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49-401: Monoplacophora / ˌ m ɒ n oʊ p l ə ˈ k ɒ f ər ə / , meaning "bearing one plate", is a polyphyletic superclass of molluscs with a cap-like shell, inhabiting deep sea environments. Extant representatives were not recognized as such until 1952; previously they were known only from the fossil record , and were thought to have become extinct 375 million years ago. Although

98-406: A cartilaginous gill arch from which projects a cartilaginous gill ray . This gill ray is the support for the sheet-like interbranchial septum , which the individual lamellae of the gills lie on either side of. The base of the arch may also support gill rakers , projections into the pharyngeal cavity that help to prevent large pieces of debris from damaging the delicate gills. A smaller opening,

147-453: A node-based clade definition , for example, could be "All descendants of the last common ancestor of species X and Y". On the other hand, polyphyletic groups can be delimited as a conjunction of several clades, for example "the flying vertebrates consist of the bat, bird, and pterosaur clades". From a practical perspective, grouping species monophyletically facilitates prediction far more than does polyphyletic grouping. For example, classifying

196-572: A pseudometamerism of bilaterally symmetrical repeated organs and muscles. The extant members of the class live only in the deep ocean (the abyssal zone , the continental shelf , and the continental slope ) at depths below 180 metres (590 ft). Cambrian forms predominately lived in shallow seas, whereas later Paleozoic forms are more commonly found in deeper waters with soft, muddy sea floors. Although superficially resembling limpets when viewed dorsally, monoplacophorans are not anatomically similar to gastropods . Some similarities are shared with

245-464: A radula , a defining characteristic of the mollusca. Tentacles are situated behind the mouth. They also have a cone-shaped stomach with a single crystalline style though no gastric shield . The intestines are long and make between four and six loops before reaching the posteriorly-positioned anus. Monoplacophorans also have oesophageal pouches . The sexes are separate with any given animal having two pair of either ovaries or testes connected to either

294-399: A bony operculum. The great majority of bony fish species have five pairs of gills, although a few have lost some over the course of evolution. The operculum can be important in adjusting the pressure of water inside of the pharynx to allow proper ventilation of the gills, so bony fish do not have to rely on ram ventilation (and hence near constant motion) to breathe. Valves inside the mouth keep

343-452: A classification is discouraged. Monophyletic groups (that is, clades ) are considered by these schools of thought to be the only valid groupings of organisms because they are diagnosed ("defined", in common parlance) on the basis of synapomorphies , while paraphyletic or polyphyletic groups are not. From the perspective of ancestry, clades are simple to define in purely phylogenetic terms without reference to clades previously introduced:

392-586: A common phenomenon in nature, particularly in plants where polyploidy allows for rapid speciation. Some cladist authors do not consider species to possess the property of "-phyly", which they assert applies only to groups of species. Gill A gill ( / ɡ ɪ l / ) is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide . The gills of some species, such as hermit crabs , have adapted to allow respiration on land provided they are kept moist. The microscopic structure of

441-451: A gill or gills. Many invertebrates, and even amphibians, use both the body surface and gills for gaseous exchange. Gills usually consist of thin filaments of tissue , lamellae (plates), branches, or slender, tufted processes that have a highly folded surface to increase surface area . The delicate nature of the gills is possible because the surrounding water provides support. The blood or other body fluid must be in intimate contact with

490-424: A gill presents a large surface area to the external environment. Branchia ( pl. : branchiae) is the zoologists' name for gills (from Ancient Greek βράγχια ). With the exception of some aquatic insects , the filaments and lamellae (folds) contain blood or coelomic fluid , from which gases are exchanged through the thin walls. The blood carries oxygen to other parts of the body. Carbon dioxide passes from

539-407: A given species), and as many as six "kidneys" (actually nephridia ). The tip or point of their low shells points forward rather than towards the back. The shell ranges from 3 mm to 37 mm in diameter depending on species. Like in chitons , the head is poorly defined, and there are no eyes. The mouth is located within the animal's undeveloped head in front of its single large foot and contains

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588-418: A lot of', and φῦλον ( phûlon ) 'genus, species', and refers to the fact that a polyphyletic group includes organisms (e.g., genera, species) arising from multiple ancestral sources. Conversely, the term monophyly , or monophyletic , employs the ancient Greek adjective μόνος ( mónos ) 'alone, only, unique', and refers to the fact that a monophyletic group includes organisms consisting of all

637-421: A newly discovered grass in the monophyletic family Poaceae , the true grasses, immediately results in numerous predictions about its structure and its developmental and reproductive characteristics, that are synapomorphies of this family. In contrast, Linnaeus' assignment of plants with two stamens to the polyphyletic class Diandria, while practical for identification, turns out to be useless for prediction, since

686-508: A secondary loss of shells caused a monoplacophoran body form to re-appear secondarily, which is plausible: At the very least, modern monoplacophorans are not closely related to vent-dwelling representatives from the Silurian. Cambrian monoplacophoran Knightoconus antarcticus is hypothesised to be an ancestor to the cephalopods . Living families: Extinct families: Many Cambrian-Devonian species have been described as "monoplacophorans", but

735-480: A separate tube which has no respiratory tissue (the pharyngocutaneous duct) develops beneath the pharynx proper, expelling ingested debris by closing a valve at its anterior end. Lungfish larvae also have external gills , as does the primitive ray-finned fish Polypterus , though the latter has a structure different from amphibians. Tadpoles of amphibians have from three to five gill slits that do not contain actual gills. Usually no spiracle or true operculum

784-421: A small mass of cells without any remaining gill-like structure. Marine teleosts also use their gills to excrete osmolytes (e.g. Na⁺, Cl ). The gills' large surface area tends to create a problem for fish that seek to regulate the osmolarity of their internal fluids. Seawater contains more osmolytes than the fish's internal fluids, so marine fishes naturally lose water through their gills via osmosis. To regain

833-451: A thin film of atmospheric oxygen in an area with small openings called spiracles that connect to the tracheal system. The plastron typically consists of dense patches of hydrophobic setae on the body, which prevent water entry into the spiracles, but may also involve scales or microscopic ridges projecting from the cuticle. The physical properties of the interface between the trapped air film and surrounding water allow gas exchange through

882-407: Is 210 cm /L. Water is 777 times more dense than air and is 100 times more viscous. Oxygen has a diffusion rate in air 10,000 times greater than in water. The use of sac-like lungs to remove oxygen from water would not be efficient enough to sustain life. Rather than using lungs, "[g]aseous exchange takes place across the surface of highly vascularised gills over which a one-way current of water

931-432: Is kept flowing by a specialised pumping mechanism. The density of the water prevents the gills from collapsing and lying on top of each other, which is what happens when a fish is taken out of water." Usually water is moved across the gills in one direction by the current, by the motion of the animal through the water, by the beating of cilia or other appendages, or by means of a pumping mechanism. In fish and some molluscs,

980-627: Is present, though many species have operculum-like structures. Instead of internal gills, they develop three feathery external gills that grow from the outer surface of the gill arches. Sometimes, adults retain these, but they usually disappear at metamorphosis . Examples of salamanders that retain their external gills upon reaching adulthood are the olm and the mudpuppy . Still, some extinct tetrapod groups did retain true gills. A study on Archegosaurus demonstrates that it had internal gills like true fish. Crustaceans , molluscs , and some aquatic insects have tufted gills or plate-like structures on

1029-405: Is renewed through the gills. In the larval dragonfly , the wall of the caudal end of the alimentary tract ( rectum ) is richly supplied with tracheae as a rectal gill, and water pumped into and out of the rectum provides oxygen to the closed tracheae. A plastron is a type of structural adaptation occurring among some aquatic arthropods (primarily insects), a form of inorganic gill which holds

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1078-412: The chitons , such as having segmented anatomy (organs arranged in series). There are eight pairs of dorso-ventral muscles (shell muscles). The nervous system is relatively simple, with no true ganglion present. The repeated organs include from three to six pairs of "gills" (actually ctenidia ) located in a curved line along each side of the foot (though the number is not always considered definitive of

1127-481: The chitons . The two classes in this new clade, with the proposed name Serialia, all show a variable number of serially repeated gills and eight sets of dorsoventral pedal retractor muscles. This study contradicts the fossil evidence, which suggests that the Monoplacophora are the sister group to the remainder of the conchiferans , and that the cephalopods ( squids , octopuses , and relatives) arose from within

1176-472: The spiracle , lies in the back of the first gill slit . This bears a small pseudobranch that resembles a gill in structure, but only receives blood already oxygenated by the true gills. The spiracle is thought to be homologous to the ear opening in higher vertebrates . Most sharks rely on ram ventilation, forcing water into the mouth and over the gills by rapidly swimming forward. In slow-moving or bottom-dwelling species, especially among skates and rays,

1225-579: The Elder held that fish respired by their gills, but observed that Aristotle was of another opinion. The word branchia comes from the Greek βράγχια , "gills", plural of βράγχιον (in singular, meaning a fin ). Many microscopic aquatic animals, and some larger but inactive ones, can absorb sufficient oxygen through the entire surface of their bodies, and so can respire adequately without gills. However, more complex or more active aquatic organisms usually require

1274-535: The ancestors of birds; "warm-blooded animals" is therefore a polyphyletic grouping. Other examples of polyphyletic groups are algae , C4 photosynthetic plants , and edentates . Many taxonomists aim to avoid homoplasies in grouping taxa together, with a goal to identify and eliminate groups that are found to be polyphyletic. This is often the stimulus for major revisions of the classification schemes. Researchers concerned more with ecology than with systematics may take polyphyletic groups as legitimate subject matter;

1323-560: The blood through the thin gill tissue into the water. Gills or gill-like organs, located in different parts of the body, are found in various groups of aquatic animals, including mollusks , crustaceans , insects, fish, and amphibians . Semiterrestrial marine animals such as crabs and mudskippers have gill chambers in which they store water, enabling them to use the dissolved oxygen when they are on land. Galen observed that fish had multitudes of openings ( foramina ), big enough to admit gases, but too fine to give passage to water. Pliny

1372-559: The definition to cap-shaped forms, excluding spiral and other shapes of shell. The inclusion of the gastropod-like Bellerophontoidea within the group is also contentious. One attempt to resolve this confusion was to separate out the predominantly coiled helcionelloids from the traditional, cap-like tergomyans , this latter group containing extant Tryblidiids. Taxonomy of Monoplacophora per Bouchet, et al. (2017): Class Monoplacophora Monoplacophorans are univalved (though not gastropodal), limpet -shaped, and are untorted . They have

1421-402: The descendants of a unique common ancestor. By comparison, the term paraphyly , or paraphyletic , uses the ancient Greek preposition παρά ( pará ) 'beside, near', and refers to the situation in which one or several monophyletic subgroups are left apart from all other descendants of a unique common ancestor. In many schools of taxonomy , the recognition of polyphyletic groups in

1470-416: The efficiency of the gills is greatly enhanced by a countercurrent exchange mechanism in which the water passes over the gills in the opposite direction to the flow of blood through them. This mechanism is very efficient and as much as 90% of the dissolved oxygen in the water may be recovered. The gills of vertebrates typically develop in the walls of the pharynx , along a series of gill slits opening to

1519-419: The exterior. Most species employ a countercurrent exchange system to enhance the diffusion of substances in and out of the gill, with blood and water flowing in opposite directions to each other. The gills are composed of comb-like filaments, the gill lamellae , which help increase their surface area for oxygen exchange. When a fish breathes, it draws in a mouthful of water at regular intervals. Then it draws

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1568-504: The fish behind the head. Originally there were many slits, but during evolution, the number reduced, and modern fish mostly have five pairs, and never more than eight. Sharks and rays typically have five pairs of gill slits that open directly to the outside of the body, though some more primitive sharks have six pairs with the Broadnose sevengill shark being the only cartilaginous fish exceeding this number. Adjacent slits are separated by

1617-411: The gills are contained in spherical pouches, with a circular opening to the outside. Like the gill slits of higher fish, each pouch contains two gills. In some cases, the openings may be fused together, effectively forming an operculum. Lampreys have seven pairs of pouches, while hagfishes may have six to fourteen, depending on the species. In the hagfish, the pouches connect with the pharynx internally and

1666-492: The insect can become oxygen-depleted if there is no water movement, so many such insects in still water actively direct a flow of water over their bodies. The inorganic gill mechanism allows aquatic arthropods with plastrons to remain constantly submerged. Examples include many beetles in the family Elmidae , aquatic weevils , and true bugs in the family Aphelocheiridae , as well as at least one species of ricinuleid arachnid and various mites. A somewhat similar mechanism

1715-501: The monoplacophoran lineage. However, some authors dispute this view and do not necessarily see modern Monoplacophora as related to their presumed fossil ancestors. The concept of Serialia is supported by other molecular studies. The fossil record does indicate that the ancestral mollusc was monoplacophoran-like and that the Polyplacophora arose from within the Monoplacophora – not the other way around. This could be reconciled if

1764-479: The mouth by the beating of cilia. Respiration in the echinoderms (such as starfish and sea urchins ) is carried out using a very primitive version of gills called papulae . These thin protuberances on the surface of the body contain diverticula of the water vascular system . The gills of aquatic insects are tracheal , but the air tubes are sealed, commonly connected to thin external plates or tufted structures that allow diffusion. The oxygen in these tubes

1813-517: The only fossil members of the crown group date to the Pleistocene. The Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005) also contains Paleozoic molluscs of uncertain systematic position . It is not known whether these were gastropods or monoplacophorans. Polyphyletic For example, the biological characteristic of warm-bloodedness evolved separately in the ancestors of mammals and

1862-400: The presence of exactly two stamens has developed convergently in many groups. Species have a special status in systematics as being an observable feature of nature itself and as the basic unit of classification. It is usually implicitly assumed that species are monophyletic (or at least paraphyletic ). However, hybrid speciation arguably leads to polyphyletic species. Hybrid species are

1911-413: The respiratory surface for ease of diffusion. A high surface area is crucial to the gas exchange of aquatic organisms, as water contains only a small fraction of the dissolved oxygen than air does, and it diffuses more slowly. A cubic meter of air contains about 275 grams of oxygen at STP . In fresh water , the dissolved oxygen content is approximately 8 cm /L compared to that of air which

1960-407: The shell of many monoplacophorans is limpet -like in shape, they are not gastropods , nor do they have any close relation to gastropods. Discussion about monoplacophorans is made difficult by the slippery definition of the taxon; some authors take it to refer to all non-gastropod molluscs with a single shell, or all single-shelled molluscs with serially repeated units; whereas other workers restrict

2009-415: The sides of its throat together, forcing the water through the gill openings, so it passes over the gills to the outside. Fish gill slits may be the evolutionary ancestors of the thymus glands , parathyroid glands , as well as many other structures derived from the embryonic branchial pouches . The gills of fish form a number of slits connecting the pharynx to the outside of the animal on either side of

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2058-448: The similarities in activity within the fungus group Alternaria , for example, can lead researchers to regard the group as a valid genus while acknowledging its polyphyly. In recent research, the concepts of monophyly, paraphyly, and polyphyly have been used in deducing key genes for barcoding of diverse groups of species. The term polyphyly , or polyphyletic , derives from the two Ancient Greek words πολύς ( polús ) 'many,

2107-417: The spiracle may be enlarged, and the fish breathes by sucking water through this opening, instead of through the mouth. Chimaeras differ from other cartilagenous fish, having lost both the spiracle and the fifth gill slit. The remaining slits are covered by an operculum , developed from the septum of the gill arch in front of the first gill. In bony fish , the gills lie in a branchial chamber covered by

2156-426: The spiracles, almost as if the insect were in atmospheric air. Carbon dioxide diffuses into the surrounding water due to its high solubility , while oxygen diffuses into the film as the concentration within the film has been reduced by respiration , and nitrogen also diffuses out as its tension has been increased. Oxygen diffuses into the air film at a higher rate than nitrogen diffuses out. However, water surrounding

2205-525: The surfaces of their bodies. Gills of various types and designs, simple or more elaborate, have evolved independently in the past, even among the same class of animals. The segments of polychaete worms bear parapodia many of which carry gills. Sponges lack specialised respiratory structures, and the whole of the animal acts as a gill as water is drawn through its spongy structure. Aquatic arthropods usually have gills which are in most cases modified appendages. In some crustaceans these are exposed directly to

2254-417: The third or fourth pair of kidneys. One genus, Micropilina , has apparently been recorded as brooding young in the distal oviduct and pallial groove, releasing the young when approximately 300 micrometers in diameter. In 2006 a molecular study on Laevipilina antarctica suggested that extant Monoplacophora and Polyplacophora form a well-supported clade with the researched Neopilina closest to

2303-404: The water from escaping. The gill arches of bony fish typically have no septum, so the gills alone project from the arch, supported by individual gill rays. Some species retain gill rakers. Though all but the most primitive bony fish lack spiracles, the pseudobranch associated with them often remains, being located at the base of the operculum. This is, however, often greatly reduced, consisting of

2352-555: The water, marine fishes drink large amounts of sea water while simultaneously expending energy to excrete salt through the Na /K -ATPase ionocytes (formerly known as mitochondrion-rich cells and chloride cells ). Conversely, fresh water contains less osmolytes than the fish's internal fluids. Therefore, freshwater fishes must utilize their gill ionocytes to attain ions from their environment to maintain optimal blood osmolarity. Lampreys and hagfish do not have gill slits as such. Instead,

2401-421: The water, while in others, they are protected inside a gill chamber . Horseshoe crabs have book gills which are external flaps, each with many thin leaf-like membranes. Many marine invertebrates such as bivalve molluscs are filter feeders . A current of water is maintained through the gills for gas exchange, and food particles are filtered out at the same time. These may be trapped in mucus and moved to

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