The Hilsenhoff Biotic Index (HBI) is a quantitative method of evaluating the abundance of arthropod fauna in stream ecosystems as a measurement of estimating water quality based on the predetermined pollution tolerances of the observed taxa. This biotic index was created by William Hilsenhoff in 1977 to measure the effects of oxygen depletion in Wisconsin streams resulting from organic or nutrient pollution.
111-413: The collection sample should contain 100+ arthropods. A tolerance value of 0 to 10 is assigned to each arthropod species (or genera ) based on its known prevalence in stream habitats with varying states of detritus contamination. A highly tolerant species would receive a value of 10, while a species collected only in unaltered streams with high water quality would receive a value of 0. The sum products of
222-475: A synthetic population . In horticulture , the term stable hybrid is used to describe an annual plant that, if grown and bred in a small monoculture free of external pollen (e.g., an air-filtered greenhouse) produces offspring that are "true to type" with respect to phenotype; i.e., a true-breeding organism. Hybridization can occur in the hybrid zones where the geographical ranges of species, subspecies, or distinct genetic lineages overlap. For example,
333-400: A = tolerance value of taxa; N = total number of specimens in the sample. Precautions should be taken to account for confounding variables , such as the effects of dominant species over-abundance, seasonal temperature stress, and water currents. Limiting the collection of individuals from each species to a maximum of 10 (10-Max BI) has been shown to minimize the effects of these phenomena on
444-460: A bactrian camel and a dromedary . There are many examples of felid hybrids , including the liger . The oldest-known animal hybrid bred by humans is the kunga equid hybrid produced as a draft animal and status symbol 4,500 years ago in Umm el-Marra , present-day Syria . The first known instance of hybrid speciation in marine mammals was discovered in 2014. The clymene dolphin ( Stenella clymene )
555-423: 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
666-400: 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
777-424: 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
888-555: A bear shot by a hunter in the Northwest Territories confirmed the existence of naturally occurring and fertile grizzly–polar bear hybrids . Hybridization between reproductively isolated species often results in hybrid offspring with lower fitness than either parental. However, hybrids are not, as might be expected, always intermediate between their parents (as if there were blending inheritance), but are sometimes stronger or perform better than either parental lineage or variety,
999-428: 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
1110-508: A considerable seed yield advantage over open pollinated varieties. Hybrid seed dominates the commercial maize seed market in the United States, Canada and many other major maize-producing countries. In a hybrid, any trait that falls outside the range of parental variation (and is thus not simply intermediate between its parents) is considered heterotic. Positive heterosis produces more robust hybrids, they might be stronger or bigger; while
1221-504: A cross between a female horse and a male donkey, and the hinny, a cross between a female donkey and a male horse. Pairs of complementary types like the mule and hinny are called reciprocal hybrids. Polar bears and brown bears are another case of a hybridizing species pairs, and introgression among non-sister species of bears appears to have shaped the Ursidae family tree. Among many other mammal crosses are hybrid camels , crosses between
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#17332019308061332-501: A degree that none of the originally genetically distinct population remains. In agriculture and animal husbandry , the Green Revolution 's use of conventional hybridization increased yields by breeding high-yielding varieties . The replacement of locally indigenous breeds, compounded with unintentional cross-pollination and crossbreeding (genetic mixing), has reduced the gene pools of various wild and indigenous breeds resulting in
1443-432: 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
1554-602: A donkey as the father. A variety of mechanisms limit the success of hybridization, including the large genetic difference between most species. Barriers include morphological differences, differing times of fertility, mating behaviors and cues, and physiological rejection of sperm cells or the developing embryo. Some act before fertilization; others after it. In plants, some barriers to hybridization include blooming period differences, different pollinator vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and structural differences of
1665-508: 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
1776-548: A group of about 50 natural hybrids between Australian blacktip shark and the larger common blacktip shark was found by Australia's eastern coast in 2012. Russian sturgeon and American paddlefish were hybridized in captivity when sperm from the paddlefish and eggs from the sturgeon were combined, unexpectedly resulting in viable offspring. This hybrid is called a sturddlefish . The two genera Asymmetron and Branchiostoma are able to produce viable hybrid offspring, even if none have lived into adulthood so far, despite
1887-500: A hybrid is interpreted differently in animal and plant breeding, where there is interest in the individual parentage. In genetics , attention is focused on the numbers of chromosomes . In taxonomy, a key question is how closely related the parent species are. Species are reproductively isolated by strong barriers to hybridization, which include genetic and morphological differences, differing times of fertility, mating behaviors and cues, and physiological rejection of sperm cells or
1998-465: 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
2109-439: A long time, both intentionally for purposes such as biological control , and unintentionally, as with accidental escapes of individuals. Introductions can drastically affect populations, including through hybridization. There is a kind of continuum with three semi-distinct categories dealing with anthropogenic hybridization: hybridization without introgression, hybridization with widespread introgression (backcrossing with one of
2220-494: A mate among the species that raised it, instead of the species of its biological parents. Cagebird breeders sometimes breed bird hybrids known as mules between species of finch , such as goldfinch × canary . Among amphibians, Japanese giant salamanders and Chinese giant salamanders have created hybrids that threaten the survival of Japanese giant salamanders because of competition for similar resources in Japan. Among fish,
2331-492: 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
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#17332019308062442-449: 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
2553-400: 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
2664-432: A phenomenon called heterosis, hybrid vigour, or heterozygote advantage . This is most common with plant hybrids. A transgressive phenotype is a phenotype that displays more extreme characteristics than either of the parent lines. Plant breeders use several techniques to produce hybrids, including line breeding and the formation of complex hybrids. An economically important example is hybrid maize (corn), which provides
2775-400: 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,
2886-404: 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
2997-523: 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
3108-400: 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
3219-405: A species, such as between different breeds . Single cross hybrids result from the cross between two true-breeding organisms which produces an F1 hybrid (first filial generation). The cross between two different homozygous lines produces an F1 hybrid that is heterozygous ; having two alleles , one contributed by each parent and typically one is dominant and the other recessive . Typically,
3330-488: 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
3441-685: 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
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3552-519: 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
3663-506: 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
3774-447: 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
3885-515: 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,
3996-438: 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
4107-400: 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
4218-577: Is a hybrid of two Atlantic species, the spinner and striped dolphins . In 2019, scientists confirmed that a skull found 30 years earlier was a hybrid between the beluga whale and narwhal , dubbed the narluga . Hybridization between species is common in birds. Hybrid birds are purposefully bred by humans, but hybridization is also common in the wild. Waterfowl have a particularly high incidence of hybridization, with at least 60% of species known to produce hybrids with another species. Among ducks , mallards widely hybridize with many other species, and
4329-407: Is a hybridization test widely used in genetics to determine whether two separately isolated mutants that have the same (or similar) phenotype are defective in the same gene or in different genes (see Complementation (genetics) article). If a hybrid organism containing the genomes of two different mutant parental organisms displays a wild type phenotype, it is ordinarily considered that
4440-571: Is a particularly common mechanism for speciation in plants, and is now known to be fundamental to the evolutionary history of plants. Plants frequently form polyploids , individuals with more than two copies of each chromosome. Whole genome doubling has occurred repeatedly in plant evolution. When two plant species hybridize, the hybrid may double its chromosome count by incorporating the entire nuclear genome of both parents, resulting in offspring that are reproductively incompatible with either parent because of different chromosome counts. Human impact on
4551-458: 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
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4662-659: Is also more occasionally done in the livestock and pet trades; some well-known wild × domestic hybrids are beefalo and wolfdogs . Human selective breeding of domesticated animals and plants has also resulted in the development of distinct breeds (usually called cultivars in reference to plants); crossbreeds between them (without any wild stock ) are sometimes also imprecisely referred to as "hybrids". Hybrid humans existed in prehistory. For example, Neanderthals and anatomically modern humans are thought to have interbred as recently as 40,000 years ago. Mythological hybrids appear in human culture in forms as diverse as
4773-451: Is always sterile worker ants (and because ants are haplodiploid , unfertilized eggs become males). Without mating with males of the other species, the queens are unable to produce workers, and will fail to establish a colony of their own. Plant species hybridize more readily than animal species, and the resulting hybrids are fertile more often. Many plant species are the result of hybridization, combined with polyploidy , which duplicates
4884-414: 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
4995-412: 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,
5106-547: Is derived from Latin hybrida , used for crosses such as of a tame sow and a wild boar. The term came into popular use in English in the 19th century, though examples of its use have been found from the early 17th century. Conspicuous hybrids are popularly named with portmanteau words , starting in the 1920s with the breeding of tiger–lion hybrids ( liger and tigon ). From the point of view of animal and plant breeders, there are several kinds of hybrid formed from crosses within
5217-403: 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
5328-671: 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
5439-726: 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,
5550-403: 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
5661-468: 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
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#17332019308065772-586: 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,
5883-406: 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. Hybrid (biology) In biology , a hybrid
5994-427: Is that hybrid individuals can form a "bridge" transmitting potentially helpful genes from one species to another when the hybrid backcrosses with one of its parent species, a process called introgression . Hybrids can also cause speciation , either because the hybrids are genetically incompatible with their parents and not each other, or because the hybrids occupy a different niche than either parent. Hybridization
6105-622: Is the offspring resulting from combining the qualities of two organisms of different varieties , subspecies , species or genera through sexual reproduction . Generally, it means that each cell has genetic material from two different organisms, whereas an individual where some cells are derived from a different organism is called a chimera . Hybrids are not always intermediates between their parents such as in blending inheritance (a now discredited theory in modern genetics by particulate inheritance ), but can show hybrid vigor , sometimes growing larger or taller than either parent. The concept of
6216-515: Is the proper time to give up on a population that is becoming a hybrid swarm, or to try and save the still existing pure individuals. Once a population becomes a complete mixture, the goal becomes to conserve those hybrids to avoid their loss. Conservationists treat each case on its merits, depending on detecting hybrids within the population. It is nearly impossible to formulate a uniform hybridization policy, because hybridization can occur beneficially when it occurs "naturally", and when hybrid swarms are
6327-594: 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 ,
6438-523: 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
6549-638: The Minotaur , blends of animals, humans and mythical beasts such as centaurs and sphinxes , and the Nephilim of the Biblical apocrypha described as the wicked sons of fallen angels and attractive women. Hybridization between species plays an important role in evolution, though there is much debate about its significance. Roughly 25% of plants and 10% of animals are known to form hybrids with at least one other species. One example of an adaptive benefit to hybridization
6660-401: 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
6771-399: 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
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#17332019308066882-498: 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
6993-464: The sea urchin Strongylocentrotus purpuratus (female) and the sand dollar Dendraster excentricus (male). When two distinct types of organisms breed with each other, the resulting hybrids typically have intermediate traits (e.g., one plant parent has red flowers, the other has white, and the hybrid, pink flowers). Commonly, hybrids also combine traits seen only separately in one parent or
7104-406: 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
7215-438: The "suture region". It is at these regions that the subspecies were formed. Other hybrid zones have formed between described species of plants and animals. From the point of view of genetics, several different kinds of hybrid can be distinguished. A genetic hybrid carries two different alleles of the same gene , where for instance one allele may code for a lighter coat colour than the other. A structural hybrid results from
7326-503: The F1 generation is also phenotypically homogeneous, producing offspring that are all similar to each other. Double cross hybrids result from the cross between two different F1 hybrids (i.e., there are four unrelated grandparents). Three-way cross hybrids result from the cross between an F1 hybrid and an inbred line. Triple cross hybrids result from the crossing of two different three-way cross hybrids. Top cross (or "topcross") hybrids result from
7437-622: 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
7548-464: The True BI. The biotic index is then ranked for water quality and degree of organic pollution, as follows: This statistics -related article is a stub . You can help Misplaced Pages by expanding it . This arthropod -related article is a stub . You can help Misplaced Pages by expanding it . Species A species ( pl. : species) is a population of organisms in which any two individuals of
7659-405: 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
7770-570: 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
7881-404: 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 the concept of
7992-474: 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
8103-505: 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,
8214-428: 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
8325-511: The butterfly Limenitis arthemis has two major subspecies in North America, L. a. arthemis (the white admiral) and L. a. astyanax (the red-spotted purple). The white admiral has a bright, white band on its wings, while the red-spotted purple has cooler blue-green shades. Hybridization occurs between a narrow area across New England, southern Ontario, and the Great Lakes,
8436-426: The chromosomes. A few animal species are the result of hybridization. The Lonicera fly is a natural hybrid. The American red wolf appears to be a hybrid of the gray wolf and the coyote , although its taxonomic status has been a subject of controversy. The European edible frog is a semi-permanent hybrid between pool frogs and marsh frogs ; its population requires the continued presence of at least one of
8547-539: The chromosomes. Chromosome duplication allows orderly meiosis and so viable seed can be produced. Plant hybrids are generally given names that include an "×" (not in italics), such as Platanus × hispanica for the London plane, a natural hybrid of P. orientalis (oriental plane) and P. occidentalis (American sycamore). The parent's names may be kept in their entirety, as seen in Prunus persica × Prunus americana , with
8658-481: The common pheasant and hen golden pheasant ( Chrysolophus pictus ). Spurs are absent in hybrids of the former type, although present in both parents. Hybridization is greatly influenced by human impact on the environment, through effects such as habitat fragmentation and species introductions. Such impacts make it difficult to conserve the genetics of populations undergoing introgressive hybridization . Humans have introduced species worldwide to environments for
8769-433: 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
8880-400: The crossing of a top quality or pure-bred male and a lower quality female, intended to improve the quality of the offspring, on average. Population hybrids result from the crossing of plants or animals in one population with those of another population. These include interspecific hybrids or crosses between different breeds. In biology, the result of crossing of two populations is called
8991-444: The developing embryo . Some act before fertilization and others after it. Similar barriers exist in plants, with differences in flowering times, pollen vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and the structure of the chromosomes. A few animal species and many plant species, however, are the result of hybrid speciation , including important crop plants such as wheat , where
9102-407: The diverse Heliconius butterflies , but that is disputed. The two closely related harvester ant species Pogonomyrmex barbatus and Pogonomyrmex rugosus have evolved to depend on hybridization. When a queen fertilizes her eggs with sperm from males of her own species, the offspring is always new queens. And when she fertilizes the eggs with sperm from males of the other species, the offspring
9213-413: The doubling of chromosome sets, causing immediate genetic isolation. Hybridization may be important in speciation in some plant groups. However, homoploid hybrid speciation (not increasing the number of sets of chromosomes) may be rare: by 1997, only eight natural examples had been fully described. Experimental studies suggest that hybridization offers a rapid route to speciation, a prediction confirmed by
9324-611: The environment has resulted in an increase in the interbreeding between regional species, and the proliferation of introduced species worldwide has also resulted in an increase in hybridization. This has been referred to as genetic pollution out of concern that it may threaten many species with extinction. Similarly, genetic erosion from monoculture in crop plants may be damaging the gene pools of many species for future breeding. The conservation impacts of hybridization between species are highly debated. While hybridization could potentially threaten rare species or lineages by "swamping"
9435-435: 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
9546-429: The fact that early generation hybrids and ancient hybrid species have matching genomes, meaning that once hybridization has occurred, the new hybrid genome can remain stable. Many hybrid zones are known where the ranges of two species meet, and hybrids are continually produced in great numbers. These hybrid zones are useful as biological model systems for studying the mechanisms of speciation. Recently DNA analysis of
9657-560: The fusion of gametes that have differing structure in at least one chromosome, as a result of structural abnormalities . A numerical hybrid results from the fusion of gametes having different haploid numbers of chromosomes . A permanent hybrid results when only the heterozygous genotype occurs, as in Oenothera lamarckiana , because all homozygous combinations are lethal. In the early history of genetics, Hugo de Vries supposed these were caused by mutation . Genetic complementation
9768-507: The genetic relationships between ducks are further complicated by the widespread gene flow between wild and domestic mallards. One of the most common interspecific hybrids in geese occurs between Greylag and Canada geese ( Anser anser x Branta canadensis ). One potential mechanism for the occurrence of hybrids in these geese is interspecific nest parasitism , where an egg is laid in the nest of another species to be raised by non-biological parents. The chick imprints upon and eventually seeks
9879-630: The genetically "pure" individuals with hybrids, hybridization could also save a rare lineage from extinction by introducing genetic diversity. It has been proposed that hybridization could be a useful tool to conserve biodiversity by allowing organisms to adapt, and that efforts to preserve the separateness of a "pure" lineage could harm conservation by lowering the organisms' genetic diversity and adaptive potential, particularly in species with low populations. While endangered species are often protected by law, hybrids are often excluded from protection, resulting in challenges to conservation. The term hybrid
9990-401: The hybrids are more fit and have breeding advantages over the indigenous ecotype or species. These hybridization events can result from the introduction of non-native genotypes by humans or through habitat modification, bringing previously isolated species into contact. Genetic mixing can be especially detrimental for rare species in isolated habitats, ultimately affecting the population to such
10101-400: The loss of genetic diversity . Since the indigenous breeds are often well-adapted to local extremes in climate and have immunity to local pathogens, this can be a significant genetic erosion of the gene pool for future breeding. Therefore, commercial plant geneticists strive to breed "widely adapted" cultivars to counteract this tendency. Familiar examples of equid hybrids are the mule,
10212-452: The number of chromosomes has been doubled. A form of often intentional human-mediated hybridization is the crossing of wild and domesticated species. This is common in both traditional horticulture and modern agriculture ; many commercially useful fruits, flowers, garden herbs, and trees have been produced by hybridization. One such flower, Oenothera lamarckiana , was central to early genetics research into mutationism and polyploidy. It
10323-446: The number of individuals in each species (or genera) multiplied by the tolerance of the species is divided by the total number of specimens in the sample to determine the HBI value. H B I = Σ ( n i a i ) N {\displaystyle HBI={\frac {\Sigma (n_{i}a_{i})}{N}}} ; where n = number of specimens in taxa;
10434-478: 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
10545-420: The only remaining evidence of prior species, they need to be conserved as well. Regionally developed ecotypes can be threatened with extinction when new alleles or genes are introduced that alter that ecotype. This is sometimes called genetic mixing. Hybridization and introgression, which can happen in natural and hybrid populations, of new genetic material can lead to the replacement of local genotypes if
10656-438: The other (e.g., a bird hybrid might combine the yellow head of one parent with the orange belly of the other). Interspecific hybrids are bred by mating individuals from two species, normally from within the same genus. The offspring display traits and characteristics of both parents, but are often sterile , preventing gene flow between the species. Sterility is often attributed to the different number of chromosomes between
10767-585: 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
10878-573: The parent species), and hybrid swarms (highly variable populations with much interbreeding as well as backcrossing with the parent species). Depending on where a population falls along this continuum, the management plans for that population will change. Hybridization is currently an area of great discussion within wildlife management and habitat management. Global climate change is creating other changes such as difference in population distributions which are indirect causes for an increase in anthropogenic hybridization. Conservationists disagree on when
10989-532: The parent species. Cave paintings indicate that the European bison is a natural hybrid of the aurochs and the steppe bison . Plant hybridization is more commonplace compared to animal hybridization. Many crop species are hybrids, including notably the polyploid wheats : some have four sets of chromosomes (tetraploid) or six (hexaploid), while other wheat species have (like most eukaryotic organisms) two sets ( diploid ), so hybridization events likely involved
11100-506: The parents' common ancestor living tens of millions of years ago. Among insects, so-called killer bees were accidentally created during an attempt to breed a strain of bees that would both produce more honey and be better adapted to tropical conditions. It was done by crossing a European honey bee and an African bee . The Colias eurytheme and C. philodice butterflies have retained enough genetic compatibility to produce viable hybrid offspring. Hybrid speciation may have produced
11211-674: 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
11322-637: The point of view of taxonomy , hybrids differ according to their parentage. Hybrids between different subspecies (such as between the dog and Eurasian wolf ) are called intra-specific hybrids. Interspecific hybrids are the offspring from interspecies mating ; these sometimes result in hybrid speciation. Intergeneric hybrids result from matings between different genera, such as between sheep and goats . Interfamilial hybrids, such as between chickens and guineafowl or pheasants , are reliably described but extremely rare. Interordinal hybrids (between different orders) are few, but have been engineered between
11433-487: 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
11544-573: 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
11655-508: 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
11766-529: 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
11877-502: 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
11988-470: The term negative heterosis refers to weaker or smaller hybrids. Heterosis is common in both animal and plant hybrids. For example, hybrids between a lion and a tigress (" ligers ") are much larger than either of the two progenitors, while " tigons " (lioness × tiger) are smaller. Similarly, the hybrids between the common pheasant ( Phasianus colchicus ) and domestic fowl ( Gallus gallus ) are larger than either of their parents, as are those produced between
12099-400: The two parental mutant organisms are defective in different genes. If the hybrid organism displays a distinctly mutant phenotype, the two mutant parental organisms are considered to be defective in the same gene. However, in some cases the hybrid organism may display a phenotype that is only weakly (or partially) wild-type, and this may reflect intragenic (interallelic) complementation. From
12210-462: The two species. For example, donkeys have 62 chromosomes , horses have 64 chromosomes, and mules or hinnies have 63 chromosomes. Mules, hinnies, and other normally sterile interspecific hybrids cannot produce viable gametes, because differences in chromosome structure prevent appropriate pairing and segregation during meiosis , meiosis is disrupted, and viable sperm and eggs are not formed. However, fertility in female mules has been reported with
12321-537: 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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