has been attributed by some authors to increased environmental stress resulting from decreasing oxygen levels in the deeper parts of the water column. Among vertebrates, jawless armored fish ( ostracoderms ) declined in diversity, while

6480-662: The east. Major tectonic events include the closure of the Rheic Ocean , the separation of South China from Gondwana, and the resulting expansion of the Paleo-Tethys Ocean . The Devonian experienced several major mountain-building events as Laurussia and Gondwana approached; these include the Acadian Orogeny in North America and the beginning of the Variscan Orogeny in Europe. These early collisions preceded

6588-637: The eastern tip of India ( Chennai ) to the Bangladesh and Myanmar borders —as well as along the coasts of northeastern South America and Bangladesh, due to the freshwater release from the Amazon and Ganges Rivers respectively. Significant coral reefs include: When alive, corals are colonies of small animals embedded in calcium carbonate shells. Coral heads consist of accumulations of individual animals called polyps , arranged in diverse shapes. Polyps are usually tiny, but they can range in size from

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6696-569: The end of the Devonian, the first seed-forming plants had appeared. This rapid appearance of many plant groups and growth forms has been referred to as the Devonian Explosion or the Silurian-Devonian Terrestrial Revolution. The 'greening' of the continents acted as a carbon sink , and atmospheric concentrations of carbon dioxide may have dropped. This may have cooled the climate and led to

6804-412: The equator to the poles was not as large as it is today. The weather was also very arid, mostly along the equator where it was the driest. Reconstruction of tropical sea surface temperature from conodont apatite implies an average value of 30 °C (86 °F) in the Early Devonian. Early Devonian mean annual surface temperatures were approximately 16 °C. CO 2 levels dropped steeply throughout

6912-479: The exchange of seawater , sediments , nutrients and marine life. Most coral reefs exist in waters less than 50 m deep. Some inhabit tropical continental shelves where cool, nutrient-rich upwelling does not occur, such as the Great Barrier Reef . Others are found in the deep ocean surrounding islands or as atolls, such as in the Maldives . The reefs surrounding islands form when islands subside into

7020-401: The floor often supports seagrass meadows which are important foraging areas for reef fish. The reef drop-off is, for its first 50 m, habitat for reef fish who find shelter on the cliff face and plankton in the water nearby. The drop-off zone applies mainly to the reefs surrounding oceanic islands and atolls. The reef face is the zone above the reef floor or the reef drop-off. This zone

7128-459: The formation of the single supercontinent Pangaea in the Late Paleozoic. 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 throughout the county was resolved by adding the Devonian Period to the geological timescale. The Great Devonian Controversy was a lengthy debate between Roderick Murchison , Adam Sedgwick and Henry De la Beche over

7236-492: The free water column as well as high ecological competition in benthic habitats, which were extremely saturated; this diversification has been labeled the Devonian Nekton Revolution by many researchers. However, other researchers have questioned whether this revolution existed at all; a 2018 study found that although the proportion of biodiversity constituted by nekton increased across the boundary between

7344-403: The fringing reef, barrier reef, atoll and table reef based on Stoddart, D.R. (1969). Spalding et al. list four main reef types that can be clearly illustrated – the fringing reef, barrier reef, atoll, and "bank or platform reef"—and notes that many other structures exist which do not conform easily to strict definitions, including the "patch reef". A fringing reef, also called a shore reef,

7452-463: The fringing reefs keep pace by growing upwards on a base of older, dead coral, forming a barrier reef enclosing a lagoon between the reef and the land. A barrier reef can encircle an island, and once the island sinks below sea level a roughly circular atoll of growing coral continues to keep up with the sea level, forming a central lagoon. Barrier reefs and atolls do not usually form complete circles but are broken in places by storms. Like sea level rise ,

7560-419: The growth of the coral. This technique is often used by climate scientists to infer a region's paleoclimate . Since Darwin's identification of the three classical reef formations – the fringing reef around a volcanic island becoming a barrier reef and then an atoll – scientists have identified further reef types. While some sources find only three, Thomas lists "Four major forms of large-scale coral reefs" –

7668-548: The history of the change in sea level relative to that substrate. The approximately 20,000-year-old Great Barrier Reef offers an example of how coral reefs formed on continental shelves. Sea level was then 120 m (390 ft) lower than in the 21st century. As sea level rose, the water and the corals encroached on what had been hills of the Australian coastal plain. By 13,000 years ago, sea level had risen to 60 m (200 ft) lower than at present, and many hills of

7776-744: The increased overall diversity of nekton in the Devonian to a broader, gradual trend of nektonic diversification across the entire Palaeozoic. A now-dry barrier reef, located in present-day Kimberley Basin of northwest Australia , once extended 350 km (220 mi), fringing a Devonian continent. Reefs are generally built by various carbonate -secreting organisms that can erect wave-resistant structures near sea level. Although modern reefs are constructed mainly by corals and calcareous algae , Devonian reefs were either microbial reefs built up mostly by autotrophic cyanobacteria or coral-stromatoporoid reefs built up by coral-like stromatoporoids and tabulate and rugose corals . Microbial reefs dominated under

7884-425: The increasing competition, predation, and diversity of jawed fishes . The shallow, warm, 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 and the ability to crawl out of the water and onto the land for short periods of time. Finally, the Late Devonian started with

7992-666: The island erodes away and sinks below sea level. Atolls may also be formed by the sinking of the seabed or rising of the sea level. A ring of reefs results, which enclose a lagoon. Atolls are numerous in the South Pacific, where they usually occur in mid-ocean, for example, in the Caroline Islands , the Cook Islands , French Polynesia , the Marshall Islands and Micronesia . Atolls are found in

8100-403: The jawed fish (gnathostomes) simultaneously increased in both the sea and fresh water . Armored placoderms were numerous during the early ages of the Devonian Period and became extinct in the Late Devonian, perhaps because of competition for food against the other fish species. Early cartilaginous ( Chondrichthyes ) and bony fishes ( Osteichthyes ) also become diverse and played a large role within

8208-481: The land lay under shallow seas, where tropical reef organisms lived. The enormous "world ocean", Panthalassa , occupied much of the Northern Hemisphere as well as wide swathes east of Gondwana and west of Laurussia. Other minor oceans were the Paleo-Tethys Ocean and Rheic Ocean . By the early Devonian, the continent Laurussia (also known as Euramerica ) was fully formed through the collision of

8316-575: The low, carpet-like vegetation during the early part of the Devonian. Also, the first possible fossils of insects appeared around 416  Ma, in the Early Devonian. Evidence for the earliest tetrapods takes the form of trace fossils in shallow lagoon environments within a marine carbonate platform/shelf during the Middle Devonian, although these traces have been questioned and an interpretation as fish feeding traces ( Piscichnus ) has been advanced. Many Early Devonian plants did not have true roots or leaves like extant plants, although vascular tissue

8424-418: The main reef, containing chunks of coral. This zone may border a lagoon and serve as a protective area, or it may lie between the reef and the shore, and in this case is a flat, rocky area. Fish tend to prefer it when it is present. The reef lagoon is an entirely enclosed region, which creates an area less affected by wave action and often contains small reef patches. However, the topography of coral reefs

8532-484: The mainland. Unlike fringing and barrier reefs which extend only seaward, platform reefs grow in all directions. They are variable in size, ranging from a few hundred metres to many kilometres across. Their usual shape is oval to elongated. Parts of these reefs can reach the surface and form sandbanks and small islands around which may form fringing reefs. A lagoon may form In the middle of a platform reef. Platform reefs are typically situated within atolls, where they adopt

8640-400: The name "patch reefs" and often span a diameter of just a few dozen meters. In instances where platform reefs develop along elongated structures, such as old and weathered barrier reefs, they tend to arrange themselves in a linear formation. This is the case, for example, on the east coast of the Red Sea near Jeddah . In old platform reefs, the inner part can be so heavily eroded that it forms

8748-562: The name "the Old Red Continent". For much of the Devonian, the majority of western Laurussia (North America) was covered by subtropical inland seas which hosted a diverse ecosystem of reefs and marine life. Devonian marine deposits are particularly prevalent in the midwestern and northeastern United States. Devonian reefs also extended along the southeast edge of Laurussia, a coastline now corresponding to southern England , Belgium , and other mid-latitude areas of Europe. In

8856-528: The name implies, coral reefs are made up of coral skeletons from mostly intact coral colonies. As other chemical elements present in corals become incorporated into the calcium carbonate deposits, aragonite is formed. However, shell fragments and the remains of coralline algae such as the green-segmented genus Halimeda can add to the reef's ability to withstand damage from storms and other threats. Such mixtures are visible in structures such as Eniwetok Atoll . The times of maximum reef development were in

8964-625: The naming of the period. Murchison and Sedgwick won the debate and named it 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 Ma, to the beginning of the Carboniferous 358.9 Ma – in North America , at

9072-411: The nearby microcontinent of Amuria (now Manchuria , Mongolia and their vicinities). Though certainly close to Siberia in the Devonian, the precise location of Amuria is uncertain due to contradictory paleomagnetic data. The Rheic Ocean, which separated Laurussia from Gondwana, was wide at the start of the Devonian, having formed after the drift of Avalonia away from Gondwana. It steadily shrunk as

9180-415: The ocean, and atolls form when an island subsides below the surface of the sea. Alternatively, Moyle and Cech distinguish six zones, though most reefs possess only some of the zones. The reef surface is the shallowest part of the reef. It is subject to surge and tides . When waves pass over shallow areas, they shoal , as shown in the adjacent diagram. This means the water is often agitated. These are

9288-528: The oceans formed the atolls. Darwin set out a sequence of three stages in atoll formation. A fringing reef forms around an extinct volcanic island as the island and ocean floor subside. As the subsidence continues, the fringing reef becomes a barrier reef and ultimately an atoll reef. Darwin predicted that underneath each lagoon would be a bedrock base, the remains of the original volcano. Subsequent research supported this hypothesis. Darwin's theory followed from his understanding that coral polyps thrive in

9396-535: The oceans' surface area. The Indo-Pacific region (including the Red Sea , Indian Ocean , Southeast Asia and the Pacific ) account for 91.9% of this total. Southeast Asia accounts for 32.3% of that figure, while the Pacific including Australia accounts for 40.8%. Atlantic and Caribbean coral reefs account for 7.6%. Although corals exist both in temperate and tropical waters, shallow-water reefs form only in

9504-490: The offshore outer reef edge formed in open water rather than next to a shoreline. Like an atoll, it is thought that these reefs are formed either as the seabed lowered or sea level rose. Formation takes considerably longer than for a fringing reef, thus barrier reefs are much rarer. The best known and largest example of a barrier reef is the Australian Great Barrier Reef . Other major examples are

9612-401: The oldest known plants with woody tissue. By the Middle Devonian, shrub-like forests of primitive plants existed: lycophytes , horsetails , ferns , and progymnosperms evolved. Most of these plants had true roots and leaves, and many were quite tall. The earliest-known trees appeared in the Middle Devonian. These included a lineage of lycopods and another arborescent, woody vascular plant,

9720-451: The open ocean, in fact anywhere where the seabed rises close enough to the surface of the ocean to enable the growth of zooxanthemic, reef-forming corals. Platform reefs are found in the southern Great Barrier Reef, the Swain and Capricorn Group on the continental shelf, about 100–200 km from the coast. Some platform reefs of the northern Mascarenes are several thousand kilometres from

9828-703: The order Agoniatitida , which in later 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. The Middle Devonian comprised two subdivisions: first the Eifelian , which then gave way to the Givetian 387.7 Ma. During this time the jawless agnathan fishes began to decline in diversity in freshwater and marine environments partly due to drastic environmental changes and partly due to

9936-403: The period continued, as the two major continents approached near the equator in the early stages of the assembly of Pangaea . The closure of the Rheic Ocean began in the Devonian and continued into the Carboniferous. As the ocean narrowed, endemic marine faunas of Gondwana and Laurussia combined into a single tropical fauna. The history of the western Rheic Ocean is a subject of debate, but there

10044-472: The period it moved northwards and began to twist clockwise, though it was not near its modern location. Siberia approached the eastern edge of Laurussia as the Devonian progressed, but it was still separated by a seaway, the Ural Ocean . Although Siberia's margins were generally tectonically stable and ecologically productive, rifting and deep mantle plumes impacted the continent with flood basalts during

10152-529: The period. Older literature on the Anglo-Welsh basin divides it into the Downtonian, Dittonian, Breconian, and Farlovian stages, the latter three of which are placed in the Devonian. The Devonian has also 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

10260-429: The polyps' tissues and provide organic nutrients that nourish the polyp in the form of glucose , glycerol and amino acids . Because of this relationship, coral reefs grow much faster in clear water, which admits more sunlight. Without their symbionts, coral growth would be too slow to form significant reef structures. Corals get up to 90% of their nutrients from their symbionts. In return, as an example of mutualism ,

10368-420: The precise condition under which corals flourish. The light is sufficient for photosynthesis by the symbiotic zooxanthellae, and agitated water brings plankton to feed the coral. The off-reef floor is the shallow sea floor surrounding a reef. This zone occurs next to reefs on continental shelves. Reefs around tropical islands and atolls drop abruptly to great depths and do not have such a floor. Usually sandy,

10476-613: The rise of the Caledonian Mountains of Great Britain and Scandinavia . As the Caledonian Orogeny wound down in the later part of the period, orogenic collapse facilitated a cluster of granite intrusions in Scotland. Most of Laurussia was located south of the equator, but in the Devonian it moved northwards and began to rotate counterclockwise towards its modern position. While the most northern parts of

10584-412: The sea bed begins to drop steeply. The surface of the fringe reef generally remains at the same height: just below the waterline. In older fringing reefs, whose outer regions pushed far out into the sea, the inner part is deepened by erosion and eventually forms a lagoon . Fringing reef lagoons can become over 100 metres wide and several metres deep. Like the fringing reef itself, they run parallel to

10692-609: The shallow water tropical coral reefs are best known, there are also deeper water reef-forming corals, which live in colder water and in temperate seas. Most coral reefs were formed after the Last Glacial Period when melting ice caused sea level to rise and flood continental shelves . Most coral reefs are less than 10,000 years old. As communities established themselves, the reefs grew upwards, pacing rising sea levels . Reefs that rose too slowly could become drowned, without sufficient light. Coral reefs are also found in

10800-424: The survival of their coral hosts. Each photosynthetic organism has a specific level of sensitivity to photodamage to compounds needed for survival, such as proteins. Rates of regeneration and replication determine the organism's ability to survive. Phylotype A is found more in the shallow waters. It is able to produce mycosporine-like amino acids that are UV resistant , using a derivative of glycerin to absorb

10908-628: The terminus of the Devonian, Earth rapidly cooled into an icehouse , marking the beginning of the Late Paleozoic icehouse . The Devonian world involved many continents and ocean basins of various sizes. The largest continent, Gondwana , was located entirely within the Southern Hemisphere . It corresponds to modern day South America , Africa , Australia , Antarctica , and India , as well as minor components of North America and Asia . The second-largest continent, Laurussia,

11016-535: 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. In fact the climate in the Devonian differed greatly during its epochs and between geographic regions. For example, during the Early Devonian, arid conditions were prevalent through much of the world including Siberia, Australia, North America, and China, but Africa and South America had

11124-521: The warmer conditions of the early and late Devonian, while coral-stromatoporoid reefs dominated during the cooler middle Devonian. By the Devonian Period, life was well underway in its colonization of the land. The moss forests and bacterial and algal mats of the Silurian were joined early in the period by primitive rooted plants that created the first stable soils and harbored arthropods like mites , scorpions , trigonotarbids and myriapods (although arthropods appeared on land much earlier than in

11232-555: The world saw the disappearance of an estimated 96% of vertebrates like conodonts and bony fishes , and all of the ostracoderms and placoderms. Land plants as well as freshwater species, such as our tetrapod ancestors, were relatively unaffected by the Late Devonian extinction event (there is a counterargument that the Devonian extinctions nearly wiped out the tetrapods ). The reasons for the Late Devonian extinctions are still unknown, and all explanations remain speculative. Canadian paleontologist Digby McLaren suggested in 1969 that

11340-680: Was by far the largest continent on the planet. It was completely south of the equator, although the northeastern sector (now Australia) did reach tropical latitudes. The southwestern sector (now South America) was located to the far south, with Brazil situated near the South Pole . The northwestern edge of Gondwana was an active margin for much of the Devonian, and saw the accretion of many smaller land masses and island arcs. These include Chilenia , Cuyania , and Chaitenia , which now form much of Chile and Patagonia . These collisions were associated with volcanic activity and plutons , but by

11448-442: Was mostly a passive margin, hosting extensive marine deposits in areas such as northwest Africa and Tibet . The eastern margin, though warmer than the west, was equally active. Numerous mountain building events and granite and kimberlite intrusions affected areas equivalent to modern day eastern Australia , Tasmania , and Antarctica. Several island microcontinents (which would later coalesce into modern day Asia) stretched over

11556-438: Was northwest of Gondwana, and corresponds to much of modern-day North America and Europe . Various smaller continents, microcontinents , and terranes were present east of Laurussia and north of Gondwana, corresponding to parts of Europe and Asia. The Devonian Period was a time of great tectonic activity, as the major continents of Laurussia and Gondwana drew closer together. Sea levels were high worldwide, and much of

11664-577: Was the newest addition to the Asian microcontinents, it was not the first. North China and the Tarim Block (now northwesternmost China) were located westward and continued to drift northwards, powering over older oceanic crust in the process. Further west was a small ocean (the Turkestan Ocean), followed by the larger microcontinents of Kazakhstania , Siberia , and Amuria . Kazakhstania was

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