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| {{Other uses|Hybrid (disambiguation){{!}}Hybrid}}
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| [[Image:Liger.jpg|thumb|Hercules, a "[[Liger]]", a Lion/Tiger hybrid.]]
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| In [[biology]] and specifically, [[genetics]], the term '''hybrid''' has several meanings, all referring to the offspring of [[sexual reproduction]].<ref>Rieger, R.; Michaelis A.; Green, M. M. (1991). ''Glossary of Genetics'' (5th ed.). Springer-Verlag. ISBN 0-387-52054-6 page 256</ref>
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| #In general usage, '''hybrid''' is synonymous with [[Zygosity#Heterozygous|heterozygous]]: any offspring resulting from the mating of two genetically distinct individuals
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| #a '''genetic hybrid''' carries two different [[allele]]s of the same [[gene]]
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| #a '''structural hybrid''' results from the fusion of [[gamete]]s that have differing structure in at least one [[chromosome]], as a result of [[Chromosome abnormality#Structural abnormalities|structural abnormalities]]
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| #a '''numerical hybrid''' results from the fusion of gametes having different [[Ploidy|haploid numbers of chromosomes]]
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| #a '''permanent hybrid''' is a situation where only the heterozygous [[genotype]] occurs, because all homozygous combinations are lethal.
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| From a [[Taxonomy (biology)|taxonomic]] perspective, hybrid refers to:
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| #Offspring resulting from the interbreeding between two animals or plants of different [[species]].<ref>Keeton, William T. 1980. Biological science. New York: Norton. ISBN 0-393-95021-2 page A9.</ref> See also [[hybrid speciation]].
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| #Hybrids between different [[subspecies]] within a species (such as between the [[Bengal tiger]] and [[Siberian tiger]]) are known as '''intra-specific''' hybrids. Hybrids between different species within the same genus (such as between [[lion]]s and [[tiger]]s) are sometimes known as '''interspecific''' hybrids or crosses. Hybrids between different genera (such as between [[Domestic sheep|sheep]] and [[goat]]s) are known as '''intergeneric''' hybrids. Extremely rare '''interfamilial''' hybrids have been known to occur (such as the [[chicken hybrids|guineafowl hybrids]]).<ref name="ghigi">Ghigi A. 1936. "Galline di faraone e tacchini" Milano (Ulrico Hoepli)</ref> No '''interordinal''' (between different orders) animal hybrids are known.
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| #The third type of hybrid consists of crosses between [[population]]s, [[breed]]s or [[cultivar]]s within a single species. This meaning is often used in plant and animal breeding, where hybrids are commonly produced and selected because they have desirable characteristics not found or inconsistently present in the parent individuals or populations. This flow of genetic material between populations or races is often called hybridization.
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| ==Etymology==
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| According to the [[Oxford English Dictionary]], the word is derived from Latin ''hybrida'', meaning the "offspring of an tame sow and a wild boar", "child of a freeman and slave", etc.<ref>[http://www.askoxford.com/concise_oed/hybrid?view=uk askoxford.com]</ref> The term entered into popular use in English in the 19th century, though examples of its use have been found from the early 17th century.<ref>Oxford English Dictionary Online, Oxford University Press 2007.</ref>
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| ==Types of hybrids==
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| Depending on the parents, there are a number of different types of hybrids;<ref>Wricke, Gunter, and Eberhard Weber. 1986.
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| ''Quantitative genetics and selection in plant breeding.'' Berlin: W. de Gruyter. Page 257.</ref>
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| *'''Single cross hybrids''' — result from the cross between two [[true breeding organism]]s and produces an F1 generation called an [[F1 hybrid]] (F1 is short for Filial 1, meaning "first offspring"). 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 [[Dominance relationship#Dominant allele|dominant]] and the other [[Dominance relationship#Recessive allele|recessive]]. Typically, the F1 generation is also [[phenotype|phenotypically]] homogeneous, producing offspring that are all similar to each other.
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| *'''Double cross hybrids''' — result from the cross between two different F1 hybrids.<ref>J. O. Rawlings, C. Clark Cockerham ''Analysis of Double Cross Hybrid Populations.'' J. O. Rawlings, C. Clark Cockerham Biometrics, Vol. 18, No. 2 (Jun., 1962), pp. 229-244 {{doi|10.2307/2527461}}</ref>
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| *'''Three-way cross hybrids''' — result from the cross between one parent that is an F1 hybrid and the other is from an inbred line.<ref>Roy, Darbeshwar. 2000. ''Plant breeding analysis and exploitation of variation''. Pangbourne, UK: Alpha Science International. Page 446.</ref>
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| *'''Triple cross hybrids''' — result from the crossing of two different three-way cross hybrids.
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| *'''Population hybrids''' — result from the crossing of plants or animals in a [[population]] with another population. These include crosses between organisms such as interspecific hybrids or crosses between different races.
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| *'''Stable hybrid''' - a [[horticultural]] term which typically refers to an [[annual plant]] that, if grown and bred in a small [[monoculture]] free of external [[pollen]] (e.g., an air-filtered greenhouse) will produce offspring that are "true to type" with respect to [[phenotype]]; i.e., a [[true breeding organism]].<ref>Toogood, A.(ed.) (1999). ''Plant Propagation'' (1st American ed.). American Horticultural Society. ISBN 0-7894-5520-X page 21</ref>
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| *'''Hybrid species''' - results from hybrid populations evolving reproductive barriers against their parent species through [[hybrid speciation]].<ref name=Arnold>{{cite book|last=Arnold|first=M.L.|title=Natural Hybridization and Evolution|year=1996|publisher=Oxford University Press|location=New York|isbn=978-0-19-509975-1|pages=232}}</ref>
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| ==Interspecific hybrids==
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| Interspecific hybrids are bred by mating two species, normally from within the same [[genus]]. The offspring display traits and characteristics of both parents. The offspring of an interspecific cross are very often [[Sterility (physiology)|sterile]]; thus, hybrid sterility prevents the movement of genes from one species to the other, keeping both species distinct.<ref>Keeton, William T. 1980. ''Biological science''. New York: Norton. ISBN 0-393-95021-2 Page 800</ref> Sterility is often attributed to the different number of chromosomes the two species have, for example [[donkey]]s have 62 [[chromosomes]], while [[horse]]s have 64 chromosomes, and [[mule]]s and [[hinny|hinnies]] have 63 chromosomes. Mules, hinnies, and other normally sterile interspecific hybrids cannot produce viable [[gamete]]s, 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 a donkey as the father.<ref>{{cite web|url=http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=3378453&dopt=Abstract|title=McBeath S, Tan PP, Bai Q, Speed RM.|year=1988}}</ref>
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| Most often other processes occurring in plants and animals keep gametic isolation and species distinction. Species often have different mating or courtship patterns or behaviors, the breeding seasons may be distinct and even if mating does occur antigenic reactions to the sperm of other species prevent fertilization or embryo development. Hybridisation is much more common among organisms that spawn indiscriminately, like [[soft corals]] and among [[plant]]s.
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| While it is possible to predict the genetic composition of a backcross ''on average'', it is not possible to accurately predict the composition of a particular backcrossed individual, due to random segregation of chromosomes. In a species with two pairs of chromosomes, a twice backcrossed individual would be predicted to contain 12.5% of one species' genome (say, species A). However, it may, in fact, still be a 50% hybrid if the chromosomes from species A were lucky in two successive segregations, and meiotic crossovers happened near the [[telomere]]s. The chance of this is fairly high: <math>\left(\frac{1}{2}\right)^{(2 \times 2)} = \frac{1}{16}</math> (where the "two times two" comes about from two rounds of meiosis with two chromosomes); however, this probability declines markedly with chromosome number and so the actual composition of a hybrid will be increasingly closer to the predicted composition.
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| Hybrids are often named by the [[portmanteau]] method, combining the names of the two parent species. For example, a [[zeedonk]] is a cross between a [[zebra]] and a [[donkey]]. Since the traits of hybrid offspring often vary depending on which species was mother and which was father, it is traditional to use the father's species as the first half of the portmanteau. For example, a [[liger]] is a cross between a male [[lion]] and a female [[tiger]], while a [[tiglon]] is a cross between a male tiger and a female lion.
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| ===Domestic and wild hybrids===
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| {{Unreferenced section|date=January 2010}}
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| Hybrids between domesticated and wild animals in particular may be problematic. Breeders of [[domesticated]] species discourage [[crossbreeding]] with wild species, unless a deliberate decision is made to incorporate a trait of a wild ancestor back into a given breed or strain. Wild populations of animals and plants have evolved naturally over millions of years through a process of [[natural selection]] in contrast to human controlled [[selective breeding]] or [[artificial selection]] for desirable traits from the human point of view. Normally, these two methods of reproduction operate independently of one another. However, an intermediate form of selective breeding, wherein animals or plants are bred by humans, but with an eye to adaptation to natural region-specific conditions and an acceptance of natural selection to weed out undesirable traits, created many ancient domesticated breeds or types now known as [[landrace]]s.
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| Many times, [[domesticated]] species live in or near areas which also still hold naturally evolved, region-specific [[Wildlife|wild]] ancestor species and subspecies. In some cases, a domesticated species of plant or animal may become [[feral]], living wild. Other times, a wild species will come into an area inhabited by a domesticated species. Some of these situations lead to the creation of hybridized plants or animals, a cross between the native species and a domesticated one. This type of crossbreeding, termed [[genetic pollution]] by those who are concerned about preserving the genetic base of the wild species, has become a major concern. Hybridization is also a concern to the breeders of purebred species as well, particularly if the gene pool is small and if such crossbreeding or hybridization threatens the genetic base of the domesticated purebred population.
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| The concern with genetic pollution of a wild population is that hybridized animals and plants may not be as genetically strong as naturally evolved region specific wild ancestors [[wildlife]] which can survive without human [[animal husbandry|husbandry]] and have high immunity to natural diseases. The concern of purebred breeders with wildlife hybridizing a domesticated species is that it can coarsen or degrade the specific qualities of a breed developed for a specific purpose, sometimes over many generations. Thus, both purebred breeders and wildlife biologists share a common interest in preventing accidental hybridization.
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| ==Hybrid species==
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| {{Main|Hybrid speciation}}
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| While not very common, a few animal species have been recognized as being the result of hybridization. The [[Lonicera fly]] is an example of a novel [[animal]] [[species]] that resulted from natural hybridization. The American [[red wolf]] appears to be a hybrid species between [[gray wolf]] and [[coyote]].<ref name=Hybrid>{{cite web|last=Esch|first=Mary|title=Study: Eastern wolves are hybrids with coyotes|url=http://www.google.com/hostednews/ap/article/ALeqM5hIGn3MyVkj3dkd4W2UljCK0y4sVg?docId=787311a8475a4d3aae6e8498aa2edd80|publisher=AP News|accessdate=1 June 201`}}</ref> The European [[edible frog]] appears to be a species, but is actually a semi-permanent hybrid between [[pool frog]]s and [[marsh frog]]s. The edible frog population is dependent on the presence of at least one of the parents species to be maintained.<ref name="Frost-etal2006">Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History. Number 297. New York. Issued March 15, 2006.</ref>
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| Hybrid species of [[plant]]s are much more common than animals. Many of the crop species are hybrids, and hybridization appear to be an important factor in speciation in some plant groups.
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| ==Examples of hybrid animals==
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| [[Image:Zeedonk 800.jpg|thumb|A "[[Zonkey]]", a zebra/donkey hybrid.]]
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| [[Image:Jaglion.jpg|thumb|A "[[Jaglion]]", a Jaguar/Lion hybrid.]]
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| [[File:Goldfinch Canary hybrid.JPG|thumb|A [[Mule (bird)|mule]], a Domestic Canary/Goldfinch hybrid.]]
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| ===Mammalian hybrids===
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| * Equid hybrids
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| ** [[Mule]], a cross of female [[horse]] and a male [[donkey]].
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| ** [[Hinny]], a cross between a female donkey and a male horse. [[Mule]] and [[hinny]] are examples of reciprocal hybrids.
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| ** [[Zebroid]]s
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| *** Zeedonk or Zonkey, a [[zebra]]/[[donkey]] cross.
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| *** Zorse, a zebra/horse cross
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| *** Zony or Zetland, a zebra/[[pony]] cross ("zony" is a generic term; "zetland" is specifically a hybrid of the Shetland pony breed with a zebra)
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| * [[Bovid hybrid]]s
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| ** [[Dzo]], zo or yakow; a cross between a domestic [[cattle|cow/bull]] and a [[yak]].
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| ** [[Beefalo]], a cross of an [[American bison]] and a domestic [[cattle|cow]]. This is a fertile breed; this along with genetic evidence has caused them to be recently reclassified into the same genus, ''Bos''.{{citation needed|date=December 2010}}
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| ** [[Zubron]], a hybrid between [[wisent]] ([[European bison]]) and domestic [[cattle|cow]].
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| * [[Sheep-goat hybrid]] is the cross between a [[sheep]] and a [[goat]], which belong to different genera. <!--the geep is a chimera not a hybrid--->
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| * [[Ursid hybrid]]s, such as the [[grizzly-polar bear hybrid]], occur between black bears, brown bears, and polar bears.
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| * [[Felid hybrid]]s
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| ** [[Savannah (cat)|Savannah cats]] are the hybrid cross between an [[serval|African serval cat]] and a [[domestic cat]]
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| **A hybrid between a [[Bengal Tiger|Bengal tiger]] and a [[Siberian Tiger|Siberian tiger]] is an example of an '''intra-specific''' hybrid.
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| ** [[Liger]]s and [[Tiglon]]s (crosses between a [[lion]] and a [[tiger]]) and other [[Panthera hybrid]]s such as the [[lijagulep]]. Various other wild cat crosses are known involving the [[lynx]], [[bobcat]], [[leopard]], [[serval]], etc.
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| *** [[Liliger]]s are the hybrid cross between a male [[lion]] and a [[liger|ligress]].
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| ** [[Bengal cat]], a cross between the Asian [[leopard cat]] and the domestic cat, one of many hybrids between the domestic cat and wild cat species. The domestic [[cat]], African wild cat and European wildcat may be considered variant populations of the same species (''Felis silvestris''), making such crosses non-hybrids.
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| * Fertile [[canid hybrid]]s occur between [[coyote]]s, [[Wolf|wolves]], [[dingo]]es, [[jackal]]s and [[Dog|domestic dogs]].
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| * Hybrids between [[Black rhinoceros|black]] and [[white rhinoceros]]es have been recognized.
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| * Hybrid camel, a cross between a [[two-humped, Bactrian camel]] and a [[single-humped, dromedary camel]], just like the mule a more powerful creature than its parents.<ref>R.W. Bulliet ''The Camel and the Wheel'' (Cambridge Mass. '75) 164-75</ref>
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| * [[Cama (animal)|Cama]], a cross between a [[camel]] and a [[llama]], also an intergeneric hybrid.
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| * [[Wholphin]], a fertile but very rare cross between a [[false killer whale]] and a [[bottlenose dolphin]].
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| * At [[Chester Zoo]] in the United Kingdom, a cross between an [[African elephant]] (male) and an [[Asian elephant]] (female). The male calf was named [[Motty]]. It died of intestinal infection after twelve days.
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| *[[Homininae]] hybrids
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| **Hybrids between [[modern humans]] and with "at least two groups" of [[ancient humans]]: [[Neanderthals]] and [[Denisovans]]<ref name="NYT-01302012">{{cite news |last=Mitchell |first=Alanna |title=DNA Turning Human Story Into a Tell-All |url=http://www.nytimes.com/2012/01/31/science/gains-in-dna-are-speeding-research-into-human-origins.html |date=January 30, 2012 |publisher=[[NYTimes]] |accessdate=January 31, 2012}}</ref>
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| **[[Humanzee]], hypothetical cross between a [[human]] and a [[chimpanzee]]
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| **Humilla, hypothetical cross between a [[human]] and a [[gorilla]]{{citation needed|date=December 2013}}
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| **[[Koolakamba]], alleged cross between a [[gorilla]] and a [[chimpanzee]]
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| ===Avian hybrids===
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| {{see also|Bird hybrid}}
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| * Hybrids between [[spotted owl]]s and [[barred owl]]s
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| * Cagebird breeders sometimes breed hybrids between species of [[finch]], such as [[Carduelis|goldfinch]] × [[Domestic Canary|canary]]. These birds are known as [[Mule (bird)|mules]].
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| * The [[Perlin (falconry)|Perlin]] is a [[Peregrine falcon]] - [[Merlin (bird)|Merlin]] hybrid.
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| * [[Gamebird hybrids]], hybrids between gamebirds and domestic fowl, including [[chickens]], [[guineafowl]] and [[peafowl]], interfamilial hybrids.
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| * Numerous [[Macaw#Hybrids|macaw hybrids]] and [[Hybrid Lovebird|lovebird hybrids]] are also known in aviculture.
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| * [[Red kite]] × [[black kite]]: five bred unintentionally at a [[falconry]] center in England. (It is reported{{Weasel-inline|date=May 2010|reason=If it has been reported, then that's a source to cite.}} that the black kite (the male) refused female black kites but mated with two female red kites.)
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| * The [[mulard]] duck, hybrid of the domestic [[pekin duck]] and domesticated [[muscovy duck]]s.
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| * In Australia, New Zealand and other areas where the [[Pacific Black Duck]] occurs, it is hybridised by the much more aggressive introduced [[Mallard]]. This is a concern to wildlife authorities throughout the affected area, as it is seen as [[Genetic pollution]] of the Black Duck gene pool.
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| * [[Hybridisation in gulls]] is a reasonably frequent occurrence in the wild.
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| ===Reptilian hybrids===
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| * [[Hybrid Iguana]], a single‐cross hybrid resulting from natural interbreeding between [[male]] [[marine iguana]]s and [[female]] [[Galapagos Land Iguana|land iguana]]s since the late 2000s.
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| * Crestoua, a cross between a Rhacodactylus Ciliatus ([[crested gecko]]) and a Rhacodactylus Chahoua.
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| * Colubrid snakes of the tribe [[Lampropeltini]] have been shown to produce fertile hybrid offspring.
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| * Hybridization between the endemic [[Cuban crocodile]] (''Crocodilus rhombifer'') and the widely distributed [[American crocodile]] (''Crocodilus acutus'') is causing conservation problems for the former species as a threat to its genetic integrity.<ref>http://www.savingwildplaces.com/swp-home/swp-crocodile/8287793?preview=&psid=&ph=class%2525253dawc-148772</ref>{{Clarify|date=May 2010|reason=This is not a proper reference citation. Use Template:Cite web or similar to provide source details.}}
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| * [[Saltwater crocodile]]s (''Crocodylus porosus'') have mated with [[Siamese crocodile]]s (''Crocodylus siamensis'') in captivity producing offspring which in many cases have grown over {{convert|20|ft|m|abbr=off}} in length. It is likely that wild hybridization occurred historically in parts of southeast Asia.
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| * Many species of [[Boidae|boas]] and [[Python (genus)|pythons]] are known to produce hybrids,such as carball (a cross between a ball python and a carpet python) or a bloodball (a cross between a blood python and a ball python) however, most of these only occur in captivity. Contrary to popular belief, boa-python hybrids are not possible due to their differing reproductive functions. Boas only produce hybrids with other species of boas, and pythons only produce hybrids with other species of pythons.
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| ===Piscine hybrids===
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| * [[Blood parrot cichlid]], which is probably created by crossing a red head cihclid and a Midas cichlid or red devil cichlid
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| * A group of about 50 hybrids between [[Australian blacktip shark]] and the larger [[Blacktip shark|common blacktip shark]] was found by Australia's East Coast in 2012. This is the only known case of hybridization in sharks.<ref>{{cite news|last=Voloder|first=Dubravka|title=Print Email Facebook Twitter More World-first hybrid sharks found off Australia|url=http://www.abc.net.au/news/2012-01-03/hybrid-sharks-found-off-australia/3757226?section=nsw|accessdate=5 January 2012|newspaper=ABC News|date=3 January 1012}}</ref>
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| * [[Silver bream]] and [[Common bream]] commonly produce sterile hybrids.
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| * [[Tiger muskie]] is a sterile hybrid between [[Northern pike]] and [[Muskellunge]].
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| ===Insect hybrids===
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| * [[Africanized bee|Killer bees]] were created in an attempt to breed tamer and more manageable bees. This was done by crossing a [[european honey bee]] and an [[african bee]], but instead the offspring became more aggressive and highly defensive.
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| * The ''[[Colias eurytheme]]'' and ''[[Colias philodice|C. philodice]]'' butterflies have retained enough genetic compatibility to produce viable hybrid offspring.<ref>Grula, John W., and Orley R. Taylor. (1980) The Effect of X-Chromosome Inheritance on Mate-Selection Behavior in the Sulfur Butterflies, Colias eurytheme and C. Philodice. Evolution 34(4):688-95</ref>
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| Hybrids should not be confused with [[Chimera (genetics)|genetic chimeras]] such as that between [[Domestic sheep|sheep]] and [[goat]] known as the [[Sheep–goat chimera|geep]]. Wider interspecific hybrids can be made via [[in vitro fertilization]] or somatic hybridization, however the resulting cells are not able to develop into a full organism. An example of interspecific hybrid cell lines is [[humster]] (hamster × human) cells.
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| ==Hybrid plants==
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| <!--[[Template:Infobox Cultivar]] links to this section, so if updates are made to section names, please remember to update the infobox-->
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| Many hybrids are created by humans, but natural hybrids occur as well. Plant species hybridize more readily than animal species, and the resulting hybrids are more often fertile hybrids and may reproduce, though there still exist sterile hybrids and selective hybrid elimination where the offspring are less able to survive and are thus eliminated before they can reproduce. A number of plant species are the result of hybridization and [[polyploidy]] with many plant species easily cross pollinating and producing viable seeds, the distinction between each species is often maintained by geographical isolation or differences in the flowering period. Since plants hybridize frequently without much work, they are often created by humans in order to produce improved plants. These improvements can include the production of more or improved seeds, fruits or other plant parts for consumption, or to make a plant more winter or heat hardy or improve its growth and/or appearance for use in horticulture. Much work is now being done with hybrids to produce more disease resistant plants for both agricultural and horticultural crops. In many groups of plants hybridization has been used to produce larger and more showy flowers and new flower colors.
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| [[Image:Trilliumhybrid2.jpg|right|thumb|200px|A sterile ''Trillium'' hybrid between ''[[Trillium cernuum]]'' and ''[[Trillium grandiflorum]]''{{Or|date=March 2011}}]]
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| Many plant genera and species have their origins in polyploidy. Autopolyploidy results from the sudden multiplication in the number of chromosomes in typical normal populations caused by unsuccessful separation of the chromosomes during meiosis. Tetraploids (plants with four sets of chromosomes rather than two) are common in a number of different groups of plants and over time these plants can differentiate into distinct species from the normal diploid line. In ''Oenothera lamarchiana'' the diploid species has 14 chromosomes, this species has spontaneously given rise to plants with 28 chromosomes that have been given the name ''Oenothera gigas''. When hybrids are formed between the tetraploids and the diploid population, the resulting offspring tend to be sterile triploids, thus effectively stopping the intermixing of genes between the two groups of plants (unless the diploids, in rare cases, produce unreduced gametes).
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| [[File:Lily Lilium 'Citronella' Flower.jpg|thumb|right|200px|An ornamental lily hybrid known as ''Lilium'' 'Citronella'<ref>http://www.pacificbulbsociety.org/pbswiki/index.php/LiliumHybrids</ref>]]
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| Another form of polyploidy called [[allopolyploid]]y occurs when two different species mate and produce polyploid hybrids. Usually the typical chromosome number is doubled, and the four sets of chromosomes can pair up during meiosis, thus the polyploids can produce offspring. Usually, these offspring can mate and reproduce with each other but cannot back-cross with the parent species. Allopolyploids may be able to adapt to new habitats that neither of their parent species inhabited.
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| Sterility in a non-polyploid hybrid is often a result of [[chromosome]] number; if parents are of differing chromosome pair number, the offspring will have an odd number of [[chromosomes]], leaving them unable to produce chromosomally balanced [[gametes]].<ref>[http://www.colorado.edu/MCDB/MCDB2150Fall/notes00/L0033.html University of Colorado Principles of Genetics (MCDB 2150) Lecture 33: Chromosomal changes: Monosomy, Trisomy, Polyploidy, Structural Changes]</ref> While this is undesirable in a crop such as [[wheat]], where growing a crop which produces no seeds would be pointless, it is an attractive attribute in some fruits. [[Triploid]] [[banana]]s and [[watermelon]]s are intentionally bred because they produce no seeds (and are [[parthenocarpy|parthenocarpic]]).
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| ===Heterosis===
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| {{Main| heterosis }}
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| Hybrids are sometimes stronger than either parent variety, a phenomenon most common with plant hybrids, which when present is known as ''[[heterosis|hybrid vigor]]'' (heterosis) or heterozygote advantage.<ref>''Evaluating the utility of Arabidopsis thaliana as a model for understanding heterosis in hybrid crops''
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| Journal Euphytica
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| Publisher Springer Netherlands
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| ISSN 0014-2336 (Print) 1573-5060 (Online)
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| Issue Volume 156, Numbers 1-2 / July, 2007
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| DOI 10.1007/s10681-007-9362-1
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| Pages 157-171</ref> A [[Transgressive segregation|transgressive phenotype]] is a [[phenotype]] displaying more extreme characteristics than either of the parent lines.<ref name="Rieseberg1999">{{cite journal | title = Transgressive segregation, adaptation and speciation | journal = Heredity | date = July 1999 | first = Loren H. | last = Rieseberg | pmid = 10583537 | coauthors = Margaret A. Archer and Robert K. Wayne | volume = 83 | pages = 363–372| doi = 10.1038/sj.hdy.6886170 | issue=4}}</ref> [[Plant breeding|Plant breeders]] make use of a number of techniques to produce hybrids, including line breeding and the formation of complex hybrids. An economically important example is hybrid [[maize]] (corn), which provides 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.<ref>Smith C. Wayne. ''Corn: Origin, History, Technology, and Production.'' Wiley Series in Crop Science, 2004, p. 332.</ref>
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| ===Examples of plant hybrids===
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| The multiplication symbol × (not italicised) indicates a hybrid in the Latin [[binomial nomenclature]]. Placed before the binomial it indicates a hybrid between species from different genera (intergeneric hybrid):-
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| *× ''[[Fatshedera lizei]]'', a hybrid between ''[[Hedera helix]]'' and ''[[Fatsia japonica]]''
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| *× ''[[Heucherella]]'', a hybrid genus between ''[[Heuchera]]'' and ''[[Tiarella]]''
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| * × ''Philageria veitchii'' is a hybrid between ''[[Lapageria rosea]]'' and ''[[Philesia magellanica]]''; it is more similar in appearance to the former
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| *[[Leyland cypress]], [× ''Cupressocyparis leylandii''] hybrid between [[Monterey cypress]] and [[Nootka cypress]]
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| * × ''[[Urceocharis]]'', a hybrid between ''[[Eucharis (plant)|Eucharis]]'' and ''[[Urceolina]]''
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| Interspecific plant hybrids include:
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| *''Dianthus'' × ''allwoodii'' (''[[Dianthus caryophyllus]]'' × ''[[Dianthus plumarius]]'')
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| *[[Limequat]] ''Citrus'' × ''floridana'', [[key lime]] ''Citrus aurantiifolia'' and [[kumquat]] ''Citrus japonica'' hybrid
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| *[[Loganberry]] ''Rubus'' × ''loganobaccus'', a hybrid between [[raspberry]] ''[[Rubus idaeus]]'' and [[blackberry]] ''[[Rubus ursinus]]''
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| *[[London plane]] (''[[Platanus orientalis]]'' × ''[[Platanus occidentalis]]''), thus forming ''Platanus'' × ''acerifolia''
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| *''Magnolia'' × ''alba'' (''[[Magnolia champaca]]'' × ''[[Magnolia montana]]'')
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| *[[Peppermint]], a hybrid between [[spearmint]] and [[water mint]]
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| *''Quercus'' × ''warei'' (''[[Quercus robur]]'' × ''[[Quercus bicolor]]'') 'Nadler' (marketed in the United States under the trade name [[Kindred Spirit hybrid oak]])
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| *[[Tangelo]], a hybrid of a [[Mandarin orange]] and a [[pomelo]] which may have been developed in Asia about 3,500 years ago
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| *[[Triticale]], a [[wheat]]–[[rye]] hybrid
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| *[[Wheat]]; most modern and ancient wheat breeds are themselves hybrids. Bread wheat is a hexaploid hybrid of three wild grasses; [[durum]] (pasta) wheat is a tetraploid hybrid of two wild grasses
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| *[[Grapefruit]], hybrid between a [[pomelo]] and the Jamaican sweet [[Orange (fruit)|orange]]
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| Some natural hybrids:
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| *''[[Iris albicans]]'', a sterile hybrid which spreads by rhizome division
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| *[[Oenothera|Evening primrose]], a flower which was the subject of famous experiments by [[Hugo de Vries]] on [[polyploidy]] and [[diploidy]]
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| ==Hybrids in nature==
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| Hybridisation between two closely related species is actually a common occurrence in nature but is also being greatly influenced by [[Human impact on the environment|anthropogenic]] changes as well.<ref name="Allendorf 2001 613–622">{{cite journal|last=Allendorf|first=Fred W.|coauthors=R.F. Leary, P. Spruell, & J.K. Wenburg|title=The problems with hybrids: setting conservation guidelines|journal=TRENDS in Ecology & Evolution|date=November 2001|volume=16|issue=11|pages=613–622|doi=10.1016/S0169-5347(01)02290-X}}</ref> Hybridization is a naturally occurring genetic process where individuals from two genetically distinct populations mate.<ref>{{cite book|last=Allendorf|first=Fred|title=Conservation and the Genetics of Populations|year=2007|publisher=Blackwell Publishing|location=Malden, MA|pages=534}}</ref> As stated above, it can occur both intraspecifically, between different distinct populations within the same species, and interspecifically, between two different species. Hybrids can be either sterile/not viable or viable/fertile. This affects the kind of effect that this hybrid will have on its and other populations that it interacts with.<ref name="Allendorf 2007 421–448">{{cite book|last=Allendorf|first=Fred|title=Conservation and the Genetics of Populations|year=2007|publisher=Blackwell Publishing|location=Malden, MA|pages=421–448}}</ref> Many [[hybrid zone]]s 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]] ([[Hybrid speciation]]). Recently DNA analysis of a bear shot by a hunter in the [[North West Territories]] confirmed the existence of naturally-occurring and fertile [[grizzly–polar bear hybrid]]s.<ref>{{cite news|url=http://news.bbc.co.uk/2/hi/science/nature/4766217.stm|publisher=[[BBC News]]|date=13 May 2006|title=Hybrid bear shot dead in Canada}}</ref> There have been reports of similar supposed hybrids, but this is the first to be confirmed by DNA analysis. In 1943, Clara Helgason described a male bear shot by hunters during her childhood. It was large and off-white with hair all over its paws. The presence of hair on the bottom of the feet suggests it was a natural hybrid of Kodiak and Polar bear.
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| ===Anthropogenic hybridization===
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| Changes to the environment caused by humans, such as [[Habitat fragmentation|fragmentation]] and [[Introduced species]], are becoming more widespread.<ref>{{cite journal|last=Ehrlich|first=Paul|coauthors=John Holdren|title=Impact of population Growth|journal=Science|date=26 March 1971|volume=171|pages=1212–1216| doi=10.1126/science.171.3977.1212|issue=3977 }}</ref> This increases the challenges in managing certain populations that are experiencing [[introgression]], and is a focus of [[conservation genetics]].
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| ====Introduced species and habitat fragmentation====
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| Humans have been [[Introduced species|introducing species]] world wide to environments for a long time both directly such as establishing a population to be used as a [[biological control]] and indirectly such as accidental escapes of individuals out of agriculture. This causes drastic global effects on various populations with hybridization being one of the reasons introduced species can be so detrimental.<ref name="Allendorf 2007 421–448"/><ref>{{cite journal|last=Vitousek|first=Peter|coauthors=Carla M. D'Antonio, Lloyd L. Loope, Marcel Rejmánek, & Randy Westbrooks|title=Introduced Species: A Significant Component of Human-cause Global Change|journal=New Zealand Journal of Ecology|year=1997|volume=21|issue=1|pages=1–16}}</ref>
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| When habitats become broken apart, one of two things can occur, genetically speaking. The first is that populations that were once connected can be cut off from one another, preventing their genes from interacting. Occasionally, this will result in a population of one species breeding with a population of another species as a means of surviving such as the case with the [[Red Wolf|red wolves]]. Their population numbers being so small, they needed another means of survival. Habitat fragmentation also led to the influx of generalist species into areas where they would not have been, leading to competition and in some cases interbreeding/incorporation of a population into another. In this way, habitat fragmentation is essentially an indirect method of introducing species to an area.
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| ===The hybridization continuum===
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| There is a kind of continuum with three semi-distinct categories dealing with anthropogenic hybridization: hybridization without [[Introgression]], hybridization with widespread introgression, and essentially a [[Hybrid swarm]].<ref name="Allendorf 2001 613–622"/> 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 fields. [[Global warming|Global climate change]] is creating other changes such as difference in population distributions which are indirect causes for an increase in anthropogenic hybridization.
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| ===Consequences===
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| Hybridization can be a less discussed way toward [[extinction]] than within detection of where a population lies along the hybrid continuum. The dispute of hybridization is how to manage the resulting hybrids. When a population experiences hybridization with substantial introgression, there still exists parent types of each set of individuals. When a complete hybrid swarm is created, all the individuals are hybrids.{{citation needed|date=February 2013}}
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| ===Management of hybrids===
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| Conservationists disagree on when 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 it becomes a complete mixture, we should look to conserve those hybrids to avoid their loss.<ref name="Allendorf 2001 613–622"/> Most leave it as a case-by-case basis, depending on detecting of hybrids within the group. It is nearly impossible to regulate hybridization via policy because hybridization can occur beneficially when it occurs "naturally" and there is the matter of protecting those previously mentioned hybrid swarms because if they are the only remaining evidence of prior species, they need to be conserved as well.<ref name="Allendorf 2001 613–622"/>
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| {{Unreferenced section|date=August 2009}}
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| In some species, hybridisation plays an important role in evolutionary biology. While most hybrids are [[disadvantaged]] as a result of genetic incompatibility, the fittest survive, regardless of species boundaries. They may have a beneficial combination of traits allowing them to exploit new habitats or to succeed in a marginal habitat where the two parent species are disadvantaged. This has been seen in experiments on sunflower species. Unlike mutation, which affects only one gene, hybridisation creates multiple variations across genes or gene combinations simultaneously. Successful hybrids could evolve into new species within 50-60 generations. This leads some scientists to speculate that life is a genetic continuum rather than a series of self-contained species.
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| Where there are two closely related species living in the same area, less than 1 in 1000 individuals are likely to be hybrids because animals rarely choose a mate from a different species (otherwise species boundaries would completely break down). In some closely related species there are recognized "hybrid zones".
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| Some species of Heliconius butterflies exhibit dramatic geographical polymorphism of their wing patterns, which act as aposematic signals advertising their unpalatability to potential predators. Where different-looking geographical races abut, inter-racial hybrids are common, healthy and fertile. Heliconius hybrids can breed with other hybrid individuals and with individuals of either parental race. These hybrid backcrosses are disadvantaged by natural selection because they lack the parental form's warning coloration, and are therefore not avoided by predators.
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| A similar case in mammals is hybrid White-Tail/Mule Deer. The hybrids don't inherit either parent's escape strategy. White-tail Deer dash while Mule Deer bound. The hybrids are easier prey than the parent species.
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| In birds, healthy Galapagos Finch hybrids are relatively common, but their beaks are intermediate in shape and less efficient feeding tools than the specialised beaks of the parental species so they lose out in the competition for food. Following a major storm in 1983, the local habitat changed so that new types of plants began to flourish, and in this changed habitat, the hybrids had an advantage over the birds with specialised beaks - demonstrating the role of hybridization in exploiting new ecological niches. If the change in environmental conditions is permanent or is radical enough that the parental species cannot survive, the hybrids become the dominant form. Otherwise, the parental species will re-establish themselves when the environmental change is reversed, and hybrids will remain in the minority.
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| Natural hybrids may occur when a species is introduced into a new habitat. In Britain, there is hybridisation of native European Red Deer and introduced Chinese Sika Deer. Conservationists want to protect the Red Deer, but the environment favors the Sika Deer genes. There is a similar situation with White-headed Ducks and Ruddy Ducks.
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| ===Expression of parental traits in hybrids===
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| When two distinct types of organisms breed with each other, the resulting hybrids typically have intermediate traits (e.g., one parent has red flowers, the other has white, and the hybrid, pink flowers).<ref name="McCarthy, Eugene M 2006. Pp. 16-17">McCarthy, Eugene M. 2006. ''Handbook of Avian Hybrids of the World''. Oxford: Oxford University Press. Pp. 16-17.</ref> Commonly, hybrids also combine traits seen only separately in one parent or the other (e.g., a [[bird hybrid]] might combine the yellow head of one parent with the orange belly of the other).<ref name="McCarthy, Eugene M 2006. Pp. 16-17"/> Most characteristics of the typical hybrid are of one of these two types, and so, in a strict sense, are not really new. However, an intermediate trait does differ from those seen in the parents (e.g., the pink flowers of the intermediate hybrid just mentioned are not seen in either of its parents). Likewise, combined traits are new when viewed as a combination.
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| In a hybrid, any trait that falls outside the range of parental variation is termed [[Heterosis|heterotic]]. Heterotic hybrids do have new traits, that is, they are not intermediate. ''Positive heterosis'' produces more robust hybrids, they might be stronger or bigger; while the term ''negative heterosis'' refers to weaker or smaller hybrids.<ref>McCarthy, Eugene M. 2006. ''Handbook of Avian Hybrids of the World''. Oxford: Oxford University Press. P. 17.</ref> 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 a tigon (lioness × tiger) is smaller. Also 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 the Common Pheasant and hen Golden Pheasant (''[[Chrysolophus pictus]]'').<ref>Darwin, C. 1868. ''Variation of Animals and Plants under Domestication,'' vol. II, p. 125</ref> Spurs are absent in hybrids of the former type, although present in both parents.<ref>Spicer, J. W. G. 1854. Note on hybrid gallinaceous birds. ''The Zoologist,'' 12: 4294-4296 (see p. 4295).</ref>
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| When populations hybridize, often the first generation (F<sub>1</sub>) hybrids are very uniform. Typically, however, the individual members of subsequent hybrid generations are quite variable. High levels of variability in a natural population, then, are indicative of hybridity. Researchers use this fact to ascertain whether a population is of hybrid origin. Since such variability generally occurs only in later hybrid generations, the existence of variable hybrids is also an indication that the hybrids in question are fertile.{{Citation needed|date=August 2009}}
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| ===Genetic mixing and extinction===
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| {{Main|Genetic pollution}}
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| Regionally developed [[ecotype]]s can be threatened with [[extinction]] when new [[allele]]s or genes are introduced that alter that ecotype. This is sometimes called [[genetic mixing]].<ref>{{cite journal | last1 = Mooney | first1 = H. A. | last2 = Cleland | first2 = E. E. | year = 2001 | title = The evolutionary impact of invasive species | url = http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=33232 | journal = Proc Natl Acad Sci U S A. | volume = 98 | issue = 10| pages = 5446–5451 | doi = 10.1073/pnas.091093398 | pmid = 11344292 | pmc = 33232 }}</ref> Hybridization and [[introgression]] of new genetic material can lead to the replacement of local [[genotypes]] if the hybrids are more [[Fitness (biology)|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 a degree that none of the originally genetically distinct population remains.<ref>Rhymer JM and Simberloff, D. (1996) [http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.ecolsys.27.1.83 Extinction by Hybridization and Introgression]. ''Annual Review of Ecology and Systematics'' '''27''': 83-109 {{doi|10.1146/annurev.ecolsys.27.1.83}}, [http://links.jstor.org/sici?sici=0066-4162(1996)27%3C83:EBHAI%3E2.0.CO;2-A#abstract]</ref><ref>Brad M. Potts, Robert C. Barbour, Andrew B. Hingston (2001) [http://www.rirdc.gov.au/reports/AFT/01-114.pdf Genetic Pollution from Farm Forestry using eucalypt species and hybrids]; A report for the RIRDC/L&WA/FWPRDC; Joint Venture Agroforestry Program; RIRDC Publication No 01/114; RIRDC Project No CPF - 3A; ISBN 0-642-58336-6; ISSN 1440-6845; Australian Government, Rural Industrial Research and Development Corporation</ref>
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| ===Effect on biodiversity and food security===
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| {{Main|biodiversity|food security}}
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| 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 the loss of [[genetic diversity]].<ref>Devinder Sharma [http://www.farmedia.org/bulletins/bulletin28.html “Genetic Pollution: The Great Genetic Scandal”]; Bulletin 28. hosted by www.farmedia.org</ref> 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.<ref>Troyer, A. Forrest. ''Breeding Widely Adapted Cultivars: Examples from Maize.'' Encyclopedia of Plant and Crop Science, 27 February 2004.</ref>
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| ==Limiting factors==
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| A number of conditions exist that limit the success of hybridization, the most obvious is great genetic diversity between most species. But in animals and plants that are more closely related hybridization barriers can include morphological differences, differing times of fertility, mating behaviors and cues, physiological rejection of sperm cells or the developing embryo.{{Citation needed|date=August 2009}}
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| In plants, 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 the chromosomes.<ref>Barriers to hybridization of Solanum bulbocastanumDun. and S. VerrucosumSchlechtd. and structural hybridity in their F1 plants
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| Journal Euphytica [googilygoogilygoo..bwahahaa]
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| Publisher Springer Netherlands
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| ISSN 0014-2336 (Print) 1573-5060 (Online)
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| Issue Volume 25, Number 1 / January, 1976
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| Category Articles
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| DOI 10.1007/BF00041523
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| Pages 1-10</ref>
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| ==Mythical, legendary and religious hybrids==
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| {{Unreferenced section|date= October 2010}}
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| {{main|Mythological hybrid}}
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| Ancient folktales often contain mythological creatures, sometimes these are described as hybrids (e.g., [[Hippogriff]] as the offspring of a [[griffin]] and a [[horse]], and the [[Minotaur]] which is the offspring of [[Pasiphaë]] and a white bull). More often they are kind of [[Chimera (mythology)|chimera]], i.e., a composite of the physical attributes of two or more kinds of animals, mythical beasts, and often humans, with no suggestion that they are the result of interbreeding, e.g., [[Harpy|Harpies]], [[mermaid]]s, and [[centaur]]s.
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| In the [[Bible]], the [[Old Testament]] contains several passages which talk about a first generation of hybrid giants who were known as the [[Nephilim]].<ref>[[James L. Kugel]] (2009), "[http://books.google.es/books?id=y8AjDrIkBG4C&printsec=frontcover&hl=es#v=onepage&q&f=false Traditions of the Bible: A Guide to the Bible As It Was at the Start of the Common Era]", [[Harvard University Press]], p. 198</ref> The [[Book of Genesis]] (6:4) states that "the sons of God went to the daughters of humans and had children by them". As a result, the offspring was born as hybrid [[giant (mythology)|giant]]s who became mighty heroes of old and legendary famous figures of ancient times.<ref>[[James L. Kugel]] (1997), [http://books.google.es/books?id=i8ppE9olz_AC&printsec=frontcover&hl=es#v=onepage&q&f=false The Bible as it was], [[Harvard University Press]], p. 110</ref> Several verses later the writer states that God saw that "''all flesh had corrupted his way upon the earth''." In addition, the [[Book of Numbers]] (13:33) says that the descendants of [[Anak]] came from the Nephilim, whose bodies looked exactly like men, but with an enormous height. According to the "apocryphal" [[Book of Enoch]] and some theological interpretations, the hybrid Nephilim were wicked sons of "[[fallen angel]]s" who had lusted with attractive women.<ref>Gregory A. Boyd, God at War: The Bible & Spiritual Conflict, p. 177</ref>
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| ==See also==
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| {{Portal|Biology}}
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| <div style="column-count:3;-moz-column-count:3;-webkit-column-count:3">
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| * [[Artificial selection]]
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| * [[Bird hybrid]]s
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| * [[Cabbit]]
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| * [[Canid hybrid]]
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| * [[Chimera (genetics)]]
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| * [[Chloroplast capture]] (botany)
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| * [[Felid hybrid]]s
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| * [[F1 hybrid]]s
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| * [[Genetic admixture]]
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| * [[Genetic erosion]]
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| * [[Grex (horticulture)]]
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| * [[Heterosis]] (hybrid vigor)
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| * [[Human-animal hybrid]] (parahuman)
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| * [[Hybrid Lovebird]]
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| * [[Mythological hybrid|Hybrid (mythology)]]
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| * [[Hybrid name]]s (botany)
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| * [[Hybrid speciation]]
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| * [[Hybrid swarm]]
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| * [[Hybrid zone]]
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| * [[Hybrot]]
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| * [[Inbreeding]]
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| * [[Interspecific pregnancy]]
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| * [[Intraspecific breeding]]
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| * [[Macropod hybrids]]
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| * [[Purebred]]
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| * [[Selective breeding]]
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| * [[Sheep-goat hybrid]]s
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| * [[Species barrier]]
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| * [[Synergy]]
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| </div>
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| ==References==<!-- CurrBiol16:R245. SystBiol55:46[hybrids and cladistic analyses]. -->
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| {{Reflist|2}}
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| {{commons category|Hybridization}}
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| ==External links==
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| * [http://www.orchids.co.in/plant-facts/artificial-hybridisation.shtm Artificial Hybridisation] - Artificial Hybridisation in orchids
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| * [http://www.feathersite.com/Poultry/BRKHybrids.html Domestic Fowl Hybrids]
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| * [http://www.messybeast.com/genetics/hybrid-mammals.html Hybrid Mammals]
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| * [http://news.nationalgeographic.com/news/2005/07/0727_050727_evolution.html Hybridisation in animals] Evolution Revolution: Two Species Become One, Study Says (nationalgeographic.com)
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| * [http://www.messybeast.com/small-hybrids/hybrids.htm Hybrids of wildcats with domestic cats]
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| * [http://www.livescience.com/animalworld/060614_butterfly_hybrid.html Scientists Create Butterfly Hybrid] - Creation of new species through hybridization was thought to be common only in plants, and rare in animals.
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| * [http://www.cambridgenetwork.co.uk/views/biolines What is a human admixed embryo?]
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| * [http://www.carp-fishingholidayfrance.co.uk/videos.htm Video of Mirror carp & Gold fish spawning at a fishing venue in France]
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| {{evolution}}
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| {{Population genetics}}
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| {{Mammal hybrids}}
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| {{Lysenkoism}}
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| {{Use dmy dates|date=March 2011}}
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| {{DEFAULTSORT:Hybrid (Biology)}}
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| [[Category:Hybridisation (biology)| ]]
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| [[Category:Biology terminology]]
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| [[Category:Botanical nomenclature]]
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| [[Category:Evolutionary biology]]
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| [[Category:Population genetics]]
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| [[Category:Breeding]]
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