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Allopatric speciation
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== Laboratory evidence == {{see also|Laboratory experiments of speciation}} [[File:Drosophila speciation experiment.svg|right|thumb|upright=1.8|A simplification of an experiment where two vicariant lines of [[Drosophila melanogaster|fruit flies]] were raised on harsh [[maltose]] and [[starch]] mediums respectively. The experiment was replicated with 8 populations; 4 with maltose and 4 with starch. Differences in adaptations were found for each population corresponding to the different mediums.<ref>{{Citation |title=Population genetics of ''Drosophila'' amylase. IV. Selection in laboratory populations maintained on different carbohydrates | author=J. R. Powell | author2= M. Andjelkovic | journal=Genetics | year=1983 | volume=103 | issue= 4| pages=675β689 | doi=10.1093/genetics/103.4.675 | pmid=6189764 | pmc=1202048 }}</ref> Later investigation found that the populations evolved behavioral isolation as a [[Pleiotropy|pleiotropic]] by-product from this adaptive divergence.<ref name=Dodd1989/> This form of pre-zygotic isolation is a prerequisite for speciation to occur.]] Experiments on allopatric speciation are often complex and do not simply divide a species population into two. This is due to a host of defining parameters: measuring reproductive isolation, sample sizes (the number of matings conducted in reproductive isolation tests), bottlenecks, length of experiments, number of generations allowed,<ref name=ABF&AO2002>{{Citation |title=Laboratory environments are not conducive for allopatric speciation | author=Ann-Britt Florin | author2=Anders Γdeen | journal=Journal of Evolutionary Biology | year=2002 | volume=15 | pages=10β19 | doi=10.1046/j.1420-9101.2002.00356.x| s2cid=85410953 | doi-access=free }}</ref> or insufficient genetic diversity.<ref name=MK&VR2002>{{Citation |title=Speciation by Natural and Sexual Selection: Models and Experiments | author=Mark Kirkpatrick | author2=Virginie RavignΓ© | journal=The American Naturalist | year=2002 | volume=159 | issue=3 | pages=S22 | doi=10.2307/3078919| jstor=3078919 }}</ref> Various isolation indices have been developed to measure reproductive isolation (and are often employed in laboratory speciation studies) such as here (index <math>Y</math><ref>{{Citation | title=Discrete Multivariate Analysis: Theory and Practice | author=Bishop, Y. M.|author1-link= Yvonne Bishop |author2=Fienberg, S. E.| author3=Holland, P. W. | date=1975 | publisher=MIT Press: Cambridge, MA. }}</ref> and index <math>I</math><ref>{{Citation |title=Sexual isolation studies in the species complex ''Drosophila virilis'' | author=H. D. Stalker | journal=Genetics | year=1942 | volume=27 | issue= 2| pages=238β257 | doi=10.1093/genetics/27.2.238 | pmid=17247038 | pmc=1209156 }}</ref>): <math>Y= {\sqrt{(AD/BC)}-1 \over \sqrt{(AD/BC)+1}}</math> <math>I= {A+D-B-C \over A+D+B+C} </math> Here, <math>A</math> and <math>D</math> represent the number of [[Heterogametic sex|matings in heterogameticity]] where <math>B</math> and <math>C</math> represent [[Heterogametic sex|homogametic matings]]. <math>A</math> and <math>B</math> is one population and <math>D</math> and <math>C</math> is the second population. A negative value of <math>Y</math> denotes negative assortive mating, a positive value denotes positive assortive mating (i. e. expressing reproductive isolation), and a [[Null (mathematics)|null value]] (of zero) means the populations are experiencing [[Panmixia|random mating]].<ref name=ABF&AO2002/> The experimental evidence has solidly established the fact that reproductive isolation evolves as a by-product of selection.<ref name="Rice&Hostert"/><ref name="Speciation"/>{{rp|90}} Reproductive isolation has been shown to arise from [[pleiotropy]] (''i.e.'' indirect selection acting on genes that code for more than one trait)βwhat has been referred to as [[genetic hitchhiking]].<ref name="Rice&Hostert" /> Limitations and controversies exist relating to whether laboratory experiments can accurately reflect the long-scale process of allopatric speciation that occurs in nature. Experiments often fall beneath 100 generations, far less than expected, as [[Speciation in the fossil record|rates of speciation]] in nature are thought to be much larger.<ref name="Speciation"/>{{rp|87}} Furthermore, rates specifically concerning the evolution of reproductive isolation in ''Drosophila'' are significantly higher than what is practiced in laboratory settings.<ref>{{Citation |title="Patterns of Speciation in Drosophila" Revisited | author=Jerry A. Coyne |author1-link=Jerry Coyne | author2=H. Allen Orr |author2-link=H. Allen Orr | journal=Evolution | year=1997 | volume=51 | issue=1 | pages=295β303 | doi= 10.1111/j.1558-5646.1997.tb02412.x| pmid=28568795 | s2cid=40390753 | doi-access=free }}</ref> Using index ''Y'' presented previously, a survey of 25 allopatric speciation experiments (included in the table below) found that reproductive isolation was not as strong as typically maintained and that laboratory environments have not been well-suited for modeling allopatric speciation.<ref name=ABF&AO2002/> Nevertheless, numerous experiments have shown pre-zygotic and post-zygotic isolation in vicariance, some in less than 100 generations.<ref name="Speciation"/>{{rp|87}} Below is a non-exhaustive table of the laboratory experiments conducted on allopatric speciation. The first column indicates the species used in the referenced study, where the "Trait" column refers to the specific characteristic selected for or against in that species. The "Generations" column refers to the number of generations in each experiment performed. If more than one experiment was formed generations are separated by semicolons or dashes (given as a range). Some studies provide a duration in which the experiment was conducted. The "Selection type" column indicates if the study modeled vicariant or peripatric speciation (this may not be explicitly). [[Directional selection|Direct selection]] refers to selection imposed to promote reproductive isolation whereas indirect selection implies isolation occurring as a pleiotropic byproduct of natural selection; whereas divergent selection implies deliberate selection of each allopatric population in opposite directions (''e.g.'' one line with more bristles and the other line with less). Some studies performed experiments modeling or controlling for genetic drift. Reproductive isolation occurred pre-zygotically, post-zygotically, both, or not at all. It is important to note that many of the studies conducted contain multiple experiments withinβa resolution of which this table does not reflect. {| class="wikitable sortable" style="margin-left: auto; margin-right: auto; border: none;" |+Laboratory studies of allopatric speciation<ref name="Speciation"/>{{rp|88β89}}<ref name="Rice&Hostert" /><ref name=MK&VR2002/><ref name=ABF&AO2002/> !Species !Trait !~Generations (duration) !Selection type !Studied Drift !Reproductive isolation !Year & Reference |- | rowspan="17" |''[[Drosophila]]<br>[[Drosophila melanogaster|melanogaster]]'' |[[Escape response]] |18 |Indirect; divergent |Yes |Pre-zygotic |1969<ref>{{Citation |title=Disruptive and stabilizing selection on the" escape" behavior of ''Drosophila melanogaster'' | author=B. S. Grant | author2=L. E. Mettler | journal=Genetics | year=1969 | volume=62 | issue=3 | pages=625β637 | doi=10.1093/genetics/62.3.625 | pmid=17248452 | pmc=1212303 }}</ref> |- |[[Animal locomotion|Locomotion]] |112 |Indirect; divergent |No |Pre-zygotic |1974<ref>{{Citation |title=Activity and sexual behaviour in ''Drosophila melanogaster'' | author=B. Burnet | author2=K. Connolly | journal=The Genetics of Behaviour | year=1974 | pages=201β258 }}</ref> |- |Temperature, [[humidity]] |70β130 |Indirect; divergent |Yes |Pre-zygotic |1980<ref>{{Citation |title=A Multifactorial Genetic Investigation of Speciation Theory Using ''Drosophila melanogaster'' | author=G. Kilias|author2= S. N. Alahiotis|author3=M. Pelecanos | journal=Evolution | year=1980 | volume=34 | issue=4 | pages=730β737 | doi=10.2307/2408027 | pmid=28563991| jstor=2408027}}</ref> |- |[[Dichlorodiphenyltrichloroethane|DDT]] adaptation |600 (25 years, +15 years) |Direct |No |Pre-zygotic |2003<ref>{{Citation |title=Forty years of solitude: life-history divergence and behavioural isolation between laboratory lines of ''Drosophila melanogaster'' | author=C. R. B. Boake|author2= K. Mcdonald|author3=S. Maitra|author4=R. Ganguly | journal=Journal of Evolutionary Biology | year=2003 | volume=16 | issue=1 | pages=83β90 | doi=10.1046/j.1420-9101.2003.00505.x | pmid=14635883| s2cid=24040182| doi-access=free }}</ref> |- | |17, 9, 9, 1, 1, 7, 7, 7, 7 |Direct, divergent | |Pre-zygotic |1974<ref>{{Citation |title=Effects of population size and selection intensity on responses to disruptive selection in ''Drosophila melanogaster'' | author=J. S. F. Barker | author2=L. J. E. Karlsson | journal=Genetics | year=1974 | volume=78 | issue=2 | pages=715β735 | doi=10.2307/2407287 | jstor=2407287 | pmid=4217303 | pmc=1213230 }}</ref> |- | |40; 50 |Direct; divergent | |Pre-zygotic |1974<ref>{{Citation |title=Changes in Mating Behavior Produced by Selection for Ethological Isolation Between Ebony and Vestigial Mutants of ''Drosophila melanogaster'' | author=Stella A. Crossley | journal=Evolution | year=1974 | volume=28 | issue=4 | pages=631β647 | doi= 10.1111/j.1558-5646.1974.tb00795.x| pmid=28564833 | s2cid=35867118 }}</ref> |- |Locomotion |45 |Direct; divergent |No |None |1979<ref>{{Citation |title=Divergent selection on locomotor activity in ''Drosophila melanogaster''. I. Selection response | author=F. R. van Dijken | author2=W. Scharloo | journal=Behavior Genetics | year=1979 | volume=9 | issue=6 | pages=543β553 | doi=10.1007/BF01067350 | pmid=122270 | s2cid=39352792 }}</ref><ref>{{Citation|title=Divergent selection on locomotor activity in ''Drosophila melanogaster''. II. Test for reproductive isolation between selected lines|year=1979|author=F. R. van Dijken | author2=W. Scharloo|journal=Behavior Genetics|volume=9|issue=6|pages=555β561|doi=10.1007/BF01067351|pmid=122271|s2cid=40169222}}</ref> |- | | |Direct; divergent | |Pre-zygotic |1953<ref>{{Citation |title=Genetic divergence of isolated populations of ''Drosophila melanogaster'' | author=B. Wallace | journal=Proceedings of the Ninth International Congress of Genetics | year=1953 | volume=9 | pages=761β764}}</ref> |- | |36; 31 |Direct; divergent | |Pre-zygotic |1956<ref>{{Citation |title=Selection for sexual isolation within a species | author=G. R. Knight | display-authors=etal | journal=Evolution | year=1956 | volume=10 | pages=14β22 | doi=10.1111/j.1558-5646.1956.tb02825.x| s2cid=87729275 | doi-access=free }}</ref> |- |[[Ethylenediaminetetraacetic acid|EDTA]] adaptation |3 experiments, 25 each |Indirect |No |Post-zygotic |1966<ref>{{Citation |title=A test of sexual isolation in ''Drosophila'' | author=Forbes W. Robertson | journal=Genetical Research | year=1966 | volume=8 | issue=2 | pages=181β187 | doi=10.1017/s001667230001003x | pmid=5922518| doi-access=free }}</ref><ref>{{Citation|title=The ecological genetics of growth in ''Drosophila'' 8. Adaptation to a New Diet|year=1966|author=Forbes W. Robertson|journal=Genetical Research|volume=8|issue=2|pages=165β179|doi=10.1017/s0016672300010028|pmid=5922517|doi-access=free}}</ref> |- | |8 experiments, 25 each |Direct | | |1997<ref>{{Citation |title=Reinforcement: a new perspective on an old controversy | author=Ellen E. Hostert | journal=Evolution | year=1997 | volume=51 | issue=3 | pages=697β702 | doi=10.1111/j.1558-5646.1997.tb03653.x| pmid=28568598 | s2cid=21054233 | doi-access=free }}</ref> |- |Abdominal [[chaeta]] number |21β31 |Direct |Yes |None |1958<ref>{{Citation |title= The origin of sexual isolation between different lines within a species | author=Koref SantibaΓ±ez, S. | author2=Waddington, C. H. | journal=Evolution | year=1958 | volume=12 | issue= 4| pages=485β493 | doi= 10.2307/2405959| jstor=2405959 }}</ref> |- |[[Morphology of Diptera|Sternopleural]] chaeta number |32 |Direct |No |None |1969<ref>{{Citation|title=The effect of selection for sternopleural bristle number in mating behaviour in ''Drosophila melanogaster''|year=1969|author=Barker, J. S. F. | author2=Cummins, L. J.|journal=Genetics|volume=61|issue=3|pages=713β719|doi=10.1093/genetics/61.3.713|pmid=17248436|pmc=1212235}}</ref> |- |[[Phototaxis]], [[geotaxis]] |20 | |No |None |1975<ref>{{Citation |title=A genetic analysis of phototactic behavior in ''Drosophila melanogaster'' | author=Markow, T. A. | journal=Genetics | year=1975 | volume=79 | issue=3 | pages=527β534 | doi=10.1093/genetics/79.3.527 | pmid=805084 | pmc=1213291 | doi-access=free }}</ref> 1981<ref>{{Citation|title=Mating preferences are not predictive of the direction of evolution in experimental populations of ''Drosophila''|year=1981|author=Markow, T. A.|s2cid=15497733|journal=Science|volume=213|issue=4514|pages=1405β1407|doi=10.1126/science.213.4514.1405|pmid=17732575|bibcode=1981Sci...213.1405M}}</ref> |- | | | |Yes | |1998<ref>{{Citation |title=Single founder-flush events and the evolution of reproductive isolation | author=Rundle, H. D.| author2=Mooers, A. Γ. | author3=Whitlock, M. C. | journal=Evolution | year=1998 | volume=52 | issue= 6| pages=1850β1855 | doi= 10.1111/j.1558-5646.1998.tb02263.x| pmid=28565304| jstor=2411356| s2cid=24502821}}</ref> |- | | | |Yes | |1999<ref>{{Citation |title=The effects of selection and bottlenecks on male mating success in peripheral isolates | author=Mooers, A. Γ.| author2=Rundle, H. D. | author3=Whitlock, M. C. | journal=American Naturalist | year=1999 | volume=153 | issue= 4| pages=437β444 | doi= 10.1086/303186| pmid=29586617| bibcode=1999ANat..153..437M| s2cid=4411105}}</ref> |- | | |Direct; divergent | |Pre-zygotic |1971<ref>{{Citation |title=Natural selection and the origin of reproductive isolation | author=Lee Ehrman | journal=American Naturalist | year=1971 | volume=105 | issue= 945| pages=479β483 | doi= 10.1086/282739| bibcode=1971ANat..105..479E | s2cid=85401244 | author-link=Lee Ehrman }}</ref> 1973<ref>{{Citation |title=More on natural selection and the origin of reproductive isolation | author=Lee Ehrman | journal=American Naturalist | year=1973 | volume=107 | issue= 954| pages=318β319 | doi= 10.1086/282835| bibcode=1973ANat..107..318E | s2cid=83780632 }}</ref> 1979<ref>{{Citation |title=Still more on natural selection and the origin of reproductive isolation | author=Lee Ehrman | journal=American Naturalist | year=1979 | volume=113 | issue=1 | pages=148β150 | doi= 10.1086/283371| bibcode=1979ANat..113..148E | s2cid=85237458 }}</ref> 1983<ref>{{Citation |title=Fourth report on natural selection for the origin of reproductive isolation | author=Lee Ehrman | journal=American Naturalist | year=1983 | volume=121 | issue=3 | pages=290β293 | doi= 10.1086/284059| bibcode=1983ANat..121..290E | s2cid=83654887 }}</ref> |- |''[[Drosophila simulans|D. simulans]]'' |[[Morphology of Diptera|Scutellar bristles]], development speed, wing width; [[Desiccation tolerance|desiccation resistance]], [[fecundity]], [[ethanol]] resistance; [[courtship display]], re-mating speed, [[Lek mating|lek behavior]]; [[pupa]]tion height, clumped egg laying, general activity |3 years | |Yes |Post-zygotic |1985<ref>{{Citation |title=An Experiment Testing Two Hypotheses of Speciation | author=John Ringo| author2=David Wood| author3=Robert Rockwell| author4=Harold Dowse | journal=The American Naturalist | year=1985 | volume=126 | issue=5 | pages=642β661 | doi=10.1086/284445| bibcode=1985ANat..126..642R| s2cid=84819968}}</ref> |- | rowspan="2" |''[[Drosophila paulistorum|D. paulistorum]]'' | |131; 131 |Direct | |Pre-zygotic |1976<ref>{{Citation |title=Partially Successful Attempt to Enhance Reproductive Isolation Between Semispecies of ''Drosophila paulistorum'' | author=T. Dobzhansky| author2=O. Pavlovsky| author3=J. R. Powell | journal=Evolution | year=1976 | volume=30 | issue=2 | pages=201β212 | doi=10.2307/2407696 | pmid=28563045| jstor=2407696}}</ref> |- | |5 years | | | |1966<ref>{{Citation |title=Spontaneous origin of an incipient species in the ''Drosophila paulistorum'' complex | author=T. Dobzhansky | author2=O. Pavlovsky | journal=PNAS | year=1966 | volume=55 | issue= 4| pages=723β733 | doi= 10.1073/pnas.55.4.727| pmid=5219677 | pmc=224220 | bibcode=1966PNAS...55..727D | doi-access=free }}</ref> |- |''[[Drosophila willistoni|D. willistoni]]'' |[[pH]] adaptation |34β122 |Indirect; divergent |No |Pre-zygotic |1980<ref>{{Citation |title=Adaptation of ''Drosophila willistoni'' experimental populations to extreme pH medium | author=Alice Kalisz de Oliveira | author2=Antonio Rodrigues Cordeiro | journal=Heredity | year=1980 | volume=44 | pages=123β130 | doi=10.1038/hdy.1980.11 | doi-access=free }}</ref> |- | rowspan="6" |''[[Drosophila pseudoobscura|D. pseudoobscura]]'' |[[Carbohydrate]] source |12 |Indirect |Yes |Pre-zygotic |1989<ref name=Dodd1989>{{Citation |title=Reproductive Isolation as a Consequence of Adaptive Divergence in ''Drosophila pseudoobscura'' | author=Diane M. B. Dodd | journal=Evolution | year=1989 | volume=43 | issue=6 | pages=1308β1311 | doi=10.2307/2409365 | pmid=28564510 | jstor=2409365 }}</ref> |- |Temperature adaptation |25β60 |Direct | | |1964<ref>{{Citation |title=Genetic divergence in M. Vetukhiv's experimental populations of ''Drosophila pseudoobscura'' | author=L. Ehrman | journal=Genetical Research | year=1964 | volume=5 | pages=150β157 | doi=10.1017/s0016672300001099| doi-access=free }}</ref> 1969<ref>{{Citation |title=Genetic divergence in M. Vetukhiv's experimental populations of ''Drosophila pseudoobscura''. 5. A further study of rudiments of sexual isolation | author=L. Ehrman | journal=American Midland Naturalist | year=1969 | volume=82 | issue=1 | pages=272β276 | doi=10.2307/2423835| jstor=2423835 }}</ref> |- |Phototaxis, geotaxis |5β11 |Indirect |No |Pre-zygotic |1966<ref>{{Citation |title=Sexual isolation caused by selection for positive and negative phototaxis and geotaxis in ''Drosophila pseudoobscura'' | author=Eduardo del Solar | journal=Proceedings of the National Academy of Sciences | year=1966 | volume=56 | issue=2 | pages=484β487 | doi = 10.1073/pnas.56.2.484 | pmid=5229969 | pmc=224398 | bibcode=1966PNAS...56..484D | doi-access=free }}</ref> |- | | | | |Pre-zygotic |1978<ref>{{Citation | title=The Founder-Flush Speciation Theory: An Experimental Approach| author=Jeffrey R. Powell| journal=Evolution| year=1978| volume=32| issue=3| pages=465β474| doi=10.1111/j.1558-5646.1978.tb04589.x| pmid=28567948| jstor=2407714| s2cid=30943286| doi-access=free}}</ref> 1985<ref>{{Citation|title=Founder-Flush Speciation: An Update of Experimental Results with ''Drosophila''|year=1985|author=Diane M. B. Dodd | author2=Jeffrey R. Powell|journal=Evolution|volume=39|issue=6|pages=1388β1392|doi=10.1111/j.1558-5646.1985.tb05704.x|pmid=28564258|jstor=2408795|s2cid=34137489|doi-access=free}}</ref> |- | | | |Yes | |1993<ref>{{Citation |title=Founder-flush speciation in ''Drosophila pseudoobscura'': a large scale experiment | author=Galiana, A.| author2=Moya, A. | author3=Ayala, F. J. | journal=Evolution | year=1993 | volume=47 | issue= 2| pages=432β444 | doi= 10.1111/j.1558-5646.1993.tb02104.x| pmid=28568735| jstor=2410062| s2cid=42232235| doi-access=free }}</ref> |- |Temperature [[Photoperiodism|photoperiod]]; food |37 |Divergent |Yes |None |2003<ref>{{Citation |title=Divergent environments and population bottlenecks fail to generate premating isolation in ''Drosophila pseudoobscura'' | author=Rundle, H. D. | journal=Evolution | year=2003 | volume=57 | issue= 11| pages=2557β2565 | doi= 10.1554/02-717| pmid=14686531 | s2cid=6162106 }}</ref> |- | rowspan="2" |''D.pseudoobscura &'' ''[[Drosophila persimilis|D. persimilis]]'' | |22; 16; 9 |Direct; divergent | |Pre-zygotic |1950<ref>{{Citation |title=Natural Selection for Reproductive Isolation Between ''Drosophila pseudoobscura'' and ''Drosophila persimilis'' | author=Karl F. Koopman | journal=Evolution | year=1950 | volume=4 | issue=2 | pages=135β148 | doi=10.2307/2405390 | jstor=2405390 }}</ref> |- | |4 experiments, 18 each |Direct | |Pre-zygotic |1966<ref>{{Citation |title=Selection For and Against Ethological Isolation Between ''Drosophila pseudoobscura'' and ''Drosophila persimilis'' | author=Seymour Kessler | journal=Evolution | year=1966 | volume=20 | issue=4 | pages=634β645 | doi=10.2307/2406597 | pmid=28562900 | jstor=2406597 }}</ref> |- | rowspan="2" |''[[Drosophila mojavensis|D. mojavensis]]'' | |12 |Direct | |Pre-zygotic |1987<ref>{{Citation |title=Selection for Sexual Isolation Between Geographic Forms of ''Drosophila mojavensis''. I Interactions Between the Selected Forms | author=H. Roberta Koepfer | journal=Evolution | year=1987 | volume=41 | issue=1 | pages=37β48 | doi=10.2307/2408971 | pmid=28563762 | jstor=2408971 }}</ref> |- |Development time |13 |Divergent |Yes |None |1998<ref>{{Citation |title=Premating isolation is determined by larval rearing substrates in cactophilis ''Drosophila mojavensis''. IV. Correlated responses in behavioral isolation to artificial selection on a life-history trait | author=Etges, W. J. | s2cid=17689372 | journal=American Naturalist | year=1998 | volume=152 | issue= 1| pages=129β144 | doi= 10.1086/286154| pmid=18811406 }}</ref> |- |''[[Drosophila adiastola|D. adiastola]]'' | | | |Yes |Pre-zygotic |1974<ref>{{Citation |title=Ethological Isolation Between Two Stocks of ''Drosophila Adiastola'' Hardy | author=Lorna H. Arita | author2=Kenneth Y. Kaneshiro | journal=Proc. Hawaii. Entomol. Soc. | year=1979 | volume=13 | pages=31β34 }}</ref> |- |''[[Drosophila silvestris|D. silvestris]]'' | | | |Yes | |1980<ref>{{Citation |title=Evolution of behavioral reproductive isolation in a laboratory stock of ''Drosophila silvestris'' | author=J. N. Ahearn | journal=Experientia | year=1980 | volume=36 | issue=1 | pages=63β64 | doi=10.1007/BF02003975 | s2cid=43809774 }}</ref> |- | rowspan="3" |''[[Housefly|Musca domestica]]'' |Geotaxis |38 |Indirect |No |Pre-zygotic |1974<ref>{{Citation |title=Evolution of Reproductive Isolation in Allopatric and Sympatric Populations | author=A. Benedict Soans| author2=David Pimentel| author3=Joyce S. Soans | journal=The American Naturalist | year=1974 | volume=108 | issue=959 | pages=117β124 | doi=10.1086/282889| bibcode=1974ANat..108..117S| s2cid=84913547}}</ref> |- |Geotaxis |16 |Direct; divergent |No |Pre-zygotic |1975<ref>{{Citation |title=Divergent Selection for Geotactic Response and Evolution of Reproductive Isolation in Sympatric and Allopatric Populations of Houseflies | author=L. E. Hurd | author2=Robert M. Eisenberg | journal=The American Naturalist | year=1975 | volume=109 | issue=967 | pages=353β358 | doi=10.1086/283002| bibcode=1975ANat..109..353H | s2cid=85084378 }}</ref> |- | | | |Yes | |1991<ref>{{Citation |title=Mating propensity and courtship behavior in serially bottlenecked lines of the housefly | author=Meffert, L. M. | author2=Bryant, E. H. | journal=Evolution | year=1991 | volume=45 | issue= 2| pages=293β306 | doi= 10.1111/j.1558-5646.1991.tb04404.x| pmid=28567864 | jstor=2409664 | s2cid=13379387 }}</ref> |- |''[[Melon fly|Bactrocera cucurbitae]]'' |Development time |40β51 |Divergent |Yes |Pre-zygotic |1999<ref>{{Citation |title=Genetic correlations between life-history and behavioral traits can cause reproductive isolation | author=Takahisa Miyatake | author2=Toru Shimizu | journal=Evolution | year=1999 | volume=53 | issue=1 | pages= 201β208 | doi=10.2307/2640932 | pmid=28565193 | jstor=2640932 }}</ref> |- |''[[Zea mays]]'' | |6; 6 |Direct; divergent | |Pre-zygotic |1969<ref>{{Citation |title=Selection for Reproductive Isolation between Two Populations of Maize, ''Zea mays'' L | author=Paterniani, E. | journal=Evolution | year=1969 | volume=23 | issue= 4| pages=534β547 | doi= 10.1111/j.1558-5646.1969.tb03539.x| pmid=28562870 | s2cid=38650254 | doi-access=free }}</ref> |- |[[Drosophila grimshawi|''D. grimshawi'']] | | | | | |<ref>{{Citation |title=Sexual selection and peripatric speciation: the Kaneshiro model revisited | author=Anders Γdeen | author2=Ann-Britt Florin | journal=Journal of Evolutionary Biology | year=2002 | volume=15 | issue= 2| pages=301β306 | doi= 10.1046/j.1420-9101.2002.00378.x| s2cid=82095639 | doi-access=free }}</ref> |}
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