Misplaced Pages

#669330

175-500: See taxonomy Brachiopods ( / ˈ b r æ k i oʊ ˌ p ɒ d / ), phylum Brachiopoda , are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs . Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate"

350-399: A ciliated frontmost lobe that becomes the body and lophophore, a rear lobe that becomes the pedicle, and a mantle like a skirt, with the hem towards the rear. On metamorphosing into an adult, the pedicle attaches to a surface, the front lobe develops the lophophore and other organs, and the mantle rolls up over the front lobe and starts to secrete the shell. In cold seas, brachiopod growth

525-425: A matrix of glycosaminoglycans (long, unbranched polysaccharides ), in which other materials are embedded: chitin in the periostracum; apatite containing calcium phosphate in the primary biomineralized layer; and a complex mixture in the innermost layer, containing collagen and other proteins, chitinophosphate and apatite. Craniids , which have no pedicle and cement themselves directly to hard surfaces, have

700-484: A phylum ( / ˈ f aɪ l əm / ; pl. : phyla ) is a level of classification or taxonomic rank below kingdom and above class . Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla,

875-417: A sister group to, the deuterostomes , a superphylum that includes chordates and echinoderms . One type of analysis of the evolutionary relationships of brachiopods has always placed brachiopods as protostomes while another type has split between placing brachiopods among the protostomes or the deuterostomes. It was suggested in 2003 that brachiopods had evolved from an ancestor similar to Halkieria ,

1050-438: A slug -like Cambrian animal with " chain mail " on its back and a shell at the front and rear end; it was thought that the ancestral brachiopod converted its shells into a pair of valves by folding the rear part of its body under its front. However, new fossils found in 2007 and 2008 showed that the "chain mail" of tommotiids formed the tube of a sessile animal; one tommotiid resembled phoronids , which are close relatives or

1225-540: A " living fossil ", as very similar genera have been found all the way back to the Ordovician . On the other hand, articulate brachiopods have produced major diversifications, and suffered severe mass extinctions —but the articulate Rhynchonellida and Terebratulida, the most diverse present-day groups, appeared at the start of the Ordovician and Carboniferous , respectively. Since 1991 Claus Nielsen has proposed

1400-409: A "primary layer" of calcite (a form of calcium carbonate ) under that, and innermost a mixture of proteins and calcite. Inarticulate brachiopod shells have a similar sequence of layers, but their composition is different from that of articulated brachiopods and also varies among the classes of inarticulate brachiopods. The Terebratulida are an example of brachiopods with a punctate shell structure;

1575-431: A 100 kyr Milankovitch cycle , and so each cyclothem represents a cycle of sea level fall and rise over a 100 kyr period. Coal forms when organic matter builds up in waterlogged, anoxic swamps, known as peat mires, and is then buried, compressing the peat into coal. The majority of Earth's coal deposits were formed during the late Carboniferous and early Permian. The plants from which they formed contributed to changes in

1750-426: A bottom-up approach that identifies genera and then groups these into intermediate groups. However, other taxonomists believe that some patterns of characteristics are sufficiently stable to make higher-level classifications worthwhile, although there are different views about what the higher-level classifications should be. The "traditional" classification was defined in 1869; two further approaches were established in

1925-556: A certain degree of morphological or developmental similarity (the phenetic definition), or a group of organisms with a certain degree of evolutionary relatedness (the phylogenetic definition). Attempting to define a level of the Linnean hierarchy without referring to (evolutionary) relatedness is unsatisfactory, but a phenetic definition is useful when addressing questions of a morphological nature—such as how successful different body plans were. The most important objective measure in

SECTION 10

#1732797615670

2100-407: A character unique to a sub-set of the crown group. Furthermore, organisms in the stem group of a phylum can possess the "body plan" of the phylum without all the characteristics necessary to fall within it. This weakens the idea that each of the phyla represents a distinct body plan. A classification using this definition may be strongly affected by the chance survival of rare groups, which can make

2275-453: A comprehensive classification of brachiopods based on morphology. The phylum also has experienced significant convergent evolution and reversals (in which a more recent group seems to have lost a characteristic that is seen in an intermediate group, reverting to a characteristic last seen in an older group). Hence some brachiopod taxonomists believe it is premature to define higher levels of classification such as order , and recommend instead

2450-486: A defined GSSP. The fusulinid Aljutovella aljutovica can be used to define the base of the Moscovian across the northern and eastern margins of Pangea, however, it is restricted in geographic area, which means it cannot be used for global correlations. The first appearance of the conodonts Declinognathodus donetzianus or Idiognathoides postsulcatus have been proposed as a boundary marking species and potential sites in

2625-568: A different opening mechanism, in which muscles reduce the length of the coelom (main body cavity) and make it bulge outwards, pushing the valves apart. Both classes open the valves to an angle of about 10 degrees. The more complex set of muscles employed by inarticulate brachiopods can also operate the valves as scissors, a mechanism that lingulids use to burrow. Each valve consists of three layers, an outer periostracum made of organic compounds and two biomineralized layers. Articulate brachiopods have an outermost periostracum made of proteins ,

2800-469: A few fossils measure up to 200 millimetres (7.9 in) wide. The earliest confirmed brachiopods have been found in the early Cambrian , inarticulate forms appearing first, followed soon after by articulate forms. Three unmineralized species have also been found in the Cambrian, and apparently represent two distinct groups that evolved from mineralized ancestors. The inarticulate Lingula is often called

2975-517: A folding of the upper surface under the body. The ventral ("lower") valve actually lies above the dorsal ("upper") valve when most brachiopods are oriented in life position. In many living articulate brachiopod species, both valves are convex, the surfaces often bearing growth lines and/or other ornamentation. However, inarticulate lingulids, which burrow into the seabed, have valves that are smoother, flatter and of similar size and shape. (R. C. Moore, 1952) Articulate ("jointed") brachiopods have

3150-402: A group ("a self-contained unity"): "perhaps such a real and completely self-contained unity is the aggregate of all species which have gradually evolved from one and the same common original form, as, for example, all vertebrates. We name this aggregate [a] Stamm [i.e., stock] ( Phylon )." In plant taxonomy , August W. Eichler (1883) classified plants into five groups named divisions,

3325-602: A group containing Viridiplantae and the algal Rhodophyta and Glaucophyta divisions. The definition and classification of plants at the division level also varies from source to source, and has changed progressively in recent years. Thus some sources place horsetails in division Arthrophyta and ferns in division Monilophyta, while others place them both in Monilophyta, as shown below. The division Pinophyta may be used for all gymnosperms (i.e. including cycads, ginkgos and gnetophytes), or for conifers alone as below. Since

3500-430: A hypothesis about the development of brachiopods, adapted in 2003 by Cohen and colleagues as a hypothesis about the earliest evolution of brachiopods. This "brachiopod fold" hypothesis suggests that brachiopods evolved from an ancestor similar to Halkieria , a slug -like animal with " chain mail " on its back and a shell at the front and rear end. The hypothesis proposes that the first brachiopod converted its shells into

3675-477: A month and have wide ranges. Brachiopods now live mainly in cold water and low light. Fish and crustaceans seem to find brachiopod flesh distasteful and seldom attack them. Among brachiopods, only the lingulids ( Lingula sp. ) have been fished commercially, on a very small scale. One brachiopod species ( Coptothyrus adamsi ) may be a measure of environmental conditions around an oil terminal being built in Russia on

SECTION 20

#1732797615670

3850-888: A month before settling, have wide ranges. Members of the discinoid genus Pelagodiscus have a cosmopolitan distribution . Brachiopods have a low metabolic rate , between one third and one tenth of that of bivalves . While brachiopods were abundant in warm, shallow seas during the Cretaceous period , most of their former niches are now occupied by bivalves, and most now live in cold and low-light conditions. Brachiopod shells occasionally show evidence of damage by predators, and sometimes of subsequent repair. Fish and crustaceans seem to find brachiopod flesh distasteful. The fossil record shows that drilling predators like gastropods attacked molluscs and echinoids 10 to 20 times more often than they did brachiopods, suggesting that such predators attacked brachiopods by mistake or when other prey

4025-476: A more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening. Brachiopod lifespans range from three to over thirty years. Ripe gametes ( ova or sperm ) float from

4200-402: A muscular heart lying in the dorsal part of the body above the stomach. The blood passes through vessels that extend to the front and back of the body, and branch to organs including the lophophore at the front and the gut, muscles, gonads and nephridia at the rear. The blood circulation seems not to be completely closed, and the coelomic fluid and blood must mix to a degree. The main function of

4375-504: A pair of valves by folding the rear part of its body under its front. However, fossils from 2007 onwards have supported a new interpretation of the Early-Cambrian tommotiids , and a new hypothesis that brachiopods evolved from tommotiids. The "armor mail" of tommotiids was well-known but not in an assembled form, and it was generally assumed that tommotiids were slug-like animals similar to Halkieria , except that tommotiids' armor

4550-429: A periostracum of chitin and mineralized layers of calcite. Shell growth can be described as holoperipheral, mixoperipheral, or hemiperipheral. In holoperipheral growth, distinctive of craniids, new material is added at an equal rate all around the margin. In mixoperipheral growth, found in many living and extinct articulates, new material is added to the posterior region of the shell with an anterior trend, growing towards

4725-400: A phylum based on body plan has been proposed by paleontologists Graham Budd and Sören Jensen (as Haeckel had done a century earlier). The definition was posited because extinct organisms are hardest to classify: they can be offshoots that diverged from a phylum's line before the characters that define the modern phylum were all acquired. By Budd and Jensen's definition, a phylum is defined by

4900-471: A phylum much more diverse than it would be otherwise. Total numbers are estimates; figures from different authors vary wildly, not least because some are based on described species, some on extrapolations to numbers of undescribed species. For instance, around 25,000–27,000 species of nematodes have been described, while published estimates of the total number of nematode species include 10,000–20,000; 500,000; 10 million; and 100 million. The kingdom Plantae

5075-706: A phylum, other phylum-level ranks appear, such as the case of Bacillariophyta (diatoms) within Ochrophyta . These differences became irrelevant after the adoption of a cladistic approach by the ISP, where taxonomic ranks are excluded from the classifications after being considered superfluous and unstable. Many authors prefer this usage, which lead to the Chromista-Protozoa scheme becoming obsolete. Currently there are 40 bacterial phyla (not including " Cyanobacteria ") that have been validly published according to

5250-401: A set of characters shared by all its living representatives. This approach brings some small problems—for instance, ancestral characters common to most members of a phylum may have been lost by some members. Also, this definition is based on an arbitrary point of time: the present. However, as it is character based, it is easy to apply to the fossil record. A greater problem is that it relies on

5425-431: A shallow, tropical seaway which stretched from Southern California to Alaska. The boundary is within a cyclothem sequence of transgressive limestones and fine sandstones , and regressive mudstones and brecciated limestones. The Moscovian Stage is named after shallow marine limestones and colourful clays found around Moscow, Russia. It was first introduced by Sergei Nikitin in 1890. The Moscovian currently lacks

Brachiopod - Misplaced Pages Continue

5600-441: A shape resembling a hand with the fingers splayed. In all species the lophophore is supported by cartilage and by a hydrostatic skeleton (in other words, by the pressure of its internal fluid), and the fluid extends into the tentacles. Some articulate brachiopods also have a brachidium, a calcareous support for the lophophore attached to the inside of the brachial valve, which have led to an extremely reduced lophophoral muscles and

5775-451: A single sedimentary cycle, with an erosional surface at its base. Whilst individual cyclothems are often only metres to a few tens of metres thick, cyclothem sequences can be many hundreds to thousands of metres thick and contain tens to hundreds of individual cyclothems. Cyclothems were deposited along continental shelves where the very gentle gradient of the shelves meant even small changes in sea level led to large advances or retreats of

5950-509: A sub-group of brachiopods. Paterimitra , another mostly assembled fossil found in 2008 and described in 2009, had two symmetrical plates at the bottom, like brachiopod valves but not fully enclosing the animal's body. At their peak in the Paleozoic , the brachiopods were among the most abundant filter-feeders and reef-builders, and occupied other ecological niches , including swimming in the jet-propulsion style of scallops . However, after

6125-400: A subgroup of brachiopods, while the other tommotiid bore two symmetrical plates that might be an early form of brachiopod valves. Lineages of brachiopods that have both fossil and extant taxa appeared in the early Cambrian , Ordovician , and Carboniferous periods , respectively. Other lineages have arisen and then become extinct, sometimes during severe mass extinctions . At their peak in

6300-401: A subjective decision about which groups of organisms should be considered as phyla. The approach is useful because it makes it easy to classify extinct organisms as " stem groups " to the phyla with which they bear the most resemblance, based only on the taxonomically important similarities. However, proving that a fossil belongs to the crown group of a phylum is difficult, as it must display

6475-424: A term that remains in use today for groups of plants, algae and fungi. The definitions of zoological phyla have changed from their origins in the six Linnaean classes and the four embranchements of Georges Cuvier . Informally, phyla can be thought of as groupings of organisms based on general specialization of body plan . At its most basic, a phylum can be defined in two ways: as a group of organisms with

6650-409: A tooth and socket arrangement by which the pedicle and brachial valves hinge, locking the valves against lateral displacement. Inarticulate brachiopods have no matching teeth and sockets; their valves are held together only by muscles. (R. C. Moore, 1952) All brachiopods have adductor muscles that are set on the inside of the pedicle valve and which close the valves by pulling on the part of

6825-648: A tropical wetland environment. Extensive coal deposits developed within the cyclothem sequences that dominated the Pennsylvanian sedimentary basins associated with the growing orogenic belt. Subduction of the Panthalassic oceanic plate along its western margin resulted in the Antler orogeny in the Late Devonian to Early Mississippian. Further north along the margin, slab roll-back , beginning in

7000-423: A variety of methods for reconstructing past atmospheric oxygen levels, including the charcoal record, halite gas inclusions, burial rates of organic carbon and pyrite , carbon isotopes of organic material, isotope mass balance and forward modelling. Depending on the preservation of source material, some techniques represent moments in time (e.g. halite gas inclusions), whilst others have a wider time range (e.g.

7175-750: A warmer climate. This rapid rise in CO 2 may have been due to a peak in pyroclastic volcanism and/or a reduction in burial of terrestrial organic matter. The LPIA peaked across the Carboniferous-Permian boundary. Widespread glacial deposits are found across South America, western and central Africa, Antarctica, Australia, Tasmania, the Arabian Peninsula, India, and the Cimmerian blocks, indicating trans-continental ice sheets across southern Gondwana that reached to sea-level. In response to

Brachiopod - Misplaced Pages Continue

7350-474: Is a paraphyletic taxon, which is less acceptable to present-day biologists than in the past. Proposals have been made to divide it among several new kingdoms, such as Protozoa and Chromista in the Cavalier-Smith system . Protist taxonomy has long been unstable, with different approaches and definitions resulting in many competing classification schemes. Many of the phyla listed below are used by

7525-556: Is a ring of tentacles mounted on a single, retracted stalk, while the basic form of the brachiopod lophophore is U-shaped, forming the brachia ("arms") from which the phylum gets its name. Brachiopod lophophores are non-retractable and occupy up to two-thirds of the internal space, in the frontmost area where the valves gape when opened. To provide enough filtering capacity in this restricted space, lophophores of larger brachiopods are folded in moderately to very complex shapes—loops and coils are common, and some species' lophophores contort into

7700-405: Is defined in various ways by different biologists (see Current definitions of Plantae ). All definitions include the living embryophytes (land plants), to which may be added the two green algae divisions, Chlorophyta and Charophyta , to form the clade Viridiplantae . The table below follows the influential (though contentious) Cavalier-Smith system in equating "Plantae" with Archaeplastida ,

7875-429: Is digested, mainly within the cells. Nutrients are transported throughout the coelom, including the mantle lobes, by cilia. The wastes produced by metabolism are broken into ammonia , which is eliminated by diffusion through the mantle and lophophore. Brachiopods have metanephridia , used by many phyla to excrete ammonia and other dissolved wastes. However, brachiopods have no sign of the podocytes , which perform

8050-481: Is generally included in kingdom Fungi, though its exact relations remain uncertain, and it is considered a protozoan by the International Society of Protistologists (see Protista , below). Molecular analysis of Zygomycota has found it to be polyphyletic (its members do not share an immediate ancestor), which is considered undesirable by many biologists. Accordingly, there is a proposal to abolish

8225-578: Is inconclusive as to the exact relations within the inarticulates. Consequently, it has been suggested to include horseshoe worms in the Brachiopoda as a class named Phoronata ( B.L.Cohen & Weydmann ) in addition to the Craniata and Lingulata, within the subphylum Linguliformea. The other subphylum, Rhynchonelliformea contains only one extant class, which is subdivided into the extant orders Rhynchonellida, Terebratulida and Thecideida. This shows

8400-516: Is located in Bed 83 of the sequence of dark grey limestones and shales at the Pengchong section, Guangxi , southern China. It is defined by the first appearance of the fusulinid Eoparastaffella simplex in the evolutionary lineage Eoparastaffella ovalis – Eoparastaffella simplex and was ratified in 2009. The Serpukhovian Stage was proposed in 1890 by Russian stratigrapher Sergei Nikitin . It

8575-528: Is named after the Russian village of Gzhel , near Ramenskoye , not far from Moscow. The name and type locality were defined by Sergei Nikitin in 1890. The Gzhelian currently lacks a defined GSSP. The first appearance of the fusulinid Rauserites rossicus and Rauserites stuckenbergi can be used in the Boreal Sea and Paleo-Tethyan regions but not eastern Pangea or Panthalassa margins. Potential sites in

8750-438: Is named after the city of Serpukhov , near Moscow. currently lacks a defined GSSP. The Visean-Serpukhovian boundary coincides with a major period of glaciation. The resulting sea level fall and climatic changes led to the loss of connections between marine basins and endemism of marine fauna across the Russian margin. This means changes in biota are environmental rather than evolutionary making wider correlation difficult. Work

8925-581: Is no evidence that the latest common ancestor of pterobranchs and other hemichordates or the latest common ancestor of hemichordates and echinoderms was sessile and fed by means of tentacles. From 1988 onwards analyses based on molecular phylogeny , which compares biochemical features such as similarities in DNA , have placed brachiopods among the Lophotrochozoa , a protostome super-phylum that includes molluscs , annelids and flatworms but excludes

SECTION 50

#1732797615670

9100-402: Is only about 1 millimetre (0.039 in) long, and lives in between gravel grains. Rhynchonelliforms, whose larvae consume only their yolks and settle and develop quickly, are often endemic to an area and form dense populations that can reach thousands per meter. Young adults often attach to the shells of more mature ones. On the other hand, inarticulate brachiopods, whose larva swim for up to

9275-400: Is radial (cells form in stacks of rings directly above each other), holoblastic (cells are separate, although adjoining) and regulative (the type of tissue into which a cell develops is controlled by interactions between adjacent cells, rather than rigidly within each cell). While some animals develop the mouth and anus by deepening the blastopore , a "dent" in the surface of the early embryo,

9450-455: Is seasonal and the animals often lose weight in winter. These variations in growth often form growth lines in the shells. Members of some genera have survived for a year in aquaria without food. Brachiopod fossils show great diversity in the morphology of the shells and lophophore, while the modern genera show less diversity but provide soft-bodied characteristics. Both fossils and extant species have limitations that make it difficult to produce

9625-413: Is solid and the flow runs from bases to tips, forming a "downstream collecting" system that catches food particles as they are about to exit. Most modern species attach to hard surfaces by means of a cylindrical pedicle ("stalk"), an extension of the body wall. This has a chitinous cuticle (non-cellular "skin") and protrudes through an opening in the hinge. However, some genera have no pedicle, such as

9800-631: Is that a delay between the development of trees with the wood fibre lignin and the subsequent evolution of lignin-degrading fungi gave a period of time where vast amounts of lignin-based organic material could accumulate. Genetic analysis of basidiomycete fungi, which have enzymes capable of breaking down lignin, supports this theory by suggesting this fungi evolved in the Permian. However, significant Mesozoic and Cenozoic coal deposits formed after lignin-digesting fungi had become well established, and fungal degradation of lignin may have already evolved by

9975-582: Is the largest extant species. The largest brachiopods known— Gigantoproductus and Titanaria , reaching 30 to 38 centimetres (12 to 15 in) in width—occurred in the upper part of the Lower Carboniferous. Brachiopods have two valves (shell sections), which cover the dorsal (top) and ventral (bottom) surface of the animal, unlike bivalve molluscs whose shells cover the lateral surfaces (sides). The valves are unequal in size and structure, with each having its own symmetrical form rather than

10150-467: Is unclear. It is constructed from a different part of the larval body, and has a compact core composed of connective tissue . Muscles at the rear of the body can straighten, bend or even rotate the pedicle. The far end of the pedicle generally has rootlike extensions or short papillae ("bumps"), which attach to hard surfaces. However, articulate brachiopods of the genus Chlidonophora use a branched pedicle to anchor in sediment . The pedicle emerges from

10325-663: Is underway in the Urals and Nashui, Guizhou Province, southwestern China for a suitable site for the GSSP with the proposed definition for the base of the Serpukhovian as the first appearance of conodont Lochriea ziegleri . The Pennsylvanian was proposed by J.J.Stevenson in 1888, named after the widespread coal-rich strata found across the state of Pennsylvania. The closure of the Rheic Ocean and formation of Pangea during

10500-444: Is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and

10675-402: Is usually larger, and near the hinge it has an opening for the stalk-like pedicle through which most brachiopods attach themselves to the substrate. ( R. C. Moore , 1952) The brachial and pedicle valves are often called the dorsal and ventral valves, respectively, but some paleontologists regard the terms "dorsal" and "ventral" as irrelevant since they believe that the "ventral" valve was formed by

SECTION 60

#1732797615670

10850-643: The Bacteriological Code Currently there are 2 phyla that have been validly published according to the Bacteriological Code Other phyla that have been proposed, but not validly named, include: Carboniferous The name Carboniferous means " coal -bearing", from the Latin carbō (" coal ") and ferō ("bear, carry"), and refers to the many coal beds formed globally during that time. The first of

11025-565: The Catalogue of Life , and correspond to the Protozoa-Chromista scheme, with updates from the latest (2022) publication by Cavalier-Smith . Other phyla are used commonly by other authors, and are adapted from the system used by the International Society of Protistologists (ISP). Some of the descriptions are based on the 2019 revision of eukaryotes by the ISP. The number of protist phyla varies greatly from one classification to

11200-717: The Gulf of Mexico in the west to Turkey in the east. The orogeny was caused by a series of continental collisions between Laurussia, Gondwana and the Armorican Terrane Assemblage (much of modern-day Central and Western Europe including Iberia ) as the Rheic Ocean closed and Pangea formed. This mountain building process began in the Middle Devonian and continued into the early Permian. The Armorican terranes rifted away from Gondwana during

11375-751: The International Commission on Stratigraphy (ICS) stage, but the Viséan is longer, extending into the lower Serpukhovian . North American geologists recognised a similar stratigraphy but divided it into two systems rather than one. These are the lower carbonate-rich sequence of the Mississippian System and the upper siliciclastic and coal-rich sequence of the Pennsylvanian . The United States Geological Survey officially recognised these two systems in 1953. In Russia, in

11550-677: The Kuznetsk Basin . The northwest to eastern margins of Siberia were passive margins along the Mongol-Okhotsk Ocean on the far side of which lay Amuria. From the mid Carboniferous, subduction zones with associated magmatic arcs developed along both margins of the ocean. The southwestern margin of Siberia was the site of a long lasting and complex accretionary orogen. The Devonian to early Carboniferous Siberian and South Chinese Altai accretionary complexes developed above an east-dipping subduction zone, whilst further south,

11725-586: The Magnitogorsk island arc , which lay between Kazakhstania and Laurussia in the Ural Ocean , collided with the passive margin of northeastern Laurussia ( Baltica craton ). The suture zone between the former island arc complex and the continental margin formed the Main Uralian Fault , a major structure that runs for more than 2,000 km along the orogen. Accretion of the island arc

11900-684: The Old Red Sandstone , Carboniferous Limestone , Millstone Grit and the Coal Measures . These four units were placed into a formalised Carboniferous unit by William Conybeare and William Phillips in 1822 and then into the Carboniferous System by Phillips in 1835. The Old Red Sandstone was later considered Devonian in age. The similarity in successions between the British Isles and Western Europe led to

12075-550: The Paleozoic era , the brachiopods were among the most abundant filter-feeders and reef-builders, and occupied other ecological niches , including swimming in the jet-propulsion style of scallops . Brachiopod fossils have been useful indicators of climate changes during the Paleozoic. However, after the Permian–Triassic extinction event , brachiopods recovered only a third of their former diversity. A study in 2007 concluded

12250-401: The Permian–Triassic extinction event , informally known as the "Great Dying", brachiopods recovered only a third of their former diversity. It was often thought that brachiopods were actually declining in diversity, and that in some way bivalves out-competed them. However, in 1980, Gould and Calloway produced a statistical analysis that concluded that both brachiopods and bivalves increased all

12425-399: The gonads into the main coelom and then exit into the mantle cavity. The larvae of inarticulate brachiopods are miniature adults, with lophophores that enable the larvae to feed and swim for months until the animals become heavy enough to settle to the seabed. The planktonic larvae of articulate species do not resemble the adults, but rather look like blobs with yolk sacs , and remain among

12600-505: The "pedicle sheath", which has no relationship to the pedicle. This structure arises from the umbo of the pedicle valve, at the centre of the earliest (metamorphic) shell at the location of the protegulum. It is sometimes associated with a fringing plate, the colleplax. The water flow enters the lophophore from the sides of the open valves and exits at the front of the animal. In lingulids the entrance and exit channels are formed by groups of chaetae that function as funnels. In other brachiopods

12775-713: The 1840s British and Russian geologists divided the Carboniferous into the Lower, Middle and Upper series based on Russian sequences. In the 1890s these became the Dinantian, Moscovian and Uralian stages. The Serpukivian was proposed as part of the Lower Carboniferous, and the Upper Carboniferous was divided into the Moscovian and Gzhelian . The Bashkirian was added in 1934. In 1975, the ICS formally ratified

12950-472: The 1990s: About 330 living species are recognized, grouped into over 100  genera . The great majority of modern brachiopods are rhynchonelliforms (Articulata). Genetic analysis performed since the 1990s has extended the understanding of the relationship between different organisms. It is now clear the brachiopods do not belong to the Deuterostomia (such as echinoderms and chordates ) as

13125-526: The Bashkirian, the late Moscovian and the latest Kasimovian to mid-Gzhelian are inferred from the disappearance of glacial sediments, the appearance of deglaciation deposits and rises in sea levels. In the early Kasimovian there was short-lived (<1 million years) intense period of glaciation, with atmospheric CO 2 concentration levels dropping as low as 180 ppm. This ended suddenly as a rapid increase in CO 2 concentrations to c. 600 ppm resulted in

13300-764: The Carboniferous Earth's atmosphere, and the coal fueled the Industrial Revolution . During the Pennsylvanian, vast amounts of organic debris accumulated in the peat mires that formed across the low-lying, humid equatorial wetlands of the foreland basins of the Central Pangean Mountains in Laurussia, and around the margins of the North and South China cratons. During glacial periods, low sea levels exposed large areas of

13475-669: The Carboniferous System, with the Mississippian and Pennsylvanian subsystems from the North American timescale, the Tournaisian and Visean stages from the Western European and the Serpukhovian, Bashkirian, Moscovian, Kasimovian and Gzhelian from the Russian. With the formal ratification of the Carboniferous System, the Dinantian, Silesian, Namurian, Westphalian and Stephanian became redundant terms, although

13650-609: The Carboniferous, the Tarim craton lay along the northwestern edge of North China. Subduction along the Kazakhstanian margin of the Turkestan Ocean resulted in collision between northern Tarim and Kazakhstania during the mid Carboniferous as the ocean closed. The South Tian Shan fold and thrust belt , which extends over 2,000 km from Uzbekistan to northwest China, is the remains of this accretionary complex and forms

13825-527: The Craniida to be a separate third group, as their outer organic layer is distinct from that of both the linguliforms ("typical" inarticulates) and rhynchonelliforms (articulates). However, some taxonomists believe it is premature to suggest higher levels of classification such as order and recommend a bottom-up approach that identifies genera and then groups these into intermediate groups. Traditionally, brachiopods have been regarded as members of, or as

14000-762: The Early Mississippian, led to the rifting of the Yukon-Tanana terrane and the opening of the Slide Mountain Ocean . Along the northern margin of Laurussia, orogenic collapse of the Late Devonian to Early Mississippian Innuitian orogeny led to the development of the Sverdrup Basin . Much of Gondwana lay in the southern polar region during the Carboniferous. As the plate moved, the South Pole drifted from southern Africa in

14175-513: The Early to Middle Mississippian, carbonate production occurred to depth across the gently dipping continental slopes of Laurussia and North and South China ( carbonate ramp architecture) and evaporites formed around the coastal regions of Laurussia, Kazakhstania, and northern Gondwana. From the late Visean, the cooling climate restricted carbonate production to depths of less than c. 10 m forming carbonate shelves with flat-tops and steep sides. By

14350-491: The Kasimovian covers a period of globally low sea level, which has resulted in disconformities within many sequences of this age. This has created difficulties in finding suitable marine fauna that can used to correlate boundaries worldwide. The Kasimovian currently lacks a defined GSSP; potential sites in the southern Urals, southwest USA and Nashui, Guizhou Province, southwestern China are being considered. The Gzhelian

14525-483: The Late Ordovician . As they drifted northwards the Rheic Ocean closed in front of them, and they began to collide with southeastern Laurussia in the Middle Devonian. The resulting Variscan orogeny involved a complex series of oblique collisions with associated metamorphism , igneous activity, and large-scale deformation between these terranes and Laurussia, which continued into the Carboniferous. During

14700-665: The Late Pennsylvanian, deformation along the Alleghanian orogen became northwesterly-directed compression . The Uralian orogeny is a north–south trending fold and thrust belt that forms the western edge of the Central Asian Orogenic Belt . The Uralian orogeny began in the Late Devonian and continued, with some hiatuses, into the Jurassic . From the Late Devonian to early Carboniferous,

14875-637: The Paleo-Tethys to the southwest and Panthalassa to the northeast. Cyclothem sediments with coal and evaporites were deposited across the passive margins that surrounded both continents. The Carboniferous climate was dominated by the Late Paleozoic Ice Age (LPIA), the most extensive and longest icehouse period of the Phanerozoic, which lasted from the Late Devonian to the Permian (365 Ma-253 Ma). Temperatures began to drop during

15050-485: The Paleozoic to modern times, with bivalves increasing faster; after the Permian–Triassic extinction, brachiopods became for the first time less diverse than bivalves. Brachiopods live only in the sea, and most species avoid locations with strong currents or waves. The larvae of articulate species settle in quickly and form dense populations in well-defined areas while the larvae of inarticulate species swim for up to

15225-463: The Pennsylvanian, cyclothems were deposited in shallow, epicontinental seas across the tropical regions of Laurussia (present day western and central US, Europe, Russia and central Asia) and the North and South China cratons . The rapid sea levels fluctuations they represent correlate with the glacial cycles of the Late Paleozoic Ice Age. The advance and retreat of ice sheets across Gondwana followed

15400-402: The Pennsylvanian, together with widespread glaciation across Gondwana led to major climate and sea level changes, which restricted marine fauna to particular geographic areas thereby reducing widespread biostratigraphic correlations. Extensive volcanic events associated with the assembling of Pangea means more radiometric dating is possible relative to the Mississippian. The Bashkirian Stage

15575-479: The Period. This was not a steady rise, but included peaks and troughs reflecting the dynamic climate conditions of the time. How the atmospheric oxygen concentrations influenced the large body size of arthropods and other fauna and flora during the Carboniferous is also a subject of ongoing debate. The changing climate was reflected in regional-scale changes in sedimentation patterns. In the relatively warm waters of

15750-485: The Permian. The Kazakhstanian microcontinent is composed of a series of Devonian and older accretionary complexes. It was strongly deformed during the Carboniferous as its western margin collided with Laurussia during the Uralian orogen and its northeastern margin collided with Siberia. Continuing strike-slip motion between Laurussia and Siberia led the formerly elongate microcontinent to bend into an orocline . During

15925-450: The Permian–Triassic extinction, as all had calcareous hard parts (made of calcium carbonate ) and had low metabolic rates and weak respiratory systems. Brachiopod fossils have been useful indicators of climate changes during the Paleozoic era. When global temperatures were low, as in much of the Ordovician , the large difference in temperature between equator and poles created different collections of fossils at different latitudes . On

16100-555: The Urals and Nashui, Guizhou Province, southwestern China are being considered. The Kasimovian is the first stage in the Upper Pennsylvanian. It is named after the Russian city of Kasimov , and was originally included as part of Nikitin's 1890 definition of the Moscovian. It was first recognised as a distinct unit by A.P. Ivanov in 1926, who named it the " Tiguliferina " Horizon after a type of brachiopod . The boundary of

16275-498: The Urals and Nashui, Guizhou Province, southwestern China for the GSSP are being considered. The GSSP for the base of the Permian is located in the Aidaralash River valley near Aqtöbe , Kazakhstan and was ratified in 1996. The beginning of the stage is defined by the first appearance of the conodont Streptognathodus postfusus . A cyclothem is a succession of non-marine and marine sedimentary rocks , deposited during

16450-467: The Visean of c. 15.3%, although with large uncertainties; and, pyrite records suggest levels of c. 15% early in the Carboniferous, to over 25% during the Pennsylvanian, before dropping back below 20% towards the end. However, whilst exact numbers vary, all models show an overall increase in atmospheric oxygen levels from a low of between 15-20% at the beginning of the Carboniferous to highs of 25-30% during

16625-522: The Zharma-Saur arc formed along the northeastern margin of Kazakhstania. By the late Carboniferous, all these complexes had accreted to the Siberian craton as shown by the intrusion of post-orogenic granites across the region. As Kazakhstania had already accreted to Laurussia, Siberia was effectively part of Pangea by 310 Ma, although major strike-slip movements continued between it and Laurussia into

16800-497: The Zygomycota phylum. Its members would be divided between phylum Glomeromycota and four new subphyla incertae sedis (of uncertain placement): Entomophthoromycotina , Kickxellomycotina , Mucoromycotina , and Zoopagomycotina . Kingdom Protista (or Protoctista) is included in the traditional five- or six-kingdom model, where it can be defined as containing all eukaryotes that are not plants, animals, or fungi. Protista

16975-458: The above definitions is the "certain degree" that defines how different organisms need to be members of different phyla. The minimal requirement is that all organisms in a phylum should be clearly more closely related to one another than to any other group. Even this is problematic because the requirement depends on knowledge of organisms' relationships: as more data become available, particularly from molecular studies, we are better able to determine

17150-485: The animals and may act as sensors . In some brachiopods groups of chaetae help to channel the flow of water into and out of the mantle cavity. In most brachiopods, diverticula (hollow extensions) of the mantle penetrate through the mineralized layers of the valves into the periostraca. The function of these diverticula is uncertain and it is suggested that they may be storage chambers for chemicals such as glycogen , may secrete repellents to deter organisms that stick to

17325-668: The average temperature in the tropics c. 24 °C (75 °F) and in polar regions c. -23 °C (-10 °F), whilst during the Early Tournaisian Warm Interval (358-353 Ma) the GAT was c. 22 °C (72 °F), the tropics c. 30 °C (86 °F) and polar regions c. 1.5 °C (35 °F). Overall, for the Ice Age the GAT was c. 17 °C (62 °F), with tropical temperatures c. 26 °C and polar temperatures c. -9.0 °C (16 °F). There are

17500-528: The base of the Carboniferous System, Mississippian Subsystem and Tournaisian Stage is located at the La Serre section in Montagne Noire , southern France. It is defined by the first appearance of the conodont Siphonodella sulcata within the evolutionary lineage from Siphonodella praesulcata to Siphonodella sulcata . This was ratified by the ICS in 1990. However, in 2006 further study revealed

17675-409: The bases of the tentacles, and its own cilia pass food along the groove towards the mouth. The method used by brachiopods is known as "upstream collecting", as food particles are captured as they enter the field of cilia that creates the feeding current. This method is used by the related phoronids and bryozoans , and also by pterobranchs . Entoprocts use a similar-looking crown of tentacles, but it

17850-433: The beginning of the Period to highs of 25-30%. The development of a Carboniferous chronostratigraphic timescale began in the late 18th century. The term "Carboniferous" was first used as an adjective by Irish geologist Richard Kirwan in 1799 and later used in a heading entitled "Coal-measures or Carboniferous Strata" by John Farey Sr. in 1811. Four units were originally ascribed to the Carboniferous, in ascending order,

18025-434: The blastopore of brachiopods closes up, and their mouth and anus develop from new openings. The larvae of lingulids (Lingulida and Discinida) are planktotrophic (feeding), and swim as plankton for months resembling miniature adults, with valves, mantle lobes, a pedicle that coils in the mantle cavity, and a small lophophore, which is used for both feeding and swimming. The larvae of craniids have no pedicle or shell. As

18200-471: The blood may be to deliver nutrients. The "brain" of adult articulates consists of two ganglia , one above and the other below the oesophagus . Adult inarticulates have only the lower ganglion. From the ganglia and the commissures where they join, nerves run to the lophophore, the mantle lobes and the muscles that operate the valves. The edge of the mantle has probably the greatest concentration of sensors. Although not directly connected to sensory neurons ,

18375-412: The brachial valve ahead of the hinge. These muscles have both "quick" fibers that close the valves in emergencies and "catch" fibers that are slower but can keep the valves closed for long periods. Articulate brachiopods open the valves by means of abductor muscles, also known as diductors, which lie further to the rear and pull on the part of the brachial valve behind the hinge. Inarticulate brachiopods use

18550-411: The brachiopods were especially vulnerable to the Permian–Triassic extinction, as they built calcareous hard parts (made of calcium carbonate ) and had low metabolic rates and weak respiratory systems. It was often thought that brachiopods went into decline after the Permian–Triassic extinction, and were out-competed by bivalves, but a study in 1980 found both brachiopod and bivalve species increased from

18725-403: The channels of the mantle lobes, while those of inarticulates lie near the gut. Ripe gametes float into the main coelom and then exit into the mantle cavity via the metanephridia , which open on either side of the mouth. Most species release both ova and sperm into the water, but females of some species keep the embryos in brood chambers until the larvae hatch. The cell division in the embryo

18900-410: The charcoal record and pyrite). Results from these different methods for the Carboniferous vary. For example: the increasing occurrence of charcoal produced by wildfires from the Late Devonian into the Carboniferous indicates increasing oxygen levels, with calculations showing oxygen levels above 21% for most of the Carboniferous; halite gas inclusions from sediments dated 337-335 Ma give estimates for

19075-579: The city of Visé , Liège Province , Belgium. In 1967, the base of the Visean was officially defined as the first black limestone in the Leffe facies at the Bastion Section in the Dinant Basin . These changes are now thought to be ecologically driven rather than caused by evolutionary change, and so this has not been used as the location for the GSSP. Instead, the GSSP for the base of the Visean

19250-471: The complexity of the geology. The ICS subdivisions from youngest to oldest are as follows: The Mississippian was proposed by Alexander Winchell in 1870 named after the extensive exposure of lower Carboniferous limestone in the upper Mississippi River valley. During the Mississippian, there was a marine connection between the Paleo-Tethys and Panthalassa through the Rheic Ocean resulting in

19425-437: The continental shelves. Major river channels, up to several kilometres wide, stretched across these shelves feeding a network of smaller channels, lakes and peat mires. These wetlands were then buried by sediment as sea levels rose during interglacials . Continued crustal subsidence of the foreland basins and continental margins allowed this accumulation and burial of peat deposits to continue over millions of years resulting in

19600-521: The development of a common European timescale with the Carboniferous System divided into the lower Dinantian , dominated by carbonate deposition and the upper Silesian with mainly siliciclastic deposition. The Dinantian was divided into the Tournaisian and Viséan stages. The Silesian was divided into the Namurian , Westphalian and Stephanian stages. The Tournaisian is the same length as

19775-482: The digestive tract is U-shaped and ends with an anus that eliminates solids from the front of the body wall. Other inarticulate brachiopods and all articulate brachiopods have a curved gut that ends blindly, with no anus. These animals bundle solid waste with mucus and periodically "sneeze" it out, using sharp contractions of the gut muscles. The lophophore and mantle are the only surfaces that absorb oxygen and eliminate carbon dioxide . Oxygen seems to be distributed by

19950-475: The early Carboniferous Kanimblan Orogeny . Continental arc magmatism continued into the late Carboniferous and extended round to connect with the developing proto-Andean subduction zone along the western South American margin of Gondwana. Shallow seas covered much of the Siberian craton in the early Carboniferous. These retreated as sea levels fell in the Pennsylvanian and as the continent drifted north into more temperate zones extensive coal deposits formed in

20125-599: The early Carboniferous in North China. However, bauxite deposits immediately above the regional mid Carboniferous unconformity indicate warm tropical conditions and are overlain by cyclothems including extensive coals. South China and Annamia (Southeast Asia) rifted from Gondwana during the Devonian. During the Carboniferous, they were separated from each other and North China by the Paleoasian Ocean with

20300-571: The early Carboniferous to eastern Antarctica by the end of the period. Glacial deposits are widespread across Gondwana and indicate multiple ice centres and long-distance movement of ice. The northern to northeastern margin of Gondwana (northeast Africa, Arabia, India and northeastern West Australia) was a passive margin along the southern edge of the Paleo-Tethys with cyclothem deposition including, during more temperate intervals, coal swamps in Western Australia. The Mexican terranes along

20475-539: The end of the Carboniferous, extension and rifting across the northern margin of Gondwana led to the breaking away of the Cimmerian Terrane during the early Permian and the opening of the Neo-Tethys Ocean . Along the southeastern and southern margin of Gondwana (eastern Australia and Antarctica), northward subduction of Panthalassa continued. Changes in the relative motion of the plates resulted in

20650-455: The end of the Devonian, even if the specific enzymes used by basidiomycetes had not. The second theory is that the geographical setting and climate of the Carboniferous were unique in Earth's history: the co-occurrence of the position of the continents across the humid equatorial zone, high biological productivity, and the low-lying, water-logged and slowly subsiding sedimentary basins that allowed

20825-560: The entry and exit channels are organized by the shape of the lophophore. The lophophore captures food particles, especially phytoplankton (tiny photosynthetic organisms), and deliver them to the mouth via the brachial grooves along the bases of the tentacles. The mouth is a tiny slit at the base of the lophophore. Food passes through the mouth, muscular pharynx ("throat") and oesophagus ("gullet"), all of which are lined with cilia and cells that secrete mucus and digestive enzymes . The stomach wall has branched ceca ("pouches") where food

21000-425: The entry channels pause and the tentacles in contact with the lumps move apart to form large gaps and then slowly use their cilia to dump the lumps onto the lining of the mantle. This has its own cilia, which wash the lumps out through the opening between the valves. If the lophophore is clogged, the adductors snap the valves sharply, which creates a "sneeze" that clears the obstructions. In some inarticulate brachiopods

21175-402: The first phase of excretion in this process, and brachiopod metanephridia appear to be used only to emit sperm and ova . The majority of food consumed by brachiopods is digestible, with very little solid waste produced. The cilia of the lophophore can change direction to eject isolated particles of indigestible matter. If the animal encounters larger lumps of undesired matter, the cilia lining

21350-507: The first publication of the APG system in 1998, which proposed a classification of angiosperms up to the level of orders , many sources have preferred to treat ranks higher than orders as informal clades. Where formal ranks have been provided, the traditional divisions listed below have been reduced to a very much lower level, e.g. subclasses . Wolf plants Hepatophyta Liver plants Coniferophyta Cone-bearing plant Phylum Microsporidia

21525-441: The fluid of the coelom, which is circulated through the mantle and driven either by contractions of the lining of the coelom or by beating of its cilia. In some species oxygen is partly carried by the respiratory pigment hemerythrin , which is transported in coelomocyte cells. The maximum oxygen consumption of brachiopods is low, and their minimum requirement is not measurable. Brachiopods also have colorless blood , circulated by

21700-480: The formation of thick and widespread coal formations. During the warm interglacials, smaller coal swamps with plants adapted to the temperate conditions formed on the Siberian craton and the western Australian region of Gondwana. There is ongoing debate as to why this peak in the formation of Earth's coal deposits occurred during the Carboniferous. The first theory, known as the delayed fungal evolution hypothesis,

21875-436: The grounds on which brachiopods were affiliated with deuterostomes: Nielsen views the brachiopods and closely related phoronids as affiliated with the deuterostome pterobranchs because their lophophores are driven by one cilium per cell, while those of bryozoans , which he regards as protostomes, have multiple cilia per cell. However, pterobranchs are hemichordates and probably closely related to echinoderms , and there

22050-414: The hinge. The rest of the space is lined with the mantle lobes , extensions that enclose a water-filled space in which sits the lophophore. The coelom (body cavity) extends into each lobe as a network of canals, which carry nutrients to the edges of the mantle. Relatively new cells in a groove on the edges of the mantle secrete material that extends the periostracum. These cells are gradually displaced to

22225-419: The inarticulate Crania and the articulate Lacazella; they cement the rear of the "pedicle" (ventral) valve to a surface so that the front is slightly inclined up away from the surface. In these brachiopods, the ventral valve lacks a pedicle opening. In a few articulate genera such as Neothyris and Anakinetica , the pedicles wither as the adults grow and finally lie loosely on the surface. In these genera

22400-456: The late Carboniferous. Land arthropods such as arachnids (e.g. trigonotarbids and Pulmonoscorpius ), myriapods (e.g. Arthropleura ) and especially insects (particularly flying insects ) also underwent a major evolutionary radiation during the late Carboniferous. Vast swaths of forests and swamps covered the land, which eventually became the coal beds characteristic of the Carboniferous stratigraphy evident today. The later half of

22575-505: The late Devonian with a short-lived glaciation in the late Famennian through Devonian–Carboniferous boundary, before the Early Tournaisian Warm Interval. Following this, a reduction in atmospheric CO 2 levels, caused by the increased burial of organic matter and widespread ocean anoxia led to climate cooling and glaciation across the south polar region. During the Visean Warm Interval glaciers nearly vanished retreating to

22750-533: The latter three are still in common use in Western Europe. Stages can be defined globally or regionally. For global stratigraphic correlation, the ICS ratify global stages based on a Global Boundary Stratotype Section and Point (GSSP) from a single formation (a stratotype ) identifying the lower boundary of the stage. Only the boundaries of the Carboniferous System and three of the stage bases are defined by global stratotype sections and points because of

22925-432: The majority of a cyclothem sequence occurred during falling sea levels, when rates of erosion were high, meaning they were often periods of non-deposition. Erosion during sea level falls could also result in the full or partial removal of previous cyclothem sequences. Individual cyclothems are generally less than 10 m thick because the speed at which sea level rose gave only limited time for sediments to accumulate. During

23100-558: The mantle's chaetae probably send tactile signals to receptors in the epidermis of the mantle. Many brachiopods close their valves if shadows appear above them, but the cells responsible for this are unknown. Some brachiopods have statocysts , which detect changes in the animals' position. Lifespans range from 3 to over 30 years. Adults of most species are of one sex throughout their lives. The gonads are masses of developing gametes ( ova or sperm ), and most species have four gonads, two in each valve. Those of articulates lie in

23275-683: The mid Carboniferous, the South American sector of Gondwana collided obliquely with Laurussia's southern margin resulting in the Ouachita orogeny. The major strike-slip faulting that occurred between Laurussia and Gondwana extended eastwards into the Appalachian Mountains where early deformation in the Alleghanian orogeny was predominantly strike-slip. As the West African sector of Gondwana collided with Laurussia during

23450-536: The mineralized layers are perforated by tiny open canals of living tissue, extensions of the mantle called caeca, which almost reach the outside of the primary layer. These shells can contain half of the animal's living tissue. Impunctate shells are solid without any tissue inside them. Pseudopunctate shells have tubercles formed from deformations unfurling along calcite rods. They are only known from fossil forms, and were originally mistaken for calcified punctate structures. Lingulids and discinids, which have pedicles, have

23625-478: The modern "system" names, it was coined by geologists William Conybeare and William Phillips in 1822, based on a study of the British rock succession. Carboniferous is the period during which both terrestrial animal and land plant life was well established. Stegocephalia (four-limbed vertebrates including true tetrapods ), whose forerunners ( tetrapodomorphs ) had evolved from lobe-finned fish during

23800-456: The near worldwide distribution of marine faunas and so allowing widespread correlations using marine biostratigraphy . However, there are few Mississippian volcanic rocks , and so obtaining radiometric dates is difficult. The Tournaisian Stage is named after the Belgian city of Tournai . It was introduced in scientific literature by Belgian geologist André Dumont in 1832. The GSSP for

23975-411: The next. The Catalogue of Life includes Rhodophyta and Glaucophyta in kingdom Plantae, but other systems consider these phyla part of Protista. In addition, less popular classification schemes unite Ochrophyta and Pseudofungi under one phylum, Gyrista , and all alveolates except ciliates in one phylum Myzozoa , later lowered in rank and included in a paraphyletic phylum Miozoa . Even within

24150-515: The northwestern Gondwana margin, were affected by the subduction of the Rheic Ocean. However, they lay to west of the Ouachita orogeny and were not impacted by continental collision but became part of the active margin of the Pacific. The Moroccan margin was affected by periods of widespread dextral strike-slip deformation, magmatism and metamorphism associated with the Variscan orogeny. Towards

24325-419: The opening of the burrow to feed, and to retract the shell when disturbed. A lingulid moves its body up and down the top two-thirds of the burrow, while the remaining third is occupied only by the pedicle, with a bulb on the end that builds a "concrete" anchor. However, the pedicles of the order Discinida are short and attach to hard surfaces. The pedicle of articulate brachiopods has no coelom, and its homology

24500-539: The other hand, the highly parasitic phylum Mesozoa was divided into two phyla ( Orthonectida and Rhombozoa ) when it was discovered the Orthonectida are probably deuterostomes and the Rhombozoa protostomes . This changeability of phyla has led some biologists to call for the concept of a phylum to be abandoned in favour of placing taxa in clades without any formal ranking of group size. A definition of

24675-537: The other hand, warmer periods, such much of the Silurian , created smaller difference in temperatures, and all seas at the low to middle latitudes were colonized by the same few brachiopod species. From about the 1940s to the 1990s, family trees based on embryological and morphological features placed brachiopods among or as a sister group to the deuterostomes . a super-phylum that includes chordates and echinoderms . Closer examination has found difficulties in

24850-549: The other protostome super-phylum Ecdysozoa , whose members include arthropods . This conclusion is unanimous among molecular phylogeny studies that use a wide selection of genes: rDNA , Hox genes , mitochondrial protein genes, single nuclear protein genes and sets of nuclear protein genes. Some combined studies in 2000 and 2001, using both molecular and morphological data, support brachiopods as Lophotrochozoa, while others in 1998 and 2004 concluded that brachiopods were deuterostomes. Phylum (biology) In biology ,

25025-408: The other shell. Hemiperipheral growth, found in lingulids, is similar to mixoperipheral growth but occurs in mostly a flat plate with the shell growing forwards and outwards. Brachiopods, as with molluscs , have an epithelial mantle which secretes and lines the shell, and encloses the internal organs. The brachiopod body occupies only about one-third of the internal space inside the shell, nearest

25200-426: The peat mires. As fully marine conditions were established, limestones succeeded these marginal marine deposits. The limestones were in turn overlain by deep water black shales as maximum sea levels were reached. Ideally, this sequence would be reversed as sea levels began to fall again; however, sea level falls tend to be protracted, whilst sea level rises are rapid, ice sheets grow slowly but melt quickly. Therefore,

25375-482: The pedicle valve, either through a notch in the hinge or, in species where the pedicle valve is longer than the brachial, from a hole where the pedicle valve doubles back to touch the brachial valve. Some species stand with the front end upwards, while others lie horizontal with the pedicle valve uppermost. Some early brachiopods—for example strophomenates , kutorginates and obolellates —do not attach using their pedicle, but with an entirely different structure known as

25550-516: The period experienced glaciations , low sea level, and 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. Atmospheric oxygen levels, originally thought to be consistently higher than today throughout the Carboniferous, have been shown to be more variable, increasing from low levels at

25725-405: The plankton for only a few days before leaving the water column upon metamorphosing . While traditional classification of brachiopods separate them into distinct inarticulate and articulate groups, two approaches appeared in the 1990s. One approach groups the inarticulate Craniida with articulate brachiopods, since both use layers of calcareous minerals their shell; the other approach considers

25900-640: The plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships among phyla within larger clades like Ecdysozoa and Embryophyta . The term phylum was coined in 1866 by Ernst Haeckel from the Greek phylon ( φῦλον , "race, stock"), related to phyle ( φυλή , "tribe, clan"). Haeckel noted that species constantly evolved into new species that seemed to retain few consistent features among themselves and therefore few features that distinguished them as

26075-451: The population of Coptothyrus adamsi useful as a measure of environmental conditions around an oil terminal being built in Russia on the shore of the Sea of Japan . Brachiopods are the state fossil of the U.S. state of Kentucky . Over 12,000 fossil species are recognized, grouped into over 5,000  genera . While the largest modern brachiopods are 100 millimetres (3.9 in) long,

26250-491: The preceding Devonian period, became pentadactylous during the Carboniferous. The period is sometimes called the Age of Amphibians because of the diversification of early amphibians such as the temnospondyls , which became dominant land vertebrates, as well as the first appearance of amniotes including synapsids (the clade to which modern mammals belong) and sauropsids (which include modern reptiles and birds) during

26425-464: The presence of Siphonodella sulcata below the boundary, and the presence of Siphonodella praesulcata and Siphonodella sulcata together above a local unconformity . This means the evolution of one species to the other, the definition of the boundary, is not seen at the La Serre site making precise correlation difficult. The Viséan Stage was introduced by André Dumont in 1832 and is named after

26600-892: The proto-Andes in Bolivia and western Argentina and the Pan-African mountain ranges in southeastern Brazil and southwest Africa. The main phase of the LPIA (c. 335-290 Ma) began in the late Visean, as the climate cooled and atmospheric CO 2 levels dropped. Its onset was accompanied by a global fall in sea level and widespread multimillion-year unconformities. This main phase consisted of a series of discrete several million-year-long glacial periods during which ice expanded out from up to 30 ice centres that stretched across mid- to high latitudes of Gondwana in eastern Australia, northwestern Argentina, southern Brazil, and central and Southern Africa. Isotope records indicate this drop in CO 2 levels

26775-420: The reduction of some brachial nerves. The tentacles bear cilia (fine mobile hairs) on their edges and along the center. The beating of the outer cilia drives a water current from the tips of the tentacles to their bases, where it exits. Food particles that collide with the tentacles are trapped by mucus , and the cilia down the middle drive this mixture to the base of the tentacles. A brachial groove runs round

26950-475: The relationships between groups. So phyla can be merged or split if it becomes apparent that they are related to one another or not. For example, the bearded worms were described as a new phylum (the Pogonophora) in the middle of the 20th century, but molecular work almost half a century later found them to be a group of annelids , so the phyla were merged (the bearded worms are now an annelid family ). On

27125-640: The sea. Cyclothem lithologies vary from mudrock and carbonate-dominated to coarse siliciclastic sediment-dominated sequences depending on the paleo-topography, climate and supply of sediments to the shelf. The main period of cyclothem deposition occurred during the Late Paleozoic Ice Age from the Late Mississippian to early Permian, when the waxing and waning of ice sheets led to rapid changes in eustatic sea level . The growth of ice sheets led global sea levels to fall as water

27300-464: The shell becomes heavier, the juvenile sinks to the bottom and becomes a sessile adult. The larvae of articulate species (Craniiformea and Rhynchonelliformea) are lecithotrophic (non-feeding) and live only on yolk , and remain among the plankton for only a few days. The Rhynchonelliformea larvae has three larval lobes, unlike the Craniiformea which only have two larval lobes. This type of larva has

27475-418: The shell or may help in respiration . Experiments show that a brachiopod's oxygen consumption drops if petroleum jelly is smeared on the shell, clogging the diverticula. Like bryozoans and phoronids , brachiopods have a lophophore, a crown of tentacles whose cilia (fine hairs) create a water current that enables them to filter food particles out of the water. However a bryozoan or phoronid lophophore

27650-402: The shells are thickened and shaped so that the opening of the gaping valves is kept free of the sediment. Pedicles of inarticulate species are extensions of the main coelom, which houses the internal organs. A layer of longitudinal muscles lines the epidermis of the pedicle. Members of the order Lingulida have long pedicles, which they use to burrow into soft substrates, to raise the shell to

27825-494: The shore of the Sea of Japan . The word "brachiopod" is formed from the Ancient Greek words brachion ("arm") and podos ("foot"). They are often known as " lamp shells ", since the curved shells of the class Terebratulida resemble pottery oil-lamps. Modern brachiopods range from 1 to 100 millimetres (0.039 to 3.937 in) long, and most species are about 10 to 30 millimetres (0.39 to 1.18 in). Magellania venosa

28000-504: The suture between Kazakhstania and Tarim. A continental magmatic arc above a south-dipping subduction zone lay along the northern North China margin, consuming the Paleoasian Ocean. Northward subduction of the Paleo-Tethys beneath the southern margins of North China and Tarim continued during the Carboniferous, with the South Qinling block accreted to North China during the mid to late Carboniferous. No sediments are preserved from

28175-585: The taxonomy of brachiopods down to the order level, including extinct groups, which make up the majority of species. Extinct groups are indicated with a (†) symbol: Brachiopods are an entirely marine phylum, with no known freshwater species. Most species avoid locations with strong currents or waves, and typical sites include rocky overhangs, crevices and caves, steep slopes of continental shelves , and in deep ocean floors. However, some articulate species attach to kelp or in exceptionally sheltered sites in intertidal zones . The smallest living brachiopod, Gwynia ,

28350-410: The thick accumulation of peat were sufficient to account for the peak in coal formation. During the Carboniferous, there was an increased rate in tectonic plate movements as the supercontinent Pangea assembled. The continents themselves formed a near circle around the opening Paleo-Tethys Ocean, with the massive Panthalassic Ocean beyond. Gondwana covered the south polar region. To its northwest

28525-434: The two being mirror images of each other. The formation of brachiopod shells during ontogeny builds on a set of conserved genes, including homeobox genes, that are also used to form the shells of molluscs. The brachial valve is usually smaller and bears brachia ("arms") on its inner surface. These brachia are the origin of the phylum's name, and support the lophophore , used for feeding and respiration . The pedicle valve

28700-417: The underside of the mantle by more recent cells in the groove, and switch to secreting the mineralized material of the shell valves. In other words, on the edge of the valve the periostracum is extended first, and then reinforced by extension of the mineralized layers under the periostracum. In most species the edge of the mantle also bears movable bristles, often called chaetae or setae , that may help defend

28875-568: The uplift and erosion of the more mafic basement rocks of the Central Pangea Mountains at this time, CO 2 levels dropped as low as 175 ppm and remained under 400 ppm for 10 Ma. Temperatures across the Carboniferous reflect the phases of the LPIA. At the extremes, during the Permo-Carboniferous Glacial Maximum (299-293 Ma) the global average temperature (GAT) was c. 13 °C (55 °F),

29050-563: The way from the Paleozoic to modern times, but bivalves increased faster; the Permian–Triassic extinction was moderately severe for bivalves but devastating for brachiopods, so that brachiopods for the first time were less diverse than bivalves and their diversity after the Permian increased from a very low base; there is no evidence that bivalves out-competed brachiopods, and short-term increases or decreases for both groups appeared synchronously. In 2007 Knoll and Bambach concluded that brachiopods were one of several groups that were most vulnerable to

29225-492: Was Laurussia. These two continents slowly collided to form the core of Pangea. To the north of Laurussia lay Siberia and Amuria . To the east of Siberia, Kazakhstania , North China and South China formed the northern margin of the Paleo-Tethys, with Annamia laying to the south. The Central Pangean Mountains were formed during the Variscan - Alleghanian - Ouachita orogeny. Today their remains stretch over 10,000 km from

29400-506: Was complete by the Tournaisian, but subduction of the Ural Ocean between Kazakhstania and Laurussia continued until the Bashkirian when the ocean finally closed and continental collision began. Significant strike-slip movement along this zone indicates the collision was oblique. Deformation continued into the Permian and during the late Carboniferous and Permian the region was extensively intruded by granites . The Laurussian continent

29575-516: Was formed by the collision between Laurentia , Baltica and Avalonia during the Devonian. At the beginning of the Carboniferous, some models show it at the equator, whilst others place it further south. In either case, the continent drifted northwards, reaching low latitudes in the northern hemisphere by the end of the Period. The Central Pangean Mountain drew in moist air from the Paleo-Tethys Ocean resulting in heavy precipitation and

29750-402: Was hypothesized earlier, but should be included in the broad group Protostomia , in a subgroup now called Lophotrochozoa . Although their adult morphology seems rather different, the nucleotide sequence of the 18S rRNA indicates that the phoronids (horseshoe worms) are the closest relatives of the inarticulate brachiopods, more so than articulate brachiopods. For now, the weight of evidence

29925-646: Was lock away in glaciers. Falling sea levels exposed large tracts of the continental shelves across which river systems eroded channels and valleys and vegetation broke down the surface to form soils . The non-marine sediments deposited on this erosional surface form the base of the cyclothem. As sea levels began to rise, the rivers flowed through increasingly water-logged landscapes of swamps and lakes. Peat mires developed in these wet and oxygen-poor conditions, leading to coal formation. With continuing sea level rise, coastlines migrated landward and deltas , lagoons and esturaries developed; their sediments deposited over

30100-433: Was made of organophosphatic compounds while that of Halkieria was made of calcite . However, fossils of a new tommotiid, Eccentrotheca , showed an assembled mail coat that formed a tube, which would indicate a sessile animal rather than a creeping slug-like one. Eccentrotheca' s organophosphatic tube resembled that of phoronids , sessile animals that feed by lophophores and are regarded either very close relatives or

30275-578: Was proposed by Russian stratigrapher Sofia Semikhatova in 1934. It was named after Bashkiria , the then Russian name of the republic of Bashkortostan in the southern Ural Mountains of Russia. The GSSP for the base of the Pennsylvanian Subsystem and Bashkirian Stage is located at Arrow Canyon in Nevada , US and was ratified in 1996. It is defined by the first appearance of the conodont Declinognathodus noduliferus . Arrow Canyon lay in

30450-405: Was scarce. In waters where food is scarce, the snail Capulus ungaricus steals food from bivalves, snails, tube worms, and brachiopods. Among brachiopods only the lingulids have been fished commercially, and only on a very small scale. It is mostly the fleshy pedicle that is eaten. Brachiopods seldom settle on artificial surfaces, probably because they are vulnerable to pollution. This may make

30625-407: Was triggered by tectonic factors with increased weathering of the growing Central Pangean Mountains and the influence of the mountains on precipitation and surface water flow. Closure of the oceanic gateway between the Rheic and Tethys oceans in the early Bashkirian also contributed to climate cooling by changing ocean circulation and heat flow patterns. Warmer periods with reduced ice volume within

#669330