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105-430: The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms ) is a class containing about 162 species of unsegmented marine annelid worms . Sipuncula was once considered a phylum , but was demoted to a class of Annelida, based on recent molecular work. Sipunculans vary in size but most species are under 10 cm (4 in) in length. The body is divided into an unsegmented, bulbous trunk and

210-407: A hydrostatic skeleton to extend or contract the animal. When threatened, Sipunculid worms can contract their body into a shape resembling a peanut kernel —a practice that has given rise to the name "peanut worm". The introvert is pulled inside the trunk by two pairs of retractor muscles that extend as narrow ribbons from the trunk wall to attachment points in the introvert. It can be protruded from

315-423: A mutation–selection balance . It is predicted that a viral quasispecies at a low but evolutionarily neutral and highly connected (that is, flat) region in the fitness landscape will outcompete a quasispecies located at a higher but narrower fitness peak in which the surrounding mutants are unfit, "the quasispecies effect" or the "survival of the flattest". There is no suggestion that a viral quasispecies resembles

420-400: A "classical" method of determining species, such as with Linnaeus, early in evolutionary theory. However, different phenotypes are not necessarily different species (e.g. a four-winged Drosophila born to a two-winged mother is not a different species). Species named in this manner are called morphospecies . In the 1970s, Robert R. Sokal , Theodore J. Crovello and Peter Sneath proposed

525-424: A 'smallest clade' idea" (a phylogenetic species concept). Mishler and Wilkins and others concur with this approach, even though this would raise difficulties in biological nomenclature. Wilkins cited the ichthyologist Charles Tate Regan 's early 20th century remark that "a species is whatever a suitably qualified biologist chooses to call a species". Wilkins noted that the philosopher Philip Kitcher called this

630-428: A connected series of neighbouring populations, each of which can sexually interbreed with adjacent related populations, but for which there exist at least two "end" populations in the series, which are too distantly related to interbreed, though there is a potential gene flow between each "linked" population. Such non-breeding, though genetically connected, "end" populations may co-exist in the same region thus closing

735-831: A day. They are sensitive to low salinities, and thus not commonly found near estuaries. They can also be abundant in coralline rock , and in Hawaii, up to seven hundred individuals have been found per square metre in burrows in the rock. Both asexual and sexual reproduction can be found in sipunculans, although asexual reproduction is uncommon and has only been observed in Aspidosiphon elegans and Sipunculus robustus . These reproduce asexually through transverse fission , followed by regeneration of vital body components, with S. robustus also reproducing by budding . One species of sipunculan, Themiste lageniformis , has been recorded as reproducing parthenogenetically ; eggs produced in

840-432: A different species from its ancestors. Viruses have enormous populations, are doubtfully living since they consist of little more than a string of DNA or RNA in a protein coat, and mutate rapidly. All of these factors make conventional species concepts largely inapplicable. A viral quasispecies is a group of genotypes related by similar mutations, competing within a highly mutagenic environment, and hence governed by

945-750: A few genera. Archaeogolfingia and Cambrosipunculus appear in the Cambrian Chengjiang biota in China. These fossils appear to belong to the crown group , and demonstrate that sipunculans have changed little (morphologically) since the early Cambrian, about 520 million years ago. An unnamed sipunculid worm from the Cambrian period has been discovered in the Burgess Shale in Alberta, Canada , and Lecthaylus has been identified from

1050-527: A fluid-filled cavity. These structures are constructed of muscle and connective fibers, densely packed into a 3-D structure. In many cases, the muscular hydrostat can be manipulated in all three dimensions. This allows for more precise movement compared to a typical hydrostatic skeleton. While in typical hydrostatic skeletons, movement is generated by applying force to a fluid-filled cavity, muscular hydrostats generate movement by muscle contractions. When one muscle contracts and decreases in area, other muscles within

1155-418: A force when squeezing through rocks and create a “prying open” gesture. There is a lightweight, flexible component to them that allows this movement with very little muscle mass. These organisms are also able to heal faster than organisms that contain hard skeletons. Healing in these organisms varies from creature to creature. However, if the cavity needs to be refilled, the “fluid” can easily be refilled if it

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1260-508: A genetic boundary suitable for defining a species concept is present. DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to use. One of the barcodes is a region of mitochondrial DNA within the gene for cytochrome c oxidase . A database, Barcode of Life Data System , contains DNA barcode sequences from over 190,000 species. However, scientists such as Rob DeSalle have expressed concern that classical taxonomy and DNA barcoding, which they consider

1365-405: A hydroskeleton possesses the ability to affect shape and movement, and involves two mechanical units: the muscle layers and the body wall. The muscular layers are longitudinal and circular, and part of the fluid-filled coelom within. Contractions of the circular muscles lengthen the organism's body, while contractions of the longitudinal muscles shorten the organism's body. Fluid within the organism

1470-399: A large amount of access to the necessary elements for survival. Terrestrial organisms that have hydrostatic skeletons generally have a lack of strength because they are not in a fluid environment. If one were to expand its body too much, it would collapse under its own weight. Hydrostatic skeletons are very common in invertebrates. A common example is the earthworm . Also, hydrostatic nature

1575-465: A lineage should be divided into multiple chronospecies , or when populations have diverged to have enough distinct character states to be described as cladistic species. Species and higher taxa were seen from the time of Aristotle until the 18th century as categories that could be arranged in a hierarchy, the great chain of being . In the 19th century, biologists grasped that species could evolve given sufficient time. Charles Darwin 's 1859 book On

1680-492: A misnomer, need to be reconciled, as they delimit species differently. Genetic introgression mediated by endosymbionts and other vectors can further make barcodes ineffective in the identification of species. A phylogenetic or cladistic species is "the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states in comparable individuals (semaphoronts)". The empirical basis – observed character states – provides

1785-561: A narrower, anterior section, called the "introvert", which can be retracted into the trunk. The mouth is at the tip of the introvert and is surrounded in most groups by a ring of short tentacles. With no hard parts, the body is flexible and mobile. Although found in a range of habitats throughout the world's oceans, the majority of species live in shallow water habitats, burrowing under the surface of sandy and muddy substrates. Others live under stones, in rock crevices or in other concealed locations. Most sipunculans are deposit feeders, extending

1890-429: A narrower, anterior section, called the "introvert". Sipunculans have a body wall somewhat similar to that of most other annelids (though unsegmented) in that it consists of an epidermis without cilia overlain by a cuticle , an outer layer of circular and an inner layer of longitudinal musculature . The body wall surrounds the coelom (body cavity) that is filled with fluid on which the body wall musculature acts as

1995-449: A particular species, including which genus (and higher taxa) it is placed in, is a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be corroborated or refuted. Sometimes, especially in the past when communication was more difficult, taxonomists working in isolation have given two distinct names to individual organisms later identified as

2100-400: A perfect model of life, it is still a useful tool to scientists and conservationists for studying life on Earth, regardless of the theoretical difficulties. If species were fixed and clearly distinct from one another, there would be no problem, but evolutionary processes cause species to change. This obliges taxonomists to decide, for example, when enough change has occurred to declare that

2205-400: A short way of saying that something applies to many species within a genus, but not to all. If scientists mean that something applies to all species within a genus, they use the genus name without the specific name or epithet. The names of genera and species are usually printed in italics . However, abbreviations such as "sp." should not be italicised. When a species' identity is not clear,

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2310-404: A specialist may use "cf." before the epithet to indicate that confirmation is required. The abbreviations "nr." (near) or "aff." (affine) may be used when the identity is unclear but when the species appears to be similar to the species mentioned after. With the rise of online databases, codes have been devised to provide identifiers for species that are already defined, including: The naming of

2415-523: A species as groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups. It has been argued that this definition is a natural consequence of the effect of sexual reproduction on the dynamics of natural selection. Mayr's use of the adjective "potentially" has been a point of debate; some interpretations exclude unusual or artificial matings that occur only in captivity, or that involve animals capable of mating but that do not normally do so in

2520-400: A species as determined by a taxonomist. A typological species is a group of organisms in which individuals conform to certain fixed properties (a type), so that even pre-literate people often recognise the same taxon as do modern taxonomists. The clusters of variations or phenotypes within specimens (such as longer or shorter tails) would differentiate the species. This method was used as

2625-491: A species. All species definitions assume that an organism acquires its genes from one or two parents very like the "daughter" organism, but that is not what happens in HGT. There is strong evidence of HGT between very dissimilar groups of prokaryotes , and at least occasionally between dissimilar groups of eukaryotes , including some crustaceans and echinoderms . The evolutionary biologist James Mallet concludes that there

2730-685: A species. Generally the term includes the unknown element of a distinct act of creation. Many authors have argued that a simple textbook definition, following Mayr's concept, works well for most multi-celled organisms , but breaks down in several situations: Species identification is made difficult by discordance between molecular and morphological investigations; these can be categorised as two types: (i) one morphology, multiple lineages (e.g. morphological convergence , cryptic species ) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity , multiple life-cycle stages). In addition, horizontal gene transfer (HGT) makes it difficult to define

2835-528: A taxonomic decision at the discretion of cognizant specialists, is not governed by the Codes of Zoological or Botanical Nomenclature, in contrast to the PhyloCode , and contrary to what is done in several other fields, in which the definitions of technical terms, like geochronological units and geopolitical entities, are explicitly delimited. The nomenclatural codes that guide the naming of species, including

2940-517: A traditional biological species. The International Committee on Taxonomy of Viruses has since 1962 developed a universal taxonomic scheme for viruses; this has stabilised viral taxonomy. Most modern textbooks make use of Ernst Mayr 's 1942 definition, known as the Biological Species Concept as a basis for further discussion on the definition of species. It is also called a reproductive or isolation concept. This defines

3045-520: A variant of the Latin siphunculus ("little tube"), a diminutive of sipho from Greek σίφων ( síphōn , " tube , pipe "). The Swedish naturalist Carl Linnaeus first described the worm Sipunculus nudus in his Systema Naturae in 1767. In 1814, the French zoologist Constantine Samuel Rafinesque used the word "Sipuncula" to describe the family (now Sipunculidae ), and in time,

3150-447: A variation on the morphological species concept, a phenetic species, defined as a set of organisms with a similar phenotype to each other, but a different phenotype from other sets of organisms. It differs from the morphological species concept in including a numerical measure of distance or similarity to cluster entities based on multivariate comparisons of a reasonably large number of phenotypic traits. A mate-recognition species

3255-515: A variety of reasons. Viruses are a special case, driven by a balance of mutation and selection , and can be treated as quasispecies . Biologists and taxonomists have made many attempts to define species, beginning from morphology and moving towards genetics . Early taxonomists such as Linnaeus had no option but to describe what they saw: this was later formalised as the typological or morphological species concept. Ernst Mayr emphasised reproductive isolation, but this, like other species concepts,

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3360-438: Is "an entity composed of organisms which maintains its identity from other such entities through time and over space, and which has its own independent evolutionary fate and historical tendencies". This differs from the biological species concept in embodying persistence over time. Wiley and Mayden stated that they see the evolutionary species concept as "identical" to Willi Hennig 's species-as-lineages concept, and asserted that

3465-500: Is a delicacy in southeast China , originally from Anhai , near Quanzhou . A sipunculid worm dish is also considered a delicacy in the islands of the Visayas region, Philippines . The muscle is first prepared by soaking it in spiced vinegar and then served with other ingredients as a dish similar to ceviche . It is a basic food for local fisherman and is sometimes seen in city restaurants as an appetizer. This style of food preparation

3570-400: Is a group of sexually reproducing organisms that recognise one another as potential mates. Expanding on this to allow for post-mating isolation, a cohesion species is the most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic cohesion mechanisms; no matter whether populations can hybridise successfully, they are still distinct cohesion species if

3675-452: Is a population of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring , typically by sexual reproduction . It is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity . Other ways of defining species include their karyotype , DNA sequence, morphology , behaviour, or ecological niche . In addition, paleontologists use

3780-458: Is a set of organisms adapted to a particular set of resources, called a niche, in the environment. According to this concept, populations form the discrete phenetic clusters that we recognise as species because the ecological and evolutionary processes controlling how resources are divided up tend to produce those clusters. A genetic species as defined by Robert Baker and Robert Bradley is a set of genetically isolated interbreeding populations. This

3885-414: Is called speciation . Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book The Origin of Species . Speciation depends on a measure of reproductive isolation , a reduced gene flow. This occurs most easily in allopatric speciation, where populations are separated geographically and can diverge gradually as mutations accumulate. Reproductive isolation

3990-412: Is called the specific name or the specific epithet (in botanical nomenclature , also sometimes in zoological nomenclature ). For example, Boa constrictor is one of the species of the genus Boa , with constrictor being the species' epithet. While the definitions given above may seem adequate at first glance, when looked at more closely they represent problematic species concepts. For example,

4095-402: Is common in marine life such as jellyfish and sea anemones . Earthworms have rings of muscles that are filled with fluid, making their entire body hydrostatic. A sea anemone has a hydrostatic head, with arms radiating out around the mouth. This structure is helpful in feeding and locomotion. An example of a simple Deuterostome containing a hydrostatic skeleton would be Enteropneusta , with

4200-403: Is described formally, in a publication that assigns it a unique scientific name. The description typically provides means for identifying the new species, which may not be based solely on morphology (see cryptic species ), differentiating it from other previously described and related or confusable species and provides a validly published name (in botany) or an available name (in zoology) when

4305-430: Is evenly concentrated so the forces of the muscle are spread throughout the whole organism and shape changes can persist. These structural factors also persist in a hydrostatic organ. A non-helical hydrostatic skeleton structure is the functional basis of the mammalian penis , which fills the corpus cavernosa with blood to maintain physical rigidity during coitus . Helically reinforced hydrostatic skeleton structure

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4410-671: Is further weakened by the existence of microspecies , groups of organisms, including many plants, with very little genetic variability, usually forming species aggregates . For example, the dandelion Taraxacum officinale and the blackberry Rubus fruticosus are aggregates with many microspecies—perhaps 400 in the case of the blackberry and over 200 in the dandelion, complicated by hybridisation , apomixis and polyploidy , making gene flow between populations difficult to determine, and their taxonomy debatable. Species complexes occur in insects such as Heliconius butterflies, vertebrates such as Hypsiboas treefrogs, and fungi such as

4515-726: Is hard or even impossible to test. Later biologists have tried to refine Mayr's definition with the recognition and cohesion concepts, among others. Many of the concepts are quite similar or overlap, so they are not easy to count: the biologist R. L. Mayden recorded about 24 concepts, and the philosopher of science John Wilkins counted 26. Wilkins further grouped the species concepts into seven basic kinds of concepts: (1) agamospecies for asexual organisms (2) biospecies for reproductively isolated sexual organisms (3) ecospecies based on ecological niches (4) evolutionary species based on lineage (5) genetic species based on gene pool (6) morphospecies based on form or phenotype and (7) taxonomic species,

4620-570: Is locally called kilawin or kinilaw , and is also used for fish, conch and vegetables. The worms, especially in dried form, are considered a delicacy in Vietnam as well, where they are caught on the coasts of Minh Chao island, in Van Don District . The relatively high market price of the worms have made them a significant source of income for the local population of fishermen families. Species A species ( pl. : species)

4725-403: Is no easy way to tell whether related geographic or temporal forms belong to the same or different species. Species gaps can be verified only locally and at a point of time. One is forced to admit that Darwin's insight is correct: any local reality or integrity of species is greatly reduced over large geographic ranges and time periods. The botanist Brent Mishler argued that the species concept

4830-478: Is not valid, notably because gene flux decreases gradually rather than in discrete steps, which hampers objective delimitation of species. Indeed, complex and unstable patterns of gene flux have been observed in cichlid teleosts of the East African Great Lakes . Wilkins argued that "if we were being true to evolution and the consequent phylogenetic approach to taxa, we should replace it with

4935-586: Is similar to Mayr's Biological Species Concept, but stresses genetic rather than reproductive isolation. In the 21st century, a genetic species could be established by comparing DNA sequences. Earlier, other methods were available, such as comparing karyotypes (sets of chromosomes ) and allozymes ( enzyme variants). An evolutionarily significant unit (ESU) or "wildlife species" is a population of organisms considered distinct for purposes of conservation. In palaeontology , with only comparative anatomy (morphology) and histology from fossils as evidence,

5040-408: Is sometimes an important source of genetic variation. Viruses can transfer genes between species. Bacteria can exchange plasmids with bacteria of other species, including some apparently distantly related ones in different phylogenetic domains , making analysis of their relationships difficult, and weakening the concept of a bacterial species. Hydrostatic skeleton As a skeletal structure,

5145-548: Is swallowed but larger particles are discarded. Species dwelling in crevices are able to withdraw their introverts, blocking the crevice entrance with their thickened trunks and presumably ingesting any food they have snared at the same time. One species, Thysanocardia procera is thought to be carnivorous , gaining entrance in some way to the interior of the sea mouse Aphrodita aculeata and sucking out its liquefied contents. Because of their soft-bodied structure, fossils of sipunculans are extremely rare, and are only known from

5250-594: Is threatened by hybridisation, but this can be selected against once a pair of populations have incompatible alleles of the same gene, as described in the Bateson–Dobzhansky–Muller model . A different mechanism, phyletic speciation, involves one lineage gradually changing over time into a new and distinct form (a chronospecies ), without increasing the number of resultant species. Horizontal gene transfer between organisms of different species, either through hybridisation , antigenic shift , or reassortment ,

5355-484: Is triggered by the presence of adults. Most sipunculans are deposit feeders employing a number of different methods to obtain their foods. Those living in burrows extend their tentacles over the surface of the sediment. Food particles get trapped in mucous secretions and the beating of cilia transport the particles to the mouth. Among those that burrow through the sand, the tentacles are replaced by fluted folds which scoop up sediment and food particles. Most of this material

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5460-498: Is typical for flexible structures as in soft-bodied animals. Hydrostatic skeletons are typically arranged in a cylinder. Hydrostatic skeletons can be controlled by several different muscle types. Length can be adjusted by longitudinal muscle fibers parallel to the longitudinal axis. The muscle fibers may be found in continuous sheets or isolated bundles, and the diameter can be manipulated by three different muscle types: circular, radial, and transverse. Circular musculature wraps around

5565-471: Is water or blood. If the fluid is some other type of liquid, it can take longer, but it is still faster than healing a bone. The common earthworm is also able to regrow damaged parts of its body. These organisms have some relatively simple pathways for circulation and respiration. Also, these organisms have a cushion to allow protection for internal organs from shock. However, it does not protect internal organs from external damage very effectively. Because

5670-690: The Granton Shrimp Bed , near Edinburgh, Scotland, dating to the Silurian period. Trace fossils of burrows that may have been formed by sipunculans have been found from the Paleozoic . Some scientists once hypothesized a close relationship between sipunculans and the extinct hyoliths , operculate shells from the Palaeozoic with which they share a helical gut; but this hypothesis has since been discounted. Sipunculid worm jelly ( 土笋凍 )

5775-530: The ICZN for animals and the ICN for plants, do not make rules for defining the boundaries of the species. Research can change the boundaries, also known as circumscription, based on new evidence. Species may then need to be distinguished by the boundary definitions used, and in such cases the names may be qualified with sensu stricto ("in the narrow sense") to denote usage in the exact meaning given by an author such as

5880-415: The coelom , which contains the respiratory pigment haemerythrin , and the separate tentacular system, the two being separated by an elaborate septum . The coelomic fluid contains five types of coelomic cells: haemocytes , granulocytes , large multinuclear cells, ciliated urn-shaped cells and immature cells. The ciliated urn cells may also be attached to the peritoneum and assist in waste filtering from

5985-399: The fly agaric . Natural hybridisation presents a challenge to the concept of a reproductively isolated species, as fertile hybrids permit gene flow between two populations. For example, the carrion crow Corvus corone and the hooded crow Corvus cornix appear and are classified as separate species, yet they can hybridise where their geographical ranges overlap. A ring species is

6090-507: The jaguar ( Panthera onca ) of Latin America or the leopard ( Panthera pardus ) of Africa and Asia. In contrast, the scientific names of species are chosen to be unique and universal (except for some inter-code homonyms ); they are in two parts used together : the genus as in Puma , and the specific epithet as in concolor . A species is given a taxonomic name when a type specimen

6195-406: The "cynical species concept", and arguing that far from being cynical, it usefully leads to an empirical taxonomy for any given group, based on taxonomists' experience. Other biologists have gone further and argued that we should abandon species entirely, and refer to the "Least Inclusive Taxonomic Units" (LITUs), a view that would be coherent with current evolutionary theory. The species concept

6300-495: The Annelida (including echiurans and pogonophorans ). It has also been shown that a rudimentary neural segmentation similar to that of annelids occurs in the early larval stage, even if these traits are absent in the adults. Sipunculans are worms ranging from 2 to 720 mm (0.1 to 28.3 in) in length, with most species being under 10 cm (4 in). The sipunculan body is divided into an unsegmented, bulbous trunk and

6405-626: The Origin of Species explained how species could arise by natural selection . That understanding was greatly extended in the 20th century through genetics and population ecology . Genetic variability arises from mutations and recombination , while organisms themselves are mobile, leading to geographical isolation and genetic drift with varying selection pressures . Genes can sometimes be exchanged between species by horizontal gene transfer ; new species can arise rapidly through hybridisation and polyploidy ; and species may become extinct for

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6510-405: The abbreviation "sp." in the singular or "spp." (standing for species pluralis , Latin for "multiple species") in the plural in place of the specific name or epithet (e.g. Canis sp.). This commonly occurs when authors are confident that some individuals belong to a particular genus but are not sure to which exact species they belong, as is common in paleontology . Authors may also use "spp." as

6615-536: The absence of sperm developed through the normal stages. Most sipunculan species are dioecious . Their gametes are produced in the coelomic lining, where they are released into the coelom to mature. These gametes are then picked up by the metanephridia system and released into the aquatic environment, where fertilisation takes place. In at least one species, Themiste pyroides , swarming behaviour occurs with adults creating compact masses among rocks immediately before spawning. Although some species hatch directly into

6720-408: The adult form, many have a trochophore larva, which metamorphoses into the adult after anything from a day to a month, depending on species. In a few species, the trochophore does not develop directly into the adult, but into an intermediate pelagosphaera stage, that possesses a greatly enlarged metatroch (ciliated band). Metamorphosis occurs only in the presence of suitable habitat conditions, and

6825-570: The amount of hybridisation is insufficient to completely mix their respective gene pools . A further development of the recognition concept is provided by the biosemiotic concept of species. In microbiology , genes can move freely even between distantly related bacteria, possibly extending to the whole bacterial domain. As a rule of thumb, microbiologists have assumed that members of Bacteria or Archaea with 16S ribosomal RNA gene sequences more similar than 97% to each other need to be checked by DNA–DNA hybridisation to decide if they belong to

6930-474: The anal shield is near the anteriorly-located anus on the trunk, just below the introvert of the animal, while the caudal shield is at the posterior of the body. In Aspidosiphon and Lithacrosiphon the anal shield is restricted to the dorsal side, causing the introvert to emerge at an angle, whereas it surrounds the anterior trunk in Cloeosiphon with the introvert emerging from its center. In Aspidosiphon

7035-433: The anterior third of the trunk. Digestion is extra-cellular , taking place in the lumen of the intestine. A rectal caecum, present in most species, is a blind ending sac at the transition between intestine and rectum with unknown function. The anus is often not visible when the introvert is retracted into the trunk. Sipunculans do not have a vascular blood system. Fluid transport and gas exchange are instead accomplished by

7140-462: The basis of developmental and larval characters. These phyla have been included in a larger group, the Lophotrochozoa , that also includes the annelids , the ribbon worms and several other phyla. Phylogenetic analyses based on 79 ribosomal proteins indicated a position of Sipuncula within Annelida. Subsequent analysis of the mitochondrion 's DNA has confirmed their close relationship to

7245-474: The biological species concept, "the several versions" of the phylogenetic species concept, and the idea that species are of the same kind as higher taxa are not suitable for biodiversity studies (with the intention of estimating the number of species accurately). They further suggested that the concept works for both asexual and sexually-reproducing species. A version of the concept is Kevin de Queiroz 's "General Lineage Concept of Species". An ecological species

7350-505: The biological species concept, a cladistic species does not rely on reproductive isolation – its criteria are independent of processes that are integral in other concepts. Therefore, it applies to asexual lineages. However, it does not always provide clear cut and intuitively satisfying boundaries between taxa, and may require multiple sources of evidence, such as more than one polymorphic locus, to give plausible results. An evolutionary species, suggested by George Gaylord Simpson in 1951,

7455-551: The body wall, especially at the forward end of the introvert which is used for exploring the surrounding environment. All sipunculid worms are marine and benthic ; they are found throughout the world's oceans including polar waters, equatorial waters and the abyssal zone , but the majority of species occur in shallow water, where they are relatively common. They inhabit a range of habitats including burrowing in sand, mud, clay and gravel, hiding under stones, in rock crevices, in hollow coral heads, in wood, in empty seashells and inside

7560-416: The bones of dead whales . Some hide in kelp holdfasts , under tangles of eelgrass , inside sponges and in the empty tubes of other organisms, and some live among fouling organisms on man-made structures. Some bore into solid rocks to make a shelter for themselves. They are common below the surface of the sediment on tidal flats . These worms may stay submerged in the sea bed for between 10 and 18 hours

7665-412: The boring process. The digestive tract of sipunculans starts with the esophagus, located between the introvert retractor muscles. In the trunk the intestine runs posteriorly, forms a loop and turns anteriorly again. The downward and upward sections of the gut are coiled around each other, forming a double helix . At the termination of the gut coil, the rectum emerges and ends in the anus , located in

7770-428: The boundaries between closely related species become unclear with hybridisation , in a species complex of hundreds of similar microspecies , and in a ring species . Also, among organisms that reproduce only asexually , the concept of a reproductive species breaks down, and each clone is potentially a microspecies. Although none of these are entirely satisfactory definitions, and while the concept of species may not be

7875-407: The circumference of the cylinder, radial musculature extends from the center of the cylinder towards the surface, and transverse musculature arrange in parallel and perpendicular sheets crossing the diameter of the cylinder. Within the cylinder lies fluid, most often water. The fluid is resistant to changes in volume. Contraction of circular, radial or transverse muscles increases the pressure within

7980-575: The coelomic fluid. Nitrogenous waste is excreted through a pair of metanephridia opening close to the anus, except in Phascolion and Onchnesoma , which have only a single nephridium. A ciliated funnel, or nephrostome, opens into the coelomic cavity at the anterior end, close to the nephridiopore. The metanephridia have an osmoregulation function but it is unclear whether the mechanism is via filtration or secretion. They also serve as gamete storage and maintenance organs. The tentacular coelom connects

8085-455: The common name of acorn worm. This organism is classified as a Hemichordate , and they are marine worms that use their hydrostatic skeleton to tunnel and anchor themselves into the ground. This can be used for locomotion, but also can aid in the defense of the organism against outside forces as the worm can try to "hide" itself within the ocean floor. The mammalian penis is a hydrostatic organ. The hydrostatic fluid, in this case blood, fills

8190-433: The concept of a chronospecies can be applied. During anagenesis (evolution, not necessarily involving branching), some palaeontologists seek to identify a sequence of species, each one derived from the phyletically extinct one before through continuous, slow and more or less uniform change. In such a time sequence, some palaeontologists assess how much change is required for a morphologically distinct form to be considered

8295-416: The concept of the chronospecies since fossil reproduction cannot be examined. The most recent rigorous estimate for the total number of species of eukaryotes is between 8 and 8.7 million. About 14% of these had been described by 2011. All species (except viruses ) are given a two-part name , called a "binomial". The first part of a binomial is the genus to which the species belongs. The second part

8400-433: The cylinder, and results in an increase in length. Contraction of longitudinal muscles can shorten the cylinder. Change in shape is limited by connective tissue fibers. Connective fibers, often collagenous, are arranged in a helical shape within the wall of the hydrostatic skeleton. The helical shape formed by these fibers allows for elongation and shortening of the skeleton, while still remaining rigid to prevent torsion. As

8505-435: The evidence to support hypotheses about evolutionarily divergent lineages that have maintained their hereditary integrity through time and space. Molecular markers may be used to determine diagnostic genetic differences in the nuclear or mitochondrial DNA of various species. For example, in a study done on fungi , studying the nucleotide characters using cladistic species produced the most accurate results in recognising

8610-402: The hydrostatic skeletons have limited ability for attachment of limbs, the organisms are relatively simple and do not have many abilities to grab or latch onto things. Organisms with complete hydrostatic skeletons need to be in an environment that allows them to re-fill themselves with their fluid that is necessary for survival. This is why hydrostatic skeletons are common in marine life. They have

8715-420: The introvert to gather food particles and draw them into the mouth, and retracting the introvert when feeding conditions are unsuitable or danger threatens. With a few exceptions, reproduction is sexual and involves a planktonic larval stage. Sipunculid worms are used as food in some countries in south-east Asia. Sipuncula is a feminine variant of the now-obsolete genus name Sipunculus , itself

8820-402: The longitudinal axis. These fibers remain folded when the penis is flaccid, but unfold as the penis fills with blood during an erection, which allows the penis to resist bending. The penises of turtles are structured similarly, although they evolved separately. Other vertebrates sometimes utilize a modified hydrostatic skeleton called a muscular hydrostat . Muscular hydrostats do not contain

8925-431: The members of this group being specialized filter feeders , unlike the other groups of sipunculans which are deposit feeders . The tentacles are hollow and are extended via hydrostatic pressure in a similar manner as the introvert, but have a different mechanism from that of the rest of the introvert, being connected, via a system of ducts, to one or two contractile sacs next to the oesophagus . Hooks are often present near

9030-539: The mouth on the introvert. These are proteinaceous, non- chitinous specializations of the epidermis , either arranged in rings or scattered. They may be involved in scraping algae off rock, or alternatively provide anchorage. Three genera ( Aspidosiphon , Lithacrosiphon and Cloeosiphon ) in the Aspidosiphonidae family possess epidermal structures, known as anal and caudal shields. These are patches of thickened, hard plates, and are used for boring into rock;

9135-492: The muscles of the body wall. In some species, there are simple light-sensitive ocelli associated with the brain. Two organs, likely functioning as a unit for chemoreception are located near its anterior margin; the non-ciliated cerebral organ, which possesses bipolar sensory cells , and the nuchal organ, located posterior to the brain. Similar light-sensing tubes have been reported in the fauveliopsid annelids. In addition, all sipunculans have numerous sensory nerve endings on

9240-483: The numerous fungi species of all the concepts studied. Versions of the phylogenetic species concept that emphasise monophyly or diagnosability may lead to splitting of existing species, for example in Bovidae , by recognising old subspecies as species, despite the fact that there are no reproductive barriers, and populations may intergrade morphologically. Others have called this approach taxonomic inflation , diluting

9345-593: The paper is accepted for publication. The type material is usually held in a permanent repository, often the research collection of a major museum or university, that allows independent verification and the means to compare specimens. Describers of new species are asked to choose names that, in the words of the International Code of Zoological Nomenclature , are "appropriate, compact, euphonious, memorable, and do not cause offence". Books and articles sometimes intentionally do not identify species fully, using

9450-426: The penis during an erection . Unlike the hydrostatic skeletons of many invertebrates, which use the bending of the animal for locomotion, the penis must resist bending and shape changes during sexual intercourse . Instead of connective fibers arranged in a helical shape, the penis contains a layer called the corpus cavernosum . The corpus cavernosum contains connective fibers arranged both parallel and perpendicular to

9555-674: The person who named the species, while the antonym sensu lato ("in the broad sense") denotes a wider usage, for instance including other subspecies. Other abbreviations such as "auct." ("author"), and qualifiers such as "non" ("not") may be used to further clarify the sense in which the specified authors delineated or described the species. Species are subject to change, whether by evolving into new species, exchanging genes with other species, merging with other species or by becoming extinct. The evolutionary process by which biological populations of sexually-reproducing organisms evolve to become distinct or reproductively isolated as species

9660-487: The result of misclassification leading to questions on whether there really are any ring species. The commonly used names for kinds of organisms are often ambiguous: "cat" could mean the domestic cat, Felis catus , or the cat family, Felidae . Another problem with common names is that they often vary from place to place, so that puma, cougar, catamount, panther, painter and mountain lion all mean Puma concolor in various parts of America, while "panther" may also mean

9765-586: The ring. Ring species thus present a difficulty for any species concept that relies on reproductive isolation. However, ring species are at best rare. Proposed examples include the herring gull – lesser black-backed gull complex around the North pole, the Ensatina eschscholtzii group of 19 populations of salamanders in America, and the greenish warbler in Asia, but many so-called ring species have turned out to be

9870-508: The same species. This concept was narrowed in 2006 to a similarity of 98.7%. The average nucleotide identity (ANI) method quantifies genetic distance between entire genomes , using regions of about 10,000 base pairs . With enough data from genomes of one genus, algorithms can be used to categorize species, as for Pseudomonas avellanae in 2013, and for all sequenced bacteria and archaea since 2020. Observed ANI values among sequences appear to have an "ANI gap" at 85–95%, suggesting that

9975-529: The same species. When two species names are discovered to apply to the same species, the older species name is given priority and usually retained, and the newer name considered as a junior synonym, a process called synonymy . Dividing a taxon into multiple, often new, taxa is called splitting . Taxonomists are often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to recognising differences or commonalities between organisms. The circumscription of taxa, considered

10080-454: The shape of the cylinder changes, the pitch of the helix will change. The angle relative to the long axis will decrease during elongation and increase during shortening. Organisms containing a hydrostatic skeleton have advantages and disadvantages. Their fluid shape allows them to move around easily while swimming and burrowing. They can fit through oddly shaped passages and hide themselves more effectively from predators. They are able to create

10185-537: The shield is a hardened, horny structure; in Lithacrosiphon it is a calcareous cone; in Cloeosiphon it is composed of separate plates. When the introvert is retracted in these animals, the anal shield blocks the entrance to its burrow. At the posterior end of the trunk, a hardened caudal shield is sometimes present in Aspidosiphon ; this may help with anchoring the animal in its burrow or may be used in

10290-440: The skin is thin and respiration is mainly through the cuticle of the trunk, where oxygen uptake is assisted by the presence of dermal coelomic canals just beneath the epidermis . The nervous system consists of dorsal cerebral ganglion, brain above the oesophagus and a nerve ring around the oesophagus, which links the brain with the single ventral nerve cord that runs the length of the body. Lateral nerves lead off this to innervate

10395-506: The species concept and making taxonomy unstable. Yet others defend this approach, considering "taxonomic inflation" pejorative and labelling the opposing view as "taxonomic conservatism"; claiming it is politically expedient to split species and recognise smaller populations at the species level, because this means they can more easily be included as endangered in the IUCN red list and can attract conservation legislation and funding. Unlike

10500-460: The structure must expand in response. Helical muscles may be present, which can create torsion, an ability that is restricted in hydrostatic skeletons. Muscular hydrostats are found in mammalian, reptilian, and amphibian tongues. Mammalian tongues have the structure of a central core of muscle fibers surrounded by bundles of longitudinal muscles and alternating parallel sheets of transverse muscle fibers. Elephant trunks and tapir proboscises also utilize

10605-439: The subclass Phascolosomatidea , the tentacles are arranged in an arc above the mouth, surrounding the nuchal organ , also located at the tip of the introvert. The tentacles each have a deep groove along which food is moved to the mouth by cilia. They are used to gather organic detritus from the water or substrate, and probably also function as gills . In the family Themistidae the tentacles form an elaborate crown-like structure,

10710-435: The tentacles at the tip of the introvert to a ring canal at their base, from which a contractile vessel runs along beside the esophagus and ends blindly posteriorly. Some evidence points towards the involvement of these structures in ultrafiltration. In crevice-dwelling sipunculans, respiration is mainly through the tentacular system, with oxygen diffusing into the trunk coelom from the tentacular coelom. However, in other species

10815-475: The term came to be used for the whole class. This is a relatively understudied group, and it is estimated there may be around 162 species worldwide. The phylogenetic placement of this group in the past has proved troublesome. Originally classified as annelids , despite the complete lack of segmentation, bristles and other annelid characters, the phylum Sipuncula was later allied with the Mollusca , mostly on

10920-433: The trunk by contracting the muscles of the trunk wall, thus forcing the fluid in the body cavity forwards. The introvert can vary in size from half the length of the trunk to several times its length, but whatever their comparative sizes, it is fully retractable. The mouth is located at the anterior end of the animal; in the subclass Sipunculidea , the mouth is surrounded by a mass of 18 to 24 ciliated tentacles , while in

11025-540: The wild. It is difficult to define a species in a way that applies to all organisms. The debate about species concepts is called the species problem. The problem was recognised even in 1859, when Darwin wrote in On the Origin of Species : I was much struck how entirely vague and arbitrary is the distinction between species and varieties. He went on to write: No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of

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