Laboratory experiments of speciation

Biological experiments

A simplification of an allopatric speciation experiment where two lines of fruit flies are raised on maltose and starch media

Laboratory experiments of speciation have been conducted for all four modes of speciation: allopatric, peripatric, parapatric, and sympatric; and various other processes involving speciation: hybridization, reinforcement, founder effects, among others. Most of the experiments have been done on flies, in particular Drosophila fruit flies.[1] However, more recent studies have tested yeasts, fungi, and even viruses.

It has been suggested that laboratory experiments are not conducive to vicariant speciation events (allopatric and peripatric) due to their small population sizes and limited generations.[2] Most estimates from studies of nature indicate that speciation takes hundreds of thousands to millions of years.[3] On the other hand, many species are thought to have speciated faster and more recently, such as the European flounders (Platichthys flesus) that spawn in pelagic and demersal zones—having allopatrically speciated in under 3000 generations.[4]

## Table of experiments

[edit]

Six publications have attempted to compile, review, and analyze the experimental research on speciation:

1. John Ringo, David Wood, Robert Rockwell, and Harold Dowse in 1985;[5]
2. William R. Rice and Ellen E. Hostert in 1993;[6]
3. Ann-Britt Florin and Anders Ödeen in 2002;[2]
4. Mark Kirkpatrick and Virginie Ravigné in 2002;[7]
5. Jerry A. Coyne and H. Allen Orr in 2004;[1] and
6. James D. Fry in 2009.[8]

The table summarizes the studies and data reviewed in these publications. It also references several contemporary experiments and is non-exhaustive. In the table, multiple numbers separated by semi-colons in the generations column indicate that multiple experiments were conducted. The replications (in parentheses) indicates the number of populations used in the experiments—i.e. how many times the experiment was replicated. Various types of selection have been imposed on experimental populations and are indicated by the selection type column. Negative or positive results of each experiment are provided by the reproductive isolation column. Pre-zygotic reproductive isolation means that the reproducing individuals in the populations were unable to produce offspring (effectively a positive result). Post-zygotic isolation means that the reproducing individuals were able to produce offspring but they were either sterile or inviable (a positive result as well). Negative results are indicated by "none"—that is, the experiments did not result in reproductive isolation.

Laboratory experiments of speciation[1][6][7][2][8] Species | Trait  | Generations (replications) [duration]  | Tested  | Selection type  | Studied genetic drift | Reproductive isolation | Reference  | Year

|
|
|
|
|
|
|
|

Drosophila melanogaster | Escape response | 18  | Vicariant, reinforcement, parapatric/

sympatric

| Indirect; divergent  | Yes  | Pre-zygotic  | Grant & Mettler[9] | 1969
D. melanogaster | Locomotion | 112  | Vicariant  | Indirect; divergent  | No  | Pre-zygotic  | Burnet & Connolly[10] | 1974
D. melanogaster | Temperature, humidity | 70–130  | Vicariant  | Indirect; divergent  | Yes  | Pre-zygotic  | Kilias et al.[11] | 1980
D. melanogaster | DDT adaptation  | 600 [25 years, +15 years]  | Vicariant  | Direct  | No  | Pre-zygotic  | Boake et al.[12] | 2003
D. melanogaster |  | 17, 9, 9, 1, 1, 7, 7, 7, 7  | Vicariant; parapatric/

sympatric

| Direct, divergent  |  | Pre-zygotic in vicariance; none with gene flow  | Barker & Karlsson[13] | 1974
D. melanogaster |  | 40; 50  | Reinforcement  | Direct; divergent  |  | Pre-zygotic  | Crossley[14] | 1974
D. melanogaster | Locomotion  | 45  | Vicariant  | Direct; divergent  | No  | None  | van Dijken & Scharloo[15][16] | 1979
D. melanogaster |  |  | Reinforcement  | Direct; divergent  |  | Pre-zygotic  | Wallace[17] | 1953
D. melanogaster |  | 36; 31  | Reinforcement  | Direct; divergent  |  | Pre-zygotic  | Knight[18] | 1956
D. melanogaster | EDTA adaptation  | 25, 25, 25, 14  | Semi-allopatric, reinforcement  | Indirect; divergent  | No  | Post-zygotic  | Robertson[19][20] | 1966
D. melanogaster |  | 25 (8)  | Vicariant; reinforcement; parapatric; sympatric  | Direct  |  | None  | Hostert[21] | 1997
D. melanogaster | Abdominal chaeta

number

| 21–31  | Vicariant  | Direct  | Yes  | None  | Santibanez & Waddington[22] | 1958
D. melanogaster | Sternopleural chaeta number  | 32  | Vicariant, reinforcement, parapatric/

sympatric

| Direct  | No  | None  | Barker & Cummins[23] | 1969
D. melanogaster | Phototaxis, geotaxis | 20  | Vicariant  |  | No  | None  | Markow[24][25] | 1975; 1981
D. melanogaster |  |  | Peripatric  |  | Yes  |  | Rundle et al.[26] | 1998
D. melanogaster |  |  | Vicariant; peripatric  |  | Yes  |  | Mooers et al.[27] | 1999
D. melanogaster |  | 12  | Reinforcement  | Divergent  |  | Pre-zygotic  | Thoday & Gibson[28] | 1962
D. melanogaster |  |  |  |  |  | None  | Thoday & Gibson[29][30] | 1970; 1971
D. melanogaster |  | 16  | Reinforcement  | Indirect  |  | None  | Spiess & Wilke[31] | 1954
D. melanogaster |  |  | Reinforcement  | Direct; divergent  |  | Pre-zygotic  | Ehrman[32][33][34][35] | 1971; 1973; 1979; 1983
D. melanogaster | Sternopleural chaeta number  | 5; 27; 27; 1; 1; 1; 1; 1  | Parapatric/

sympatric

|  |  | None  | Chabora[36] | 1968
D. melanogaster |  |  |  |  |  | None  | Scharloo[37] | 1967
D. melanogaster |  | 1, 1  |  |  |  |  | Coyne & Grant[38] | 1972
D. melanogaster |  | 25  |  |  |  |  | Rice[39] | 1985
D. melanogaster |  | 25  |  | Disruptive  |  | Pre-zygotic  | Rice & Salt[40] | 1988
D. melanogaster |  | 35; 35  | Sympatric  |  |  | Pre-zygotic  | Rice & Salt[41] | 1990
D. melanogaster | NaCl and CuSO4 levels in food  | [3 years in allopatry, 1 in sympatry]  | Allopatric; reinforcement; sympatric  |  |  | Pre-zygotic in allopatry, none in sympatry  | Wallace[42] | 1982
D. melanogaster |  |  | Reinforcement  |  |  |  | Ehrman et al.[43] | 1991
D. melanogaster |  |  | Reinforcement  |  |  |  | Fukatami & Moriwaki[44] | 1970
Drosophila simulans | Scutellar bristles, development speed, wing width; desiccation resistance, fecundity, ethanol resistance; courtship display, re-mating speed, lek behavior; pupation height, clumped egg laying, general activity  | [3 years]  | Vicariant; peripatric  |  | Yes  | Post-zygotic  | Ringo et al.[5] | 1985
Drosophila paulistorum |  | 131; 131  | Reinforcement  | Direct  |  | Pre-zygotic  | Dobzhansky et al.[45] | 1976
D. paulistorum |  | [5 years]  | Vicariant  |  |  |  | Dobzhansky and Pavlovsky[46] | 1966
Drosophila willistoni | pH adaptation  | 34–122  | Vicariant  | Indirect; divergent  | No  | Pre-zygotic  | Kalisz & Cordeiro[47] | 1980
Drosophila pseudoobscura | Carbohydrate source  | 12  | Vicariant  | Indirect  | Yes  | Pre-zygotic  | Dodd[48] | 1989
D. pseudoobscura | Temperature adaptation  | 25–60  | Vicariant  | Direct  |  |  | Ehrman[49][50][51][52][53] | 1964;

1969

D. pseudoobscura | Phototaxis, geotaxis  | 5–11  | Vicariant  | Indirect  | No  | Pre-zygotic  | del Solar[54] | 1966
D. pseudoobscura |  |  | Vicariant; peripatric  |  |  | Pre-zygotic  | Powell[55][56] | 1978; 1985
D. pseudoobscura |  |  | Peripatric; vicariant  |  | Yes  |  | Galiana et al.[57] | 1993
D. pseudoobscura | Temperature photoperiod; food  | 37 (78) [33–34 months]  | Vicariant  | Divergent  | Yes  | None  | Rundle[58] | 2003
D. pseudoobscura &

Drosophila persimilis

|  | 22; 16; 9  | Reinforcement  | Direct; divergent  |  | Pre-zygotic  | Koopman[59] | 1950
D. pseudoobscura &

D. persimilis

|  | 18 (4)  |  | Direct  |  | Pre-zygotic  | Kessler[60] | 1966
Drosophila mojavensis |  | 12  |  | Direct  |  | Pre-zygotic  | Koepfer[61] | 1987
D. mojavensis | Development time  | 13  |  | Divergent  | Yes  | None  | Etges[62] | 1998
Drosophila adiastola |  |  | Peripatric  |  | Yes  | Pre-zygotic  | Arita & Kaneshiro[63] | 1974
Drosophila silvestris |  |  | Peripatric  |  | Yes  |  | Ahearn[64] | 1980
Musca domestica | Geotaxis  | 38  | Vicariant  | Indirect  | No  | Pre-zygotic  | Soans et al.[65] | 1974
M. domestica | Geotaxis  | 16  | Vicariant  | Direct; divergent  | No  | Pre-zygotic  | Hurd & Eisenburg[66] | 1975
M. domestica |  |  | Peripatric  |  | Yes  |  | Meffert & Bryant[67] | 1991
M. domestica |  |  |  |  |  |  | Regan et al.[68] | 2003
Bactrocera cucurbitae | Development time  | 40–51  |  | Divergent  | Yes  | Pre-zygotic  | Miyatake & Shimizu[69] | 1999
Zea mays |  | 6; 6  | Reinforcement  | Direct; divergent  |  | Pre-zygotic  | Paterniani[70] | 1969
Drosophila grimshawi |  |  | Peripatric  |  |  |  | Jones, Widemo, & Arrendal[71] | N/A
Saccharomyces cerevisiae |  |  |  |  |  |  | Leu & Murry[72] | 2006
D. melanogaster |  |  | Reinforcement  |  |  |  | Harper & Lambert[73] | 1983
Tribolium castaneum | Pupal weight  | 15 (6)  |  | Disruptive  |  |  | Halliburton & Gall[74] | 1983
D. melanogaster | Geotaxis  |  |  | Divergent  |  |  | Lofdahl et al.[75] | 1992
D. pseudoobscura  |  | [10 years]  |  |  |  |  | Moya et al.[76] | 1995
Neurospora |  |  |  | Divergent  |  |  | Dettman et al.[77] | 2008
S. cerevisiae |  | 500  |  | Divergent  |  |  | Dettman et al.[78] | 2007
Sepsis cynipsea |  | 35  |  |  |  |  | Martin & Hosken[79] | 2003
D. melanogaster |  |  |  |  |  |  | Wigby & Chapman[80] | 2006
D. pseudoobscura | Sexual conflict  | 48–52 (4; 4; 4)  |  |  |  |  | Bacigalupe et al.[81] | 2007
D. serrata |  |  |  |  |  |  | Rundle et al.[82] | 2005
Drosophila serrata & D. birchii | Mate recognition  | 9 (3; 3)  | Reinforcement  | Natural  |  | Pre-zygotic  | Higgie et al.[83] | 2000
Enterobacteria phage λ | Escherichia coli receptor exploitation  | 35 cycles (6)  | Vicariant, sympatric  |  |  | Pre-zygotic  | Meyer et al.[84] | 2016
Tetranychus urticae | Resistance to host plant toxin  |  |  |  |  |  | Overmeer[85] | 1966
T. urticae | Resistance to host plant toxin  |  |  |  |  |  | Fry[86] | 1999
Helianthus annus × H. petiolaris and H. anomalus |  |  | Hybrid  |  |  |  | Rieseburg et al.[87] | 1996
S. cerevisiae |  |  |  |  |  |  | Greig et al.[88] | 2002
D. melanogaster | Life history |  |  |  |  |  | Ghosh & Joshi[89] | 2012
Drosophila subobscura | Mate behavior  |  |  |  |  |  | Bárbaro et al.[90] | 2015
Digital organisms  |  | ~42,000; ~850 (20)  | Ecological  |  |  | Post-zygotic  | Anderson & Harmon[91] | 2014
Schizosaccharomyces pombe |  |  |  |  |  | Complete reproductive isolation  | Seike et al.[92] | 2015
D. pseudoobscura | Courtship song  | 130  |  |  |  |  | Debelle et al.[93] | 2014
Callosobruchus maculatus |  | 40 (16)  |  |  |  |  | Debelle et al.[94] | 2010

## See also

[edit]

* Directed evolution

## References

[edit]

1. ^ a b c Coyne, Jerry A.; Orr, H. Allen (2004), Speciation, Sinauer Associates, pp. 1–545, ISBN 978-0-87893-091-3
2. ^ a b c Florin, Ann-Britt & Ödeen, Anders (2002), "Laboratory environments are not conducive for allopatric speciation", Journal of Evolutionary Biology, 15 (1): 10–19, doi:10.1046/j.1420-9101.2002.00356.x`{{citation}}`: CS1 maint: multiple names: authors list (link)
3. ^ Coyne, Jerry A.; Orr, H. Allen (1997), ""Patterns of Speciation in Drosophila" Revisited", Evolution, 51 (1): 295–303, doi:10.1111/j.1558-5646.1997.tb02412.x, PMID 28568795
4. ^ Momigliano, Paolo; Jokinen, Henri; Fraimout, Antoine; Florin, Ann-Britt; Norkko, Alf; Merilä, Juha (2017), "Extraordinarily rapid speciation in a marine fish" (PDF), PNAS, 114 (23): 6074–6079, Bibcode:2017PNAS..114.6074M, doi:10.1073/pnas.1615109114, PMC 5468626, PMID 28533412
5. ^ a b Ringo, John; Wood, David; Rockwell, Robert; Dowse, Harold (1985), "An Experiment Testing Two Hypotheses of Speciation", The American Naturalist, 126 (5): 642–661, Bibcode:1985ANat..126..642R, doi:10.1086/284445, S2CID 84819968
6. ^ a b Rice, William R. & Hostert, Ellen E. (1993), "Laboratory Experiments on Speciation: What Have We Learned in 40 Years?", Evolution, 47 (6): 1637–1653, doi:10.1111/j.1558-5646.1993.tb01257.x, PMID 28568007, S2CID 42100751`{{citation}}`: CS1 maint: multiple names: authors list (link)
7. ^ a b Kirkpatrick, Mark & Ravigné, Virginie (2002), "Speciation by Natural and Sexual Selection: Models and Experiments", The American Naturalist, 159 (S3): S22 – S35, Bibcode:2002ANat..159S..22K, doi:10.1086/338370, PMID 18707367, S2CID 16516804`{{citation}}`: CS1 maint: multiple names: authors list (link)
8. ^ a b Fry, James D. (2009). Laboratory Experiments on Speciation. In Garland, Theodore & Rose, Michael R. "Experimental Evolution: Concepts, Methods, and Applications of Selection Experiments". Pp. 631–656. doi:10.1525/california/9780520247666.003.0020
9. ^ Grant, B. S. & Mettler, L. E. (1969), "Disruptive and stabilizing selection on the" escape" behavior of Drosophila melanogaster", Genetics, 62 (3): 625–637, doi:10.1093/genetics/62.3.625, PMC 1212303, PMID 17248452`{{citation}}`: CS1 maint: multiple names: authors list (link)
10. ^ Burnet, B. & Connolly, K. (1974). Activity and sexual behavior in Drosophila melanogaster. In Abeelen, J. H. V. F. (eds). The Genetics of Behaviour. North-Holland, Amsterdam. Pp. 201–258.
11. ^ Kilias, G., Alahiotis, S. N., & Pelecanos, M. (1980), "A Multifactorial Genetic Investigation of Speciation Theory Using Drosophila melanogaster", Evolution, 34 (4): 730–737, doi:10.2307/2408027, JSTOR 2408027, PMID 28563991`{{citation}}`: CS1 maint: multiple names: authors list (link)
12. ^ Boake, C. R. B., Mcdonald, K., Maitra, S., Ganguly, R. (2003), "Forty years of solitude: life-history divergence and behavioural isolation between laboratory lines of Drosophila melanogaster", Journal of Evolutionary Biology, 16 (1): 83–90, doi:10.1046/j.1420-9101.2003.00505.x, PMID 14635883`{{citation}}`: CS1 maint: multiple names: authors list (link)
13. ^ Barker, J. S. F. & Karlsson, L. J. E. (1974), "Effects of population size and selection intensity on responses to disruptive selection in Drosophila melanogaster", Genetics, 78 (2): 715–735, doi:10.2307/2407287, JSTOR 2407287, PMC 1213230, PMID 4217303`{{citation}}`: CS1 maint: multiple names: authors list (link)
14. ^ Crossley, Stella A. (1974), "Changes in Mating Behavior Produced by Selection for Ethological Isolation Between Ebony and Vestigial Mutants of Drosophila melanogaster", Evolution, 28 (4): 631–647, doi:10.1111/j.1558-5646.1974.tb00795.x, PMID 28564833, S2CID 35867118
15. ^ van Dijken, F. R. & Scharloo, W. (1979), "Divergent selection on locomotor activity in Drosophila melanogaster. I. Selection response", Behavior Genetics, 9 (6): 543–553, doi:10.1007/BF01067350, PMID 122270, S2CID 39352792`{{citation}}`: CS1 maint: multiple names: authors list (link)
16. ^ van Dijken, F. R. & Scharloo, W. (1979), "Divergent selection on locomotor activity in Drosophila melanogaster. II. Test for reproductive isolation between selected lines", Behavior Genetics, 9 (6): 555–561, doi:10.1007/BF01067351, PMID 122271, S2CID 40169222`{{citation}}`: CS1 maint: multiple names: authors list (link)
17. ^ Wallace, B. (1953), "Genetic divergence of isolated populations of Drosophila melanogaster", Proceedings of the Ninth International Congress of Genetics, 9: 761–764
18. ^ Knight, G. R., Robertson, Alan, & Waddington, C. H. (1956), "Selection for sexual isolation within a species", Evolution, 10 (1): 14–22, doi:10.1111/j.1558-5646.1956.tb02825.x`{{citation}}`: CS1 maint: multiple names: authors list (link)
19. ^ Robertson, Forbes W. (1966), "A test of sexual isolation in Drosophila", Genetical Research, 8 (2): 181–187, doi:10.1017/S001667230001003X, PMID 5922518
20. ^ Robertson, Forbes W. (1966), "The ecological genetics of growth in Drosophila 8\. Adaptation to a New Diet", Genetical Research, 8 (2): 165–179, doi:10.1017/S0016672300010028, PMID 5922517
21. ^ Hostert, Ellen E. (1997), "Reinforcement: a new perspective on an old controversy", Evolution, 51 (3): 697–702, doi:10.1111/j.1558-5646.1997.tb03653.x, PMID 28568598
22. ^ Koref Santibañez, S. & Waddington, C. H. (1958), "The origin of sexual isolation between different lines within a species", Evolution, 12 (4): 485–493, doi:10.2307/2405959, JSTOR 2405959`{{citation}}`: CS1 maint: multiple names: authors list (link)
23. ^ Barker, J. S. F. & Cummins, L. J. (1969), "The effect of selection for sternopleural bristle number in mating behaviour in Drosophila melanogaster", Genetics, 61 (3): 713–719, doi:10.1093/genetics/61.3.713, PMC 1212235, PMID 17248436`{{citation}}`: CS1 maint: multiple names: authors list (link)
24. ^ Markow, Therese Ann (1975), "A genetic analysis of phototactic behavior in Drosophila melanogaster", Genetics, 79 (3): 527–534, doi:10.1093/genetics/79.3.527, PMC 1213291, PMID 805084
25. ^ Markow, Therese Ann (1981), "Mating preferences are not predictive of the direction of evolution in experimental populations of Drosophila", Science, 213 (4514): 1405–1407, Bibcode:1981Sci...213.1405M, doi:10.1126/science.213.4514.1405, PMID 17732575, S2CID 15497733
26. ^ Rundle, H. D., Mooers, Arne Ø. & Whitlock, Michael C. (1998), "Single founder-flush events and the evolution of reproductive isolation", Evolution, 52 (6): 1850–1855, doi:10.1111/j.1558-5646.1998.tb02263.x, PMID 28565304, S2CID 24502821`{{citation}}`: CS1 maint: multiple names: authors list (link)
27. ^ Mooers, Arne Ø., Rundle, Howard D. & Whitlock, Michael C. (1999), "The effects of selection and bottlenecks on male mating success in peripheral isolates", American Naturalist, 153 (4): 437–444, Bibcode:1999ANat..153..437M, doi:10.1086/303186, PMID 29586617, S2CID 4411105`{{citation}}`: CS1 maint: multiple names: authors list (link)
28. ^ Thoday, J. M. & Gibson, J. B. (1962), "Isolation by disruptive selection", Nature, 193 (4821): 1164–1166, Bibcode:1962Natur.193.1164T, doi:10.1038/1931164a0, PMID 13920720, S2CID 5156234`{{citation}}`: CS1 maint: multiple names: authors list (link)
29. ^ Thoday, J. M. & Gibson, J. B. (1970), "The probability of isolation by disruptive selection", Nature, 104 (937): 219–230, Bibcode:1970ANat..104..219T, doi:10.1086/282656, S2CID 85333360`{{citation}}`: CS1 maint: multiple names: authors list (link)
30. ^ Scharloo, W. (1971), "Reproductive isolation by disruptive selection: Did it occur?", American Naturalist, 105 (941): 83–86, Bibcode:1971ANat..105...83S, doi:10.1086/282706, S2CID 84204545
31. ^ Spiess, E. B. & Wilke, C. M. (1984), "Still another attempt to achieve assortive mating by disruptive selection in Drosophila", Evolution, 38 (3): 505–515, doi:10.2307/2408700, JSTOR 2408700, PMID 28555983`{{citation}}`: CS1 maint: multiple names: authors list (link)
32. ^ Ehrman, Lee (1971), "Natural selection and the origin of reproductive isolation", American Naturalist, 105 (945): 479–483, Bibcode:1971ANat..105..479E, doi:10.1086/282739, S2CID 85401244
33. ^ Ehrman, Lee (1973), "More on natural selection and the origin of reproductive isolation", American Naturalist, 107 (954): 318–319, Bibcode:1973ANat..107..318E, doi:10.1086/282835, S2CID 83780632
34. ^ Ehrman, Lee (1979), "Still more on natural selection and the origin of reproductive isolation", American Naturalist, 113 (1): 148–150, Bibcode:1979ANat..113..148E, doi:10.1086/283371, S2CID 85237458
35. ^ Ehrman, Lee (1983), "Fourth report on natural selection for the origin of reproductive isolation", American Naturalist, 121 (2): 290–293, Bibcode:1983ANat..121..290E, doi:10.1086/284059, S2CID 83654887
36. ^ Chabora, Alice J. (1968), "Disruptive selection for sternopleural chaeta number in various strains of Drosophila melanogaster", American Naturalist, 102 (928): 525–532, Bibcode:1968ANat..102..525C, doi:10.1086/282565, S2CID 84885812
37. ^ Scharloo, W., Hoogmoed, M. S. & Kuile, A. T. (1967), "Stabilizing and disruptive selection on a mutant character in Drosophila. I. The phenotypic variance and its components.", Genetics, 56 (4): 709–726, doi:10.1093/genetics/56.4.709, PMC 1211648, PMID 6061662`{{citation}}`: CS1 maint: multiple names: authors list (link)
38. ^ Coyne, Jerry A. & and Grant, Bruce (1972), "Disruptive selection on I-maze activity in Drosophila melanogaster", Genetics, 71 (1): 185–188, doi:10.1093/genetics/71.1.185, PMC 1212770, PMID 17248572`{{citation}}`: CS1 maint: multiple names: authors list (link)
39. ^ Rice, W. R. (1985), "Disruptive selection on habitat preference and the evolution of reproductive isolation: an exploratory experiment", Evolution, 39 (3): 645–656, doi:10.1111/j.1558-5646.1985.tb00401.x, PMID 28561974
40. ^ Rice, William R. & Salt, George, W. (1988), "Speciation via disruptive selection on habitat preference", American Naturalist, 131 (6): 911–917, doi:10.1086/284831, S2CID 84876223`{{citation}}`: CS1 maint: multiple names: authors list (link)
41. ^ Rice, William R. & Salt, George, W. (1990), "The evolution of reproductive isolation as a correlated character under sympatric conditions: experimental evidence", Evolution, 44 (5): 1140–1152, doi:10.2307/2409278, JSTOR 2409278, PMID 28563894`{{citation}}`: CS1 maint: multiple names: authors list (link)
42. ^ Wallace, B. (1982), "Drosophila melanogaster populations selected for resistances to NaCl and CuSO4 in both allopatry and sympatry", Journal of Heredity, 73 (1): 35–42, doi:10.1093/oxfordjournals.jhered.a109572, PMID 6802898
43. ^ Ehrman, Lee, White, Marney A. & Wallace, B. (1991). A long-term study involving Drosophila melanogaster and toxic media. In Hecht, M. K., Wallace, B., & Maclntyre, R. J. (eds). Evolutionary biology, vol. 25. Plenum Press, New York. Pp. 175–209
44. ^ Fukatami, A & Moriwaki, D. (1970), "Selection for sexual isolation in Drosophila melanogaster by a modification of Koopman's method", The Japanese Journal of Genetics, 45 (3): 193–204, doi:10.1266/jjg.45.193`{{citation}}`: CS1 maint: multiple names: authors list (link)
45. ^ Dobzhansky, Theodosius; Pavlovsky, O.; Powell, J. R. (1976), "Partially Successful Attempt to Enhance Reproductive Isolation Between Semispecies of Drosophila paulistorum", Evolution, 30 (2): 201–212, doi:10.2307/2407696, JSTOR 2407696, PMID 28563045
46. ^ Dobzhansky, Theodosius & Pavlovsky, O. (1966), "Spontaneous origin of an incipient species in the Drosophila paulistorum complex", Proceedings of the National Academy of Sciences, 55 (4): 723–733, Bibcode:1966PNAS...55..727D, doi:10.1073/pnas.55.4.727, PMC 224220, PMID 5219677`{{citation}}`: CS1 maint: multiple names: authors list (link)
47. ^ de Oliveira, Alice Kalisz & Cordeiro, Antonio Rodrigues (1980), "Adaptation of Drosophila willistoni experimental populations to extreme pH medium", Heredity, 44 (1): 123–130, Bibcode:1980Hered..44..123D, doi:10.1038/hdy.1980.11`{{citation}}`: CS1 maint: multiple names: authors list (link)
48. ^ Dodd, Diane M. B. (1989), "Reproductive Isolation as a Consequence of Adaptive Divergence in Drosophila pseudoobscura", Evolution, 43 (6): 1308–1311, doi:10.2307/2409365, JSTOR 2409365, PMID 28564510
49. ^ Ehrman, Lee (1964), "Genetic divergence in M. Vetukhiv's experimental populations of Drosophila pseudoobscura 1\. Rudiments of sexual isolation", Genetical Research, 5: 150–157, doi:10.1017/S0016672300001099
50. ^ Mouradael, K. (1965), "Genetic divergence in M. Vetukhiv's experimental populations of Drosophila pseudoobscura 2\. Longevity", Genetical Research, 6: 139–146, doi:10.1017/S0016672300004006, PMID 14297592
51. ^ Anderson, Wyatt, W. (1966), "Genetic divergence in M. Vetukhiv's experimental populations of Drosophila pseudoobscura 3. Divergence in Body Size", Genetical Research, 7 (2): 255–266, doi:10.1017/S0016672300009666`{{citation}}`: CS1 maint: multiple names: authors list (link)
52. ^ Kitagawa, Osamu (1967), "Genetic divergence in M. Vetukhiv's experimental populations of Drosophila pseudoobscura: IV. Relative viability", Genetical Research, 10 (7): 303–312, doi:10.1017/S001667230001106X
53. ^ Ehrman, Lee (1969), "Genetic divergence in M. Vetukhiv's experimental populations of Drosophila pseudoobscura. 5. A further study of rudiments of sexual isolation", American Midland Naturalist, 82 (1): 272–276, doi:10.2307/2423835, JSTOR 2423835
54. ^ del Solar, Eduardo (1966), "Sexual isolation caused by selection for positive and negative phototaxis and geotaxis in Drosophila pseudoobscura", Proceedings of the National Academy of Sciences, 56 (2): 484–487, doi:10.1073/pnas.56.2.484, PMC 224398, PMID 5229969
55. ^ Powell, Jeffrey R. (1978), "The Founder-Flush Speciation Theory: An Experimental Approach", Evolution, 32 (3): 465–474, doi:10.2307/2407714, JSTOR 2407714, PMID 28567948
56. ^ Dodd, Diane M. B. & Powell, Jeffrey R. (1985), "Founder-Flush Speciation: An Update of Experimental Results with Drosophila", Evolution, 39 (6): 1388–1392, doi:10.1111/j.1558-5646.1985.tb05704.x, PMID 28564258`{{citation}}`: CS1 maint: multiple names: authors list (link)
57. ^ Galiana, Augustí, Moya, Andres & Ayala, Francisco J. (1993), "Founder-flush speciation in Drosophila pseudoobscura: a large scale experiment", Evolution, 47 (2): 432–444, doi:10.1111/j.1558-5646.1993.tb02104.x, PMID 28568735`{{citation}}`: CS1 maint: multiple names: authors list (link)
58. ^ Rundle, Howard D. (2003), "Divergent environments and population bottlenecks fail to generate premating isolation in Drosophila pseudoobscura", Evolution, 57 (11): 2557–2565, doi:10.1111/j.0014-3820.2003.tb01499.x, PMID 14686531
59. ^ Koopman, Karl F. (1950), "Natural Selection for Reproductive Isolation Between Drosophila pseudoobscura and Drosophila persimilis", Evolution, 4 (2): 135–148, doi:10.2307/2405390, JSTOR 2405390
60. ^ Kessler, Seymour (1966), "Selection For and Against Ethological Isolation Between Drosophila pseudoobscura and Drosophila persimilis", Evolution, 20 (4): 634–645, doi:10.2307/2406597, JSTOR 2406597, PMID 28562900
61. ^ Koepfer, H. Roberta (1987), "Selection for Sexual Isolation Between Geographic Forms of Drosophila mojavensis. I Interactions Between the Selected Forms", Evolution, 41 (1): 37–48, doi:10.2307/2408971, JSTOR 2408971, PMID 28563762
62. ^ Etges, W. J. (1998), "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", American Naturalist, 152 (1): 129–144, doi:10.1086/286154, PMID 18811406, S2CID 17689372
63. ^ Arita, Lorna H. & Kaneshiro, Kenneth Y. (1979), "Ethological Isolation Between Two Stocks of Drosophila Adiastola Hardy", Hawaiian Entomological Society, 23 (1): 31–34`{{citation}}`: CS1 maint: multiple names: authors list (link)
64. ^ Ahearn, J. N. (1980), "Evolution of behavioral reproductive isolation in a laboratory stock of Drosophila silvestris", Experientia, 36 (1): 63–64, doi:10.1007/BF02003975, S2CID 43809774
65. ^ Soans, A. Benedict; Pimentel, David; Soans, Joyce S. (1974), "Evolution of Reproductive Isolation in Allopatric and Sympatric Populations", The American Naturalist, 108 (959): 117–124, Bibcode:1974ANat..108..117S, doi:10.1086/282889, S2CID 84913547
66. ^ Hurd, L. E. & Eisenberg, Robert M. (1975), "Divergent Selection for Geotactic Response and Evolution of Reproductive Isolation in Sympatric and Allopatric Populations of Houseflies", The American Naturalist, 109 (967): 353–358, Bibcode:1975ANat..109..353H, doi:10.1086/283002, S2CID 85084378`{{citation}}`: CS1 maint: multiple names: authors list (link)
67. ^ Meffert, L. M. & Bryant, E. H. (1991), "Mating propensity and courtship behavior in serially bottlenecked lines of the housefly", Evolution, 45 (2): 293–306, doi:10.1111/j.1558-5646.1991.tb04404.x, PMID 28567864, S2CID 13379387`{{citation}}`: CS1 maint: multiple names: authors list (link)
68. ^ Regan, J. L.; Meffert, L. M.; Bryant, E. H. (2003), "A direct experimental test of founder-flush effects on the evolutionary potential for assortative mating", Journal of Evolutionary Biology, 16 (2): 302–312, doi:10.1046/j.1420-9101.2003.00521.x, PMID 14635869
69. ^ Miyatake, Takahisa & Shimizu, Toru (1999), "Genetic correlations between life-history and behavioral traits can cause reproductive isolation", Evolution, 53 (1): 201–208, doi:10.2307/2640932, JSTOR 2640932, PMID 28565193`{{citation}}`: CS1 maint: multiple names: authors list (link)
70. ^ Paterniani, E. (1969), "Selection for Reproductive Isolation between Two Populations of Maize, Zea mays L.", Evolution, 23 (4): 534–547, doi:10.2307/2406851, JSTOR 2406851, PMID 28562870
71. ^ Ödeen, Anders & Florin, Ann-Britt (2002), "Sexual selection and peripatric speciation: the Kaneshiro model revisited", Journal of Evolutionary Biology, 15 (2): 301–306, doi:10.1046/j.1420-9101.2002.00378.x`{{citation}}`: CS1 maint: multiple names: authors list (link)
72. ^ Leu, J. Y. & Murray, A. W. (2006), "Experimental evolution of mating discrimination in budding yeast", Current Biology, 16 (3): 280–286, Bibcode:2006CBio...16..280L, doi:10.1016/j.cub.2005.12.028, PMID 16461281`{{citation}}`: CS1 maint: multiple names: authors list (link)
73. ^ Harper, A. A. & Lambert, D. M. (1983), "The population genetics of reinforcing selection", Genetica, 62 (1): 15–23, doi:10.1007/BF00123305, S2CID 7947934`{{citation}}`: CS1 maint: multiple names: authors list (link)
74. ^ Halliburton, Richard & Gall, G. A. E. (1981), "Disruptive selection and assortative mating in Tribolium castaneum", Evolution, 35 (5): 829–843, doi:10.1111/j.1558-5646.1981.tb04947.x, PMID 28581046`{{citation}}`: CS1 maint: multiple names: authors list (link)
75. ^ Lofdahl, L. Katharine; Hu, Dan; Ehrman, Lee; Hirsch, Jerry; Skoog, Linda (1992), "Incipient reproductive isolation and evolution in laboratory Drosophila melanogaster selected for geotaxis", Animal Behaviour, 44 (4): 783–786, doi:10.1016/S0003-3472(05)80307-0, S2CID 53257556
76. ^ Moya, A.; Galiana, A.; Ayala, F. J. (1995), "Founder-effect speciation theory: failure of experimental corroboration", Proceedings of the National Academy of Sciences, 92 (9): 3983–3986, Bibcode:1995PNAS...92.3983M, doi:10.1073/pnas.92.9.3983, PMC 42086, PMID 7732017
77. ^ Dettman, Jeremy R.; Anderson, James B.; Kohn, Linda M. (2008), "Divergent adaptation promotes reproductive isolation among experimental populations of the filamentous fungus Neurospora", BMC Evolutionary Biology, 8 (35): 35, Bibcode:2008BMCEE...8...35D, doi:10.1186/1471-2148-8-35, PMC 2270261, PMID 18237415
78. ^ Dettman, Jeremy R.; Sirjusingh, Caroline; Kohn, Linda M.; Anderson, James B. (2007), "Incipient speciation by divergent adaptation and antagonistic epistasis in yeast", Nature, 447 (7144): 585–588, Bibcode:2007Natur.447..585D, doi:10.1038/nature05856, PMID 17538619, S2CID 4382609
79. ^ Martin, Oliver Y. & Hosken, David J. (2003), "The evolution of reproductive isolation through sexual conflict", Nature, 423 (6943): 979–982, Bibcode:2003Natur.423..979M, doi:10.1038/nature01752, PMID 12827200, S2CID 4379725`{{citation}}`: CS1 maint: multiple names: authors list (link)
80. ^ Wigby, S. & Chapman, T. (2006), "No evidence that experimental manipulation of sexual conflict drives premating reproductive isolation in Drosophila melanogaster", Journal of Evolutionary Biology, 19 (4): 1033–1039, doi:10.1111/j.1420-9101.2006.01107.x, PMID 16780504`{{citation}}`: CS1 maint: multiple names: authors list (link)
81. ^ Bacigalupe, L. D.; Crudgington, H. S.; Hunter, F.; Moore, A. J.; Snook, R. R. (2007), "Sexual conflict does not drive reproductive isolation in experimental populations of Drosophila pseudoobscura", Journal of Evolutionary Biology, 20 (5): 1763–1771, doi:10.1111/j.1420-9101.2007.01389.x, PMID 17714294
82. ^ Rundle, Howard D.; Chenoweth, Steve F.; Doughty, Paul; Blows, Mark W. (2005), "Divergent selection and the evolution of signal traits and mating preferences", PLOS Biology, 3 (11): e368, doi:10.1371/journal.pbio.0030368, PMC 1262626, PMID 16231971
83. ^ Higgie, Megan; Chenoweth, Steve F.; Blows, Mark W. (2000), "Natural selection and the reinforcement of mate recognition" (PDF), Science, 290 (5491): 519–521, Bibcode:2000Sci...290..519H, doi:10.1126/science.290.5491.519, PMID 11039933
84. ^ Meyer, Justin R.; Dobias, Devin T.; Medina, Sarah J.; Servilio, Lisa; Gupta, Animesh; Lenski, Richard E. (2016), "Ecological speciation of bacteriophage lambda in allopatry and sympatry", Science, 354 (6317): 1301–1304, Bibcode:2016Sci...354.1301M, doi:10.1126/science.aai8446, PMID 27884940
85. ^ Overmeer, W. P. J. (1966), "Intersterility as a Consequence of Insecticide Selections in Tetranychus urticae Koch (Acari: Tetranychidae)", Nature, 209 (321): 321, Bibcode:1966Natur.209..321O, doi:10.1038/209321a0, PMID 5950361, S2CID 4190896
86. ^ Fry, James D. (1999), "The role of adaptation to host plants in the evolution of reproductive isolation: Negative evidence from Tetranychus urticae Koch", Experimental & Applied Acarology, 23 (5): 379–387, doi:10.1023/A:1006245711950, hdl:2027.42/41783, S2CID 2329637
87. ^ Rieseberg, L. H.; Sinervo B.; Linder, C. R.; Ungerer, M.C.; Arias, D. M. (1996), "Role of Gene Interactions in Hybrid Speciation: Evidence from Ancient and Experimental Hybrids", Science, 272 (5262): 741–745, Bibcode:1996Sci...272..741R, doi:10.1126/science.272.5262.741, PMID 8662570, S2CID 39005242
88. ^ Greig, Duncan; Louis, Edward J.; Borts, Rhona H.; Travisano, Michael (2002), "Hybrid speciation in experimental populations of yeast", Science, 298 (5599): 1773–1775, Bibcode:2002Sci...298.1773G, doi:10.1126/science.1076374, PMID 12459586, S2CID 29972396
89. ^ Ghosh, Shampa M. & Joshi, Amitabh (2012), "Evolution of reproductive isolation as a by-product of divergent life-history evolution in laboratory populations of Drosophila melanogaster", Ecology and Evolution, 2 (12): 3214–3226, Bibcode:2012EcoEv...2.3214M, doi:10.1002/ece3.413, PMC 3539013, PMID 23301185`{{citation}}`: CS1 maint: multiple names: authors list (link)
90. ^ Bárbaro, Margarida; Mira, Mário S.; Fragata, Inês; Simões, Pedro; Lima, Margarida; Lopes-Cunha, Miguel; Kellen, Bárbara; Santos, Josiane; Varela, Susana A. M.; Matos, Margarida; Magalhães, Sara (2015), "Evolution of mating behavior between two populations adapting to common environmental conditions", Ecology and Evolution, 5 (8): 1609–1617, Bibcode:2015EcoEv...5.1609B, doi:10.1002/ece3.1454, PMC 4409410, PMID 25937905
91. ^ Anderson, Carlos J. R. & Harmon, Luke (2014), "Ecological and Mutation-Order Speciation in Digital Organisms", The American Naturalist, 183 (2): 257–269, Bibcode:2014ANat..183..257A, doi:10.1086/674359, PMID 24464199, S2CID 30400444`{{citation}}`: CS1 maint: multiple names: authors list (link)
92. ^ Seike, Taisuke; Nakamura, Taro; Shimoda, Chikashi (2015), "Molecular coevolution of a sex pheromone and its receptor triggers reproductive isolation in Schizosaccharomyces pombe", PNAS, 112 (14): 4405–4410, Bibcode:2015PNAS..112.4405S, doi:10.1073/pnas.1501661112, PMC 4394278, PMID 25831518
93. ^ Debelle, Allan; Ritchie, Michael G.; Snook, Rhonda R. (2014), "Evolution of divergent female mating preference in response to experimental sexual selection", Evolution, 68 (9): 2524–2533, doi:10.1111/evo.12473, PMC 4262321, PMID 24931497
94. ^ Fricke, C; Andersson, C.; Arnqvist, G. (2010), "Natural selection hampers divergence of reproductive traits in a seed beetle", Journal of Evolutionary Biology, 23 (9): 1857–1867, doi:10.1111/j.1420-9101.2010.02050.x, PMID 20646133, S2CID 13815274

* v
* t
* e

Speciation

* Introduction
* History
* Laboratory experiments
* Glossary

Basic concepts|

* Species (Species complex)
* Reproductive isolation
* Anagenesis
* Cladogenesis
* Cospeciation
* Evidence of evolution

Geographic modes|

* Allopatric (Peripatric · Quantum · Centrifugal · Founder effect)
* Parapatric (Clines · Ring species)
* Sympatric

Isolating factors|

* Adaptation
* Natural selection
* Sexual selection
* Ecological speciation (Parallel speciation · Allochrony)
* Nonecological speciation
* Assortative mating
* Haldane's rule

Hybrid concepts|

* Hybrid speciation (Polyploidy · Klepton · Recombination)
* Reinforcement (evidence)
* Secondary contact
* Character displacement

Speciation in taxa|

* Birds
* Fish
* Insects
* Plants
* Fossils (Paleopolyploidy · Punctuated equilibrium · Macroevolution · Chronospecies)

* Category
* Commons
* Evolutionary biology Portal
* WikiProject

* v
* t
* e

Evolutionary biology

* Introduction
* Outline
* Timeline of evolution
* History of life
* Index

Evolution|

* Abiogenesis
* Adaptation
* Adaptive radiation
* Altruism
* Cheating
* Reciprocal
* Baldwin effect
* Cladistics
* Coevolution
* Mutualism
* Common descent
* Convergence
* Divergence
* Earliest known life forms
* Evidence of evolution
* Evolutionary arms race
* Evolutionary pressure
* Exaptation
* Extinction
* Event
* Homology
* Last universal common ancestor
* Macroevolution
* Microevolution
* Mismatch
* Non-adaptive radiation
* Origin of life
* Panspermia
* Parallel evolution
* Signalling theory
* Handicap principle
* Speciation
* Species
* Species complex
* Taxonomy
* Unit of selection
* Gene-centered view of evolution

Population
genetics|

* Artificial selection
* Biodiversity
* Evolutionarily stable strategy
* Fisher's principle
* Fitness
* Inclusive
* Gene flow
* Genetic drift
* Kin selection
* Inbreeding avoidance
* Kin recognition
* Parental investment
* Parent–offspring conflict
* Mutation
* Population
* Natural selection
* Sexual dimorphism
* Sexual selection
* Flowering plants
* Fungi
* Mate choice
* Social selection
* Trivers–Willard hypothesis
* Variation

Development|

* Canalisation
* Evolutionary developmental biology
* Genetic assimilation
* Inversion
* Modularity
* Phenotypic plasticity

Of taxa|

* Bacteria
* Birds
* origin
* Brachiopods
* Molluscs
* Cephalopods
* Dinosaurs
* Fish
* Fungi
* Insects
* butterflies
* Life
* Mammals
* cats
* canids
* wolves
* dogs
* hyenas
* dolphins and whales
* horses
* Kangaroos
* primates
* humans
* lemurs
* sea cows
* Plants
* pollinator-mediated
* Reptiles
* Spiders
* Tetrapods
* Viruses

Of organs|

* Cell
* DNA
* Flagella
* Eukaryotes
* symbiogenesis
* chromosome
* endomembrane system
* mitochondria
* nucleus
* plastids
* In animals
* eye
* hair
* auditory ossicle
* nervous system
* brain

Of processes|

* Aging
* Death
* Programmed cell death
* Avian flight
* Biological complexity
* Cooperation
* Color vision
* in primates
* Emotion
* Empathy
* Ethics
* Eusociality
* Immune system
* Metabolism
* Monogamy
* Morality
* Mosaic evolution
* Multicellularity
* Sexual reproduction
* Gamete differentiation/sexes
* Life cycles/nuclear phases
* Mating types
* Meiosis
* Sex-determination
* Snake venom

Tempo and modes|

* Gradualism/Punctuated equilibrium/Saltationism
* Micromutation/Macromutation
* Uniformitarianism/Catastrophism

Speciation|

* Allopatric
* Anagenesis
* Catagenesis
* Cladogenesis
* Cospeciation
* Ecological
* Hybrid
* Non-ecological
* Parapatric
* Peripatric
* Reinforcement
* Sympatric

History|

* Renaissance and Enlightenment
* Transmutation of species
* David Hume
* Dialogues Concerning Natural Religion
* Charles Darwin
* On the Origin of Species
* History of paleontology
* Transitional fossil
* Blending inheritance
* Mendelian inheritance
* The eclipse of Darwinism
* Neo-Darwinism
* Modern synthesis
* History of molecular evolution
* Extended evolutionary synthesis

Philosophy|

* Darwinism
* Alternatives
* Catastrophism
* Lamarckism
* Orthogenesis
* Mutationism
* Saltationism
* Structuralism
* Spandrel
* Theistic
* Vitalism
* Teleology in biology

Related|

* Biogeography
* Ecological genetics
* Evolutionary medicine
* Group selection
* Cultural evolution
* Cultural group selection
* Dual inheritance theory
* Hologenome theory of evolution
* Missing heritability problem
* Molecular evolution
* Astrobiology
* Phylogenetics
* Tree
* Polymorphism
* Protocell
* Systematics
* Transgenerational epigenetic inheritance

* Category
* Portal

*[v]: View this template
*[t]: Discuss this template
*[e]: Edit this template