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d species actually do exist in nature in both space and time implies that some fundamental feature of natural selection operates to generate and maintain species 11 a possible explanation for how these dilemmas can be resolved is discussed in the article speciation in the section effect of sexual reproduction on species formation biogeographic influence edit the german traveller moritz wagner 1813 1887 see also geographic speciation recognition of geographic factors involved in species populations was present even before darwin with many naturalists aware of the role of isolation in species relationships 14 482 in 1833 c l gloger published the variation of birds under the influence of climate in which he described geographic variations but did not recognize that geographic isolation was an indicator of past speciation events 14 482 another naturalist in 1856 wollaston studied island beetles in comparison to mainland species 14 482 he saw isolation as key to their differentiation 14 482 however he did not recognize that the pattern was due to speciation 14 483 one naturalist leopold von buch 1825 did recognize the geographic patterns and explicitly stated that geographic isolation may lead to species separating into new species 14 483 mayr suggests that von buch was likely the first naturalist to truly suggest geographic speciation 15 other naturalists such as henry walter bates 1863 recognized and accepted the patterns as evidence of speciation but in bate s case did not propose a coherent model 14 484 in 1868 moritz wagner was the first to propose the concept of geographic speciation 16 14 484 in which he used the term separationstheorie 5 edward bagnall poulton the evolutionary biologist and a strong proponent of the importance of natural selection highlighted the role of geographic isolation in promoting speciation 17 in the process coining the term sympatric speciation in 1904 18 19 wagner and other naturalists who studied the geographic distributions of animals such as karl jordan and david starr jordan noticed that closely related species were often geographically isolated from one another allopatrically distributed which lead to the advocation of the importance of geographic isolation in the origin of species 4 2 karl jordan is thought to have recognized the unification of mutation and isolation in the origin of new species in stark contrast to the prevailing views at the time 14 486 david starr jordan reiterated wagner s proposal in 1905 providing a wealth of evidence from nature to support the theory 16 20 4 2 and asserting that geographic isolation is obvious but had been unfortunately ignored by most geneticists and experimental evolutionary biologists at the time 14 487 joel asaph allen suggested the observed pattern of geographic separation of closely related species be called jordan s law or wagner s law 14 487 despite the contentions most taxonomists did accept the geographic model of speciation 14 487 many of the early terms used to describe speciation were outlined by ernst mayr 21 he was the first to encapsulate the then contemporary literature in his 1942 publication systematics and the origin of species from the viewpoint of a zoologist and in his subsequent 1963 publication animal species and evolution like jordan s works they relied on direct observations of nature documenting the occurrence of geographic speciation 4 86 he described the three modes geographic semi geographic and non geographic which today are referred to as allopatric parapatric and sympatric respectively 21 mayr s 1942 publication influenced heavily by the ideas of karl jordan and poulton was regarded as the authoritative review of speciation for over 20 years and is still valuable today 19 a major focus of mayr s works was on the importance of geography in facilitating speciation with islands often acting as a central theme to many of the speciation concepts put forth 22 one of which was the concept of peripatric speciation a variant of allopatric speciation 23 24 he has since distinguished the two modes by referring to them as peripatric and dichopatric 25 this concept arose by an interpretation of wagner s separationstheorie as a form of founder effect speciation that focused on small geographically isolated species 5 this model was later expanded and modified to incorporate sexual selection by kenneth y kaneshiro in 1976 and 1980 26 27 28 modern evolutionary synthesis edit main article modern synthesis 20th century many geneticists at the time did little to bridge the gap between the genetics of natural selection and the origin of reproductive barriers between species 4 3 ronald fisher proposed a model of speciation in his 1930 publication the genetical theory of natural selection where he described disruptive selection acting on sympatric or parapatric populations with reproductive isolation completed by reinforcement 29 other geneticists such as j b s haldane did not even recognize that species were real while sewall wright ignored the topic despite accepting allopatric speciation 4 3 the primary contributors to the incorporation of speciation into modern evolutionary synthesis were ernst mayr and theodosius dobzhansky 29 dobzhansky a geneticist published genetics and the origin of species in 1937 in which he formulated the genetic framework for how speciation could occur 4 2 he recognized that speciation was an unsolved problem in biology at the time rejecting darwin s position that new species arose by occupation of new niches contending that reproductive isolation was instead based on barriers to gene flow 4 2 subsequently mayr conducted extensive work on the geography of species emphasizing the importance of geographic separation and isolation in which he filled dobzhansky s gaps concerning the origin of biodiversity in his 1942 book 30 both of their works gave rise not without controversy to the modern understanding of speciation stimulating a wealth of research on the topic 4 3 furthermore this extended to plants as well as animals with g ledyard stebbins s book variation and evolution in plants and the much later 1981 book plant speciation by verne grant ernst mayr an influential evolutionary biologist 31 in 1947 a consensus had been achieved among geneticists paleontologists and systematists and that evolutionary biology as an independent biological discipline had been established during a princeton university conference 32 this 20th century synthesis incorporated speciation since then the ideas have been consistently and repeatedly confirmed 30 contemporary work edit after the synthesis speciation research continued largely within natural history and biogeography with much less emphasis on genetics 4 4 the study of speciation has seen its largest increase since the 1980s 4 4 with an influx of publications and a host of new terms methods concepts and theories 21 this third phase of work as jerry a coyne and h allen orr put it has led to a growing complexity of the language used to describe the many processes of speciation 21 the research and literature on speciation have become enormous scattered and increasingly technical 4 1 from the 1980s new research tools increased the robustness of research 4 4 assisted by new methods theoretical frameworks models and approaches 21 coyne and orr discuss the modern post 1980s developments centered around five major themes genetics also a primary factor in the modern synthesis molecular biology and analysis namely phylogenetics and systematics comparative analysis mathematical modeling and computer simulations and the role of ecology 4 5 ecologists became aware that the ecological factors behind speciation were under represented this saw the growth in research concerning ecology s role in facilitating speciation rightly designated ecological speciation 4 4 this focus on ecology generated a host of new terms relating to the barriers to reproduction 21 e g allochronic speciation in which gene flow is reduced or removed by timing of breeding periods or habitat isolation in which species occupy different habitats within the same area sympatric speciation regarded by mayr as unlikely has become widely accepted 33 34 35 research on the influence of natural selection on speciation including the process of reinforcement has grown 36 researchers have long debated the roles of sexual selection natural selection and genetic drift in speciation 4 383 darwin extensively discussed sexual selection with his work greatly expanded on by ronald fisher however it was not until 1983 that the biologist mary jane west eberhard recognized the importance of sexual selection in speciation 37 4 3 natural selection plays a role in that any selection towards reproductive isolation can result in speciation whether indirectly or directly genetic drift has been widely researched from the 1950s onwards especially with peak shift models of speciation by genetic drift 4 388 mayr championed founder effects in which isolated individuals like those found on islands near a mainland experience a strong population bottleneck as they contain only a small sample of the genetic variation in the main population 4 390 38 later other biologists such as hampton l carson alan templeton sergey gavrilets and alan hastings developed related models of speciation by genetic drift noting that islands were inhabited mostly by endemic species 39 selection s role in speciation is widely supported whereas founder effect speciation is not 4 410 having been subject to a number of criticisms 40 classification debate edit speciation represented as a continuum of gene flow where m displaystyle m equals the rate of gene exchange the three primary geographic modes of speciation allopatric parapatric and sympatric can exist within this continuum as well as other non geographic modes throughout the history of research concerning speciation classification and delineation of modes and processes have been debated julian huxley divided speciation into three separate modes geographical speciation genetic speciation and ecological speciation 14 427 sewall wright proposed ten different varying modes 14 427 ernst mayr championed the importance of physical geographic separation of species populations maintaining it to be of major importance to speciation he originally proposed the three primary modes known today geographic semi geographic non geographic 21 corresponding to allopatric parapatric and sympatric respectively the phrase modes of speciation is imprecisely defined most often indicating speciation occurring as a result of a species geographic distribution 41 more succinctly the modern classification of speciation is often described as occurring on a gene flow continuum i e allopatry at m 0 displaystyle m 0 and sympatry at m 0 5 displaystyle m 0 5 42 43 this gene flow concept views speciation as based on the exchange of genes between populations instead of seeing a purely geographic setting as necessarily relevant despite this concepts of biogeographic modes can be translated into models of gene flow such as that in the image at left however this translation has led to some confusion of language in the scientific literature 21 comparisons of the three classic geographic modes of speciation allopatric parapatric and sympatric with peripatric speciation included as a special case of allopatric speciation as research has expanded over the decades the geographic scheme has been challenged the traditional classification is considered by some researchers to be obsolete 44 while others argue for its merits proponents of non geographic schemes often justify non geographic classifications not by rejection of the importance of reproductive isolation or even the processes themselves but instead by the fact that it simplifies the complexity of speciation 45 one major critique of the geographic framework is that it arbitrarily separates a biological continuum into discontinuous groups 45 another criticism rests with the fact that when speciation is viewed as a continuum of gene flow parapatric speciation becomes unreasonably represented by the entire continuum 46 with allopatric and sympatric existing in the extremes 45 coyne and orr argue that the geographic classification scheme is valuable in that biogeography controls the strength of the evolutionary forces at play as gene flow and geography are clearly linked 44 james mallet and colleagues contend that the sympatric vs allopatric dichotomy is valuable to determine the degree in which natural selection acts on speciation 47 kirkpatrick and ravigné categorize speciation in terms of its genetic basis or by the forces driving reproductive isolation 4 85 here the geographic modes of speciation are classified as types of assortive mating 48 fitzpatrick and colleagues believe that the biogeographic scheme is a distraction that could be positively misleading if the real goal is to understand the influence of natural selection on divergence 44 they maintain that to fully understand speciation the spatial ecological and genetic factors involved in divergence must be explored 44 sara via recognizes the importance of geography in speciation but suggests that classification under this scheme be abandoned 34 history of modes and mechanisms edit sympatric speciation edit sympatric speciation from its beginnings with darwin who did not coin the term has been a contentious issue 41 4 125 mayr along with many other evolutionary biologists interpreted darwins s view of speciation and the origin of biodiversity as arising by species entering new ecological niches a form of sympatric speciation 1 before mayr sympatric speciation was regarded as the primary mode of speciation in 1963 mayr provided a strong criticism citing various flaws in the theory 4 126 after that sympatric speciation fell out of favor with biologists and has only recently seen a resurgence in interest 4 126 some biologists such as james mallet believe that darwin s view on speciation was misunderstood and misconstrued by mayr 1 49 today sympatric speciation is supported by evidence from laboratory experiments and observations from nature 4 127 33 hybrid speciation edit for most of the history of speciation hybridization polyploidy has been a contentious issue as botanists and zoologists have traditionally viewed hybridization s role in speciation differently 21 carl linnaeus was the earliest to suggest hybridization in 1760 50 øjvind winge was the first to confirm allopolyploidy in 1917 50 51 and a later experiment conducted by clausen and goodspeed in 1925 confirmed the findings 50 today it is widely recognized as a common mechanism of speciation 52 historically zoologists considered hybridization to be a rare phenomenon while botanists found it to be commonplace in plant species 21 the botanists g ledyard stebbins and verne grant were two of the well known botanists who championed the idea of hybrid speciation during the 1950s to the 1980s 21 ...