Laboratory experiments of speciation

Short description: 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

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 |  |  |  |  |  |  | 2002
D. melanogaster | Life history |  |  |  |  |  | Ghosh & Joshi[88] | 2012
Drosophila subobscura | Mate behavior  |  |  |  |  |  | Bárbaro et al.[89] | 2015
Digital organisms  |  | ~42,000; ~850 (20)  | Ecological  |  |  | Post-zygotic  | Anderson & Harmon[90] | 2014
Schizosaccharomyces pombe |  |  |  |  |  | Complete reproductive isolation  | Seike et al.[91] | 2015
D. pseudoobscura | Courtship song  | 130  |  |  |  |  | Debelle et al.[92] | 2014
Callosobruchus maculatus |  | 40 (16)  |  |  |  |  | Debelle et al.[93] | 2010

## See also

* Directed evolution

## References

1. ↑ 1.0 1.1 1.2 Coyne, Jerry A.; Orr, H. Allen (2004), Speciation, Sinauer Associates, pp. 1–545, ISBN 978-0-87893-091-3
2. ↑ 2.0 2.1 2.2 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
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", PNAS 114 (23): 6074–6079, doi:10.1073/pnas.1615109114, PMID 28533412, PMC 5468626, Bibcode: 2017PNAS..114.6074M, http://www.pnas.org/content/pnas/114/23/6074.full.pdf
5. ↑ 5.0 5.1 Ringo, John; Wood, David; Rockwell, Robert; Dowse, Harold (1985), "An Experiment Testing Two Hypotheses of Speciation", The American Naturalist 126 (5): 642–661, doi:10.1086/284445
6. ↑ 6.0 6.1 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
7. ↑ 7.0 7.1 Kirkpatrick, Mark & Ravigné, Virginie (2002), "Speciation by Natural and Sexual Selection: Models and Experiments", The American Naturalist 159: S22–S35, doi:10.1086/338370, PMID 18707367
8. ↑ 8.0 8.1 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, PMID 17248452
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, PMID 28563991
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
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, PMID 4217303
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
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
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
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
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
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, PMID 17248436
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, 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, doi:10.1126/science.213.4514.1405, PMID 17732575, Bibcode: 1981Sci...213.1405M
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
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, doi:10.1086/303186, PMID 29586617
28. ↑ Thoday, J. M. & Gibson, J. B. (1962), "Isolation by disruptive selection", Nature 193 (4821): 1164–1166, doi:10.1038/1931164a0, PMID 13920720, Bibcode: 1962Natur.193.1164T
29. ↑ Thoday, J. M. & Gibson, J. B. (1970), "The probability of isolation by disruptive selection", Nature 104 (937): 219–230, doi:10.1086/282656
30. ↑ Scharloo, W. (1971), "Reproductive isolation by disruptive selection: Did it occur?", American Naturalist 105 (941): 83–86, doi:10.1086/282706
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, PMID 28555983
32. ↑ Ehrman, Lee (1971), "Natural selection and the origin of reproductive isolation", American Naturalist 105 (945): 479–483, doi:10.1086/282739
33. ↑ Ehrman, Lee (1973), "More on natural selection and the origin of reproductive isolation", American Naturalist 107 (954): 318–319, doi:10.1086/282835
34. ↑ Ehrman, Lee (1979), "Still more on natural selection and the origin of reproductive isolation", American Naturalist 113 (1): 148–150, doi:10.1086/283371
35. ↑ Ehrman, Lee (1983), "Fourth report on natural selection for the origin of reproductive isolation", American Naturalist 121 (2): 290–293, doi:10.1086/284059
36. ↑ Chabora, Alice J. (1968), "Disruptive selection for sternopleural chaeta number in various strains of Drosophila melanogaster", American Naturalist 102 (928): 525–532, doi:10.1086/282565
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, PMID 6061662
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, PMID 17248572
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
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, PMID 28563894
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
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, 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, doi:10.1073/pnas.55.4.727, PMID 5219677, Bibcode: 1966PNAS...55..727D
47. ↑ de Oliveira, Alice Kalisz & Cordeiro, Antonio Rodrigues (1980), "Adaptation of Drosophila willistoni experimental populations to extreme pH medium", Heredity 44: 123–130, doi:10.1038/hdy.1980.11
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, 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
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
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, PMID 5229969
55. ↑ Powell, Jeffrey R. (1978), "The Founder-Flush Speciation Theory: An Experimental Approach", Evolution 32 (3): 465–474, doi:10.2307/2407714, 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
57. ↑ Galiana, Augustí, Moya, Andres & Ayala, Fransisco 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
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
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, 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, 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
63. ↑ Arita, Lorna H. & Kaneshiro, Kenneth Y. (1979), "Ethological Isolation Between Two Stocks of Drosophila Adiastola Hardy", Hawaiian Entomological Society 23 (1): 31–34
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
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, doi:10.1086/282889
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, doi:10.1086/283002
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
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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, PMID 28565193
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, 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
72. ↑ Leu, J. Y. & Murray, A. W. (2006), "Experimental evolution of mating discrimination in budding yeast", Current Biology 16 (3): 280–286, doi:10.1016/j.cub.2005.12.028, PMID 16461281
73. ↑ Harper, A. A. & Lambert, D. M. (1983), "The population genetics of reinforcing selection", Genetica 62 (1): 15–23, doi:10.1007/BF00123305
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
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
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, doi:10.1073/pnas.92.9.3983, PMID 7732017, Bibcode: 1995PNAS...92.3983M
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, doi:10.1186/1471-2148-8-35, 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, doi:10.1038/nature05856, PMID 17538619, Bibcode: 2007Natur.447..585D
79. ↑ Martin, Oliver Y. & Hosken, David J. (2003), "The evolution of reproductive isolation through sexual conflict", Nature 423 (6943): 979–982, doi:10.1038/nature01752, PMID 12827200, Bibcode: 2003Natur.423..979M
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
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, PMID 16231971
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Speciation

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* Species (Species problem · Species complex)
* Reproductive isolation
* Anagenesis
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