Brachiopods are rare in modern oceans, but were very common in the past (only 325 living species but more than 12,000 fossil species). Lifespans range from three to over thirty years. [7], Lifespans range from 3 to over 30 years. Lingula found near Ozamis City, Philippines. [28] [23], 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 Devonian brachiopod Tylothyris from the Milwaukee Formation, Milwaukee County, Wisconsin The origin of the brachiopods is uncertain; they either arose from reduction of a multi-plated tubular organism, or from the folding of a slug-like organism with a protective shell on either end. Brachiopods feed by filtering tiny food particles from seawater. Complete trilobite skeletons are relatively rare, and were probably preserved when the sea floor was buried by mud during major storms. The smallest living brachiopod, Gwynia, is only about 1 millimetre (0.039 in) long, and lives in between gravel. In 1883, Othniel Marsh discovered a Brachiosaurus skull near Garden Park, Colora… The front limbs being longer than hind limbs gave the … On the other hand, articulate brachiopods have produced major diversifications, and suffered severe mass extinctions[41]—but the articulate Rhynchonellida and Terebratulida, the most diverse present-day groups, appeared at the start of the Ordovician and Carboniferous respectively. Lower Carboniferous of Wooster, Ohio. It is sometimes associated with a fringing plate, the colleplax. The other subphylum Rhynchonelliformea contains only one class, which is subdivided in the orders Rhynchonellida, Terebratulida and Thecideida. Their front limbs were longer than hind limbs. [61], The phoronids feed with a lophophore, burrow or encrust on surfaces, and build three-layered tubes made of polysaccharide, possibly chitin, mixed with particles with seabed material. These cells are gradually displaced to the underside of the mantle by more recent cells in the groove, and switch to secreting the mineralized material of the shell valves. [7] Members of the order Lingulida have long pedicles, which they use to burrow into soft substrates, to raise the shell to the opening of the burrow to feed, and to retract the shell when disturbed. [14], 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. The word "brachiopod" is formed from the Ancient Greek words brachion ("arm") and podos ("foot"). The mouth is at the base of the lophophore. The Terebratulida are punctate brachiopods; their shell is perforated by tiny open canals of living tissue, extensions of the mantle called caeca, which almost reach the outside of the primary layer. 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. Modern day brachiopods do still exist in the form of lingula. About 60 percent of brachiopods live in shallow water (less than 100 fathoms—about 180 metres [600 feet]) on the shelf areas around the continents. Today they live in every ocean and major benthic (ocean-bottom) habitat. All but a few brachiopods fall into two basic types, the rhynchonelliform (or articulate) brachiopods and the lingulate (or inarticulate) brachiopods. Eccentrotheca's organophosphatic tube resembled that of phoronids,[45] sessile animals that feed by lophophores and are regarded either very close relatives or a sub-group of brachiopods. If the animal encounters larger lumps of undesired matter, the cilia lining 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. Brachiopods are suspension feeders, which means that they extract food (plankton, particles of dead organic matter, etc.) Depsite their relative obscurity today, brachiopods have a long and rich paleontological history. However, after the Permian–Triassic extinction event, brachiopods recovered only a third of their former diversity. The Devonian Period was a time of extensive reef building in the shallow water that surrounded each continent and separated Gondwana from Euramerica. Spiriferids and brachiopods in general, hit the height of diversity during the Devonian Period. [7] Brachiopods have metanephridia, used by many phyla to excrete ammonia and other dissolved wastes. 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). The oldest brachiopods can be found in rocks of early Cambrian age (about 530 million years old). More than 35 percent occupy waters deeper than 100 fathoms, and a few live in the abyss down to more than 6,000 metres (about 20,000 feet). However, there are a few species that can live in depths exceeding 5000m. [7], The cell division in the embryo 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). [2] 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 was scarce. Rhynchonelliform brachiopods have shells made calcium carbonate and interlocking pegs (teeth) and sockets that form a hinge between the valves. You likely haven't seen one because the vast majority inhabit areas not frequented by human activity. A Carboniferous brachiopod Neospirifer condor, from Bolivia. Two major groups are recognized, articulate and inarticulate. [27][7] While some animals develop the mouth and anus by deepening the blastopore, a "dent" in the surface of the early embryo, the blastopore of brachiopods closes up, and their mouth and anus develop from new openings. 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 word "articulate" is used to describe the tooth-and-groove features of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. Although many rhynchonelliform brachiopods are held in place by a pedicle, some extinct forms lost the pedicle and lay freely on the sea bottom. They are unable to move. The valves are termed brachial and pedicle. [46] 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. [47], At their peak in the Paleozoic, the brachiopods were among the most abundant filter-feeders and reef-builders,[48] and occupied other ecological niches, including swimming in the jet-propulsion style of scallops. [19] However, the pedicles of the order Discinida are short and attach to hard surfaces. Oxygen seems to be distributed by 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. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. [40], Since 1991 Claus Nielsen has proposed a hypothesis about the development of brachiopods, adapted in 2003 by Cohen and colleagues as a hypothesis about the earliest evolution of brachiopods. [7] [7] 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. [4] Other lineages have arisen and then become extinct, sometimes during severe mass extinctions. They were extremely diverse during the Devonian period and later went extinct during the Jurassic period. (405) 325-4712, The Sam Noble Museum: [email protected], Paleobotany, Micropaleontology & Mineralogy. Brachiopods now live mainly in cold water and low light. [2] However, after the Permian–Triassic extinction event, informally known as the "Great Dying",[48] brachiopods recovered only a third of their former diversity. In some species oxygen is partly carried by the respiratory pigment hemerythrin, which is transported in coelomocyte cells. The first Brachiosaurus (B. altithorax) fossil was found in Grand River Valley in western Colorado in 1900. Ripe gametes (ova or sperm) float from the gonads into the main coelom and then exit into the mantle cavity. [14] In most species the edge of the mantle also bears movable bristles, often called chaetae or setae, that may help defend the animals and may act as sensors. Most brachiopods live in relatively shallow marine water, up to about 650 feet (200 m), but some species have been found at depths of more than a mile. Traditionally they have been regarded as a separate phylum, but increasingly detailed molecular phylogeny studies between 1997 and 2000 have concluded that phoronids are a sub-group of brachiopods. [2] Rhynchonelliforms (Articulata excluding Craniida), whose larvae consume only their yolks and settle and develop quickly, specialize in specific areas and form dense populations that can reach thousands per meter. Brachiopods are virtually defenceless and their shell, enclosing the animal’s organs, is the only protection against predators. However, in 1980, Gould and Calloway produced a statistical analysis that concluded that: both brachiopods and bivalves increased all 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. A layer of longitudinal muscles lines the epidermis of the pedicle. The paleontologist who discovered this partial skeleton, Elmer Riggs, named this new find Brachiosaurus in 1903. Entoprocts use a similar-looking crown of tentacles, but it 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. In the "traditional" classification, the Articulata have toothed hinges between the valves, while the hinges of the Inarticulata are held together only by muscles. [2], Genetic analysis performed since the 1990s has extended the understanding of the relationship between different organisms. The body is covered in a shell that is made of two halves (valves) that are held in place by muscles. [14], The tentacles bear cilia (fine mobile hairs) on their edges and along the center. Barroisella, a lingulid from the Middle Devonian of Wisconsin. [14] As the shell becomes heavier, the juvenile sinks to the bottom and becomes a sessile adult. Hence some brachiopod taxonomists believe it is premature to define higher levels of classification such as order, and recommend instead a bottom-up approach that identifies genera and then groups these into intermediate groups. Brachiopod fossils have been useful indicators of climate changes during the Paleozoic. The gonads are masses of developing gametes (ova or sperm), and most species have four gonads, two in each valve. Although not directly connected to sensory neurons, the mantle's chaetae probably send tactile signals to receptors in the epidermis of the mantle. In a typical brachiopod a stalk-like pedicle projects from an opening in one of the valves near the hinges, known as the pedicle valve, keeping the animal anchored to the seabed but clear of silt that would obstruct the opening. live in relatively shallow coastal waters. The pedicle valve has on its inner surface the attachment to the stalk-like pedicle by which most brachiopods attach themselves to the substrate. The specimen is 7 cm across. [3] They are often known as "lamp shells", since the curved shells of the class Terebratulida resemble pottery oil-lamps.[2]. In addition to the traditional classification of brachiopods into inarticulate and articulate, two approaches appeared in the 1990s: one approach groups the inarticulate Craniida with articulate brachiopods, since both use the same material in the mineral layers of their shell; the other approach makes the Craniida a third group, as their outer organic layer is different from that in either of the other two. If one were to judge by the superb col- lection of recent brachiopods in the U. S. National Museum he might be led to believe that shallow \/ater brachiopcds were more abundant as to individuals than deep water brachiopods. These shells can contain half of the animal's living tissue. [7] The larvae of articulate species live only on yolk, and remain among the plankton for only a few days. [2], 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. Adult inarticulates have only the lower ganglion. They were much more abundant in seas of the Silurian Period. HOW DID BRACHIOPODS LIVE? [60] However, pterobranchs are hemichordates and probably closely related to echinoderms, and there 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. Brachiopods have a shell made of two halves. Although their adult morphology seems rather different, the nucleotid sequence of the 18S rRNA indicates that the horseshoe worms are the closest relatives of the inarticulate brachiopods rather than articulate brachiopods. It was suggested in 2003 that brachiopods had evolved from an ancestor similar to Halkieria, 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. [34][35], Brachiopods live only in the sea. [46] However, an analysis in 2005 concluded that phoronids are a sub-group of bryozoans. Fossil spiriferids first appear in the Ordovician period, as illustrated in the fossil range chart for brachiopods. 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 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). They are unable to move. Brachiopods are benthic (bottom dwelling), marine (ocean), bivalves (having two shells). Number of families 20. In lingulids the entrance and exit channels are formed by groups of chaetae that function as funnels. They were represented in the Ordovician (about 488 million to 444 million years ago) but decreased thereafter. Instead, their mouths contained bony structures used to crush or shear prey. [14] The wastes produced by metabolism are broken into ammonia, which is eliminated by diffusion through the mantle and lophophore. Interestingly, though Riggs may have identified the new dinosaur species, his find was technically not the first discovery of a Brachiosaurus fossil. The pedicle emerges from 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. Lineages of brachiopods that have both fossil and extant taxa appeared in the early Cambrian, Ordovician, and Carboniferous periods, respectively. A study in 2007 concluded 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. Many brachiopods close their valves if shadows appear above them, but the cells responsible for this are unknown. [50], Brachiopod fossils have been useful indicators of climate changes during the Paleozoic era. This order belongs to the class of brachiopods called Articulata, all of which have an articulated hinge. De Rosa (2001) cites the following examples of brachiopods as close to deuterostomes: sfn error: no target: CITEREFValentine:_Cleavage_patterns (, Zvyagintsev etc: Brachio fouling & (2007), Shorter Oxford English Dictionary & (2002), "Braquiópodos discínidos (Lingulida, Discinoidea) de la Formación Ixtaltepec, Carbonífero del área de Santiago Ixtaltepec, Oaxaca", Marine Species Identification Portal : Brachiopoda of the North Sea, "Apatite Ca5(PO4, CO3)3(F, Cl, OH) Hexagonal", Parkinson etc: Brachiopod shells & (2005), Williams etc: Suprafamilial Classif & (2000), "Molecular evidence that phoronids are a subtaxon of brachiopods (Brachiopoda: Phoronata) and that genetic divergence of metazoan phyla began long before the early Cambrian", Ushatinskaya: Earliest brachiopods & (2008), Balthasar etc: Brachios stem Phoronids & (2009), Conway Morris etc: Articulated Halkieriids & (1995), "The scleritome of Paterimitra: an Early Cambrian stem group brachiopod from South Australia", "Molecular Data Indicate the Protostome Affinity of Brachiopods", Helmkampf etc: Lophotrochozoa concept & (2008), Peterson etc: Combined phylogeny & (2001), Wood etc: Phylactolaemate Phylog & (2005), Bourlat etc: Close to Nemertines & (2008), 10.1666/0022-3360(2001)075<1109:GOTPPA>2.0.CO;2, "Monophyly of brachiopods and phoronids: reconciliation of molecular evidence with Linnaean classification (the subphylum Phoroniformea nov.)", "The brachiopod fold: a neglected body plan hypothesis", "Triploblastic relationships with emphasis on the acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: a combined approach of 18S rDNA sequences and morphology", "Phylogenomic analyses of lophophorates (brachiopods, phoronids and bryozoans) confirm the Lophotrochozoa concept", "Paleophysiology and end-Permian mass extinction", "The Phylogenetic Position of Entoprocta, Ectoprocta, Phoronida, and Brachiopoda", "Downstream collecting in ciliary suspension feeders: the catch-up principle", "Cleavage patterns and the topology of the metazoan tree of life", "Outline of Suprafamilial Classification and Authorship", "Earliest rhynchonelliform brachiopod parasite from the Late Ordovician of northern Estonia (Baltica)", Information from the Kansas Geological Survey, Archaeal Richmond Mine acidophilic nanoorganisms, https://en.wikipedia.org/w/index.php?title=Brachiopod&oldid=997295478, Short description is different from Wikidata, Articles using Template:Background color with invalid colour combination, Creative Commons Attribution-ShareAlike License. [7], The majority of food consumed by brachiopods is digestible, with very little solid waste produced. Traditionally, brachiopods have been regarded as members of, or as a sister group to, the deuterostomes, a superphylum that includes chordates and echinoderms. [52][55] This conclusion is unanimous among molecular phylogeny studies, which use a wide selection of genes: rDNA, Hox genes, mitochondrial protein genes, single nuclear protein genes and sets of nuclear protein genes. What era did Lingula live? [7], Nutrients are transported throughout the coelom, including the mantle lobes, by cilia. Muscles at the rear of the body can straighten, bend or even rotate the pedicle. 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. Brachiopods live on the ocean floor. Both fossils and extant species have limitations that make it difficult to produce a comprehensive classification of brachiopods based on morphology. However they did not become abundant until the Devonian. The teeth are in one valve (the pedicle or ventral valve) and the sockets are in the other (the brachial or dorsal valve). [18], Most modern species attach to hard surfaces by means of a cylindrical pedicle ("stalk"), an extension of the body wall. [7]The maximum oxygen consumption of brachiopods is low, and their minimum requirement is not measurable. [7] Experiments show that a brachiopod's oxygen consumption drops if petroleum jelly is smeared on the shell, clogging the diverticula. [65][66] Another study in 2008 also concluded that brachiopods are closely related to nemertines, casting doubt on the idea that brachiopods are part of a clade Lophophorata of lophophore-feeding animals within the lophotrochozoans.[61]. Members of some genera have survived for a year in aquaria without food. [7], The larvae of inarticulates swim as plankton for months[2] and are like 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[7]—except that Craniids have no pedicle. Studies of modern-day brachiopods show us the soft tissues and other internal structures of the animal. These variations in growth often form growth lines in the shells. In a prominent example, the traditional view that brachiopods were ecologically dominant over bivalves in the Paleozoic has been disputed on both taphonomic and metabolic grounds. [14] 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. This has a chitinous cuticle (non-cellular "skin") and protrudes through an opening in the hinge. That accomplished, they sat where they were, filtered water, grew, reproduced, and did little else. The beating of the outer cilia drives a water current from the tips of the tentacles to their bases, where it exits. Articulated brachiopods have an outermost periostracum made of proteins, a "primary layer" of calcite (a form of calcium carbonate) under that, and innermost a mixture of proteins and calcite. [6] Each has two valves (shell sections) which cover the dorsal and ventral surface of the animal, unlike bivalve molluscs whose shells cover the lateral surfaces. Among brachiopods, only the lingulids (Lingula sp. The Sam Noble Museum at The University of Oklahoma inspires minds to understand the world through collection-based research, interpretation, and education. [7] The coelom extends into each lobe as a network of canals, which carry nutrients to the edges of the mantle. Norman, OK 73072-7029 [13], Brachiopods, as with molluscs, have an epithelial mantle that secretes and lines the shell, and encloses the internal organs. This "brachiopod fold" hypothesis suggests that brachiopods evolved from an ancestor similar to Halkieria,[23] a slug-like animal with "chain mail" on its back and a shell at the front and rear end. There are about 300 living species of brachiopods. The plane of symmetry passes through the valves, rather than between them. [49] In 2007 Knoll and Bambach concluded that brachiopods were one of several groups that were most vulnerable to the Permian–Triassic extinction, as all had calcareous hard parts (made of calcium carbonate) and had low metabolic rates and weak respiratory systems. [17] A brachial groove runs round the bases of the tentacles, and its own cilia pass food along the groove towards the mouth. Brachiopods also have colorless blood, circulated by a muscular heart lying in the dorsal part of the body above the stomach. Many other types of brachipods, like those shown above, are found in the Pennsylvanian rocks of Kentucky. A Devonian spiriferid brachiopod from Ohio that served as a host substrate for a colony of hederellids. The brachiopod body occupies only about one-third of the internal space inside the shell, nearest the hinge. They are rare today but during the Paleozoic Era they dominated the sea floors. [7] 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. Some have spines that serve as anchors. 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. However, articulate brachiopods of genus Chlidonophora use a branched pedicle to anchor in sediment. Trilobites could roll up into a ball for protection by bending the thorax and bringing the tail underneath the head. Many species are deep water specialists, though some species such as the burrow living Lingula sp. The planktonic larvae of articulate species do not resemble the adults, but rather look like blobs with yolk sacs, and remain among the plankton for only a few days before leaving the water column upon metamorphosing. Most brachiopods became extinct about 250 million years ago during the P-T Extinction period. [37] In waters where food is scarce, the snail Capulus ungaricus steals food from bivalves, snails, tube worms, and brachiopods. [46][61] However, in 2008 two analyses found that brachiopods' closest lophotrochozoan relatives were nemertines. On the other hand, inarticulate brachiopods, whose larva swim for up to a month before settling, have wide ranges. On 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. Many species were anchored to the sea floor by a muscular stalk (pedicle) which emerged through a hole in the pedicle valve (pedicle opening). However, the majority of brachiopods lived and still live in shallow water and intertidal zones. [7] In other brachiopods the entry and exit channels are organized by the shape of the lophophore. Most brachiopods live attached to the sea floor by a fleshy stalk that is an extension of the soft body. Most brachiopods tolerate only normal marine salinity, but a few species, such as the ligulides, can live in brackish salinities. Brachiosaurus is one of the rarer sauropods of the Morrison Formation. Most brachiopods live on the shallow continental shelf. The Fossil RecordTheir first appearance in the fossil record is in the Ordovician Period. The blood circulation seems not to be completely closed, and the coelomic fluid and blood must mix to a degree. Although relatively rare, modern brachiopods occupy a variety of sea-bed habitats ranging from the tropics to the cold waters of the Arctic and, especially, the Antarctic. [7] 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. Rafinesquina is a brachiopod, which are small filter-feeding sea creatures that have a two-part shell with a hinge at the back. 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 resemble a hand with the fingers splayed. [2], Pedicles of inarticulate species are extensions of the main coelom, which houses the internal organs. Members of the discinoid genus Pelagodiscus have a cosmopolitan distribution. [39] Brachiopods seldom settle on artificial surfaces, probably because they are vulnerable to pollution. The chart below depicts the geological periods during which trilobites existed. [52][53] a super-phylum that includes chordates and echinoderms. Articulata (Articulate lampshells) Phylum Brachiopoda. Some forms contain calcium phosphate and others have calcium carbonate. [7] Some articulate brachiopods also have a brachidium, a calcareous support for the lophophore attached to the inside of the brachial valve. In some brachiopods groups of chaetae help to channel the flow of water into and out of the mantle cavity. 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 a subgroup of brachiopods, while the other tommotiid bore two symmetrical plates that might be an early form of brachiopod valves. Brachiopods live in all parts of the sea, mostly as sessile animals attached to the substrate by a fleshy or horny pedicel. [2], The rates of metabolism of Brachiopoda are between one third and one tenth of those of bivalves. [7] However, some genera such as the inarticulate Crania and the articulate Lacazella have no pedicle, and cement the rear of the "pedicle" valve to a surface so that the front is slightly inclined up away from the surface. Petrocrania brachiopods attached to a strophomenid brachiopod; Upper Ordovician of southeastern Indiana.
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