Adaptive Dynamics in a 2-patch Environment: a Simple Model for Allopatric and Parapatric Speciation

Adaptation to an environment consisting of two patches (each with different optimal strategy) is investigated. The patches have independent density regulation ('soft selection'). If the patches are similar enough and migration between them is strong, then evolution ends up with a generalist ESS. If either the difference between the patches increases or migration weakens, then the generalist strategy represent a branching singularity: The initially monomorphic population first evolves towards the generalist strategy, there it undergoes branching, and finally two specialist strategies form an evolutionarily stable coalition. Further increasing the between-patch difference or decreasing migration causes the generalist to lose its convergence stability as well, and an initially monomorphic population evolves towards one of the specialists optimally adapted to one of the two patches. Bifurcation pattern of the singularities is presented as a function of patch difference and migration rate. Connection to speciation theory is discussed. The transition from the generalist ESS to the coexisting pair of specialist strategies is regarded as a clonal prototype of parapatric (if the between-patch difference increases) or allopatric (if the migration decreases) speciation. We conclude that the geographic and the competitive speciation modes are not distinct classes.

[1]  Karel F. Liem,et al.  Evolutionary Strategies and Morphological Innovations: Cichlid Pharyngeal Jaws , 1973 .

[2]  D. Krakauer,et al.  SEXUAL SELECTION, SPACE, AND SPECIATION , 1997, Evolution; international journal of organic evolution.

[3]  J. Metz,et al.  Adaptive Dynamics: A Geometrical Study of the Consequences of Nearly Faithful Reproduction , 1995 .

[4]  Peter D. Taylor,et al.  Evolutionary stability in one-parameter models under weak selection , 1989 .

[5]  Daniel R. Brooks Evolution: A biological and Paleontological approach: edited by Peter Skelton Addison Wesley, 1993. £19.95 pbk (1064 pages) ISBN 0 201 54423 7 , 1994 .

[6]  Marino Gatto,et al.  Lyapunov exponents and the mathematics of invasion in oscillatory or chaotic populations , 1995 .

[7]  C. Woese,et al.  Bacterial evolution , 1987, Microbiological reviews.

[8]  O. Seehausen,et al.  Cichlid Fish Diversity Threatened by Eutrophication That Curbs Sexual Selection , 1997 .

[9]  G. Fox,et al.  Chaos and evolution. , 1995, Trends in ecology & evolution.

[10]  F. Galis The application of functional morphology to evolutionary studies. , 1996, Trends in ecology & evolution.

[11]  D. Tautz,et al.  Sympatric speciation suggested by monophyly of crater lake cichlids , 1994, Nature.

[12]  Odo Diekmann,et al.  On evolutionarily stable life histories, optimization and the need to be specific about density dependence , 1995 .

[13]  F. B. Christiansen,et al.  Evolution and intraspecific exploitative competition I. One-locus theory for small additive gene effects , 1980 .

[14]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[15]  P. Hammerstein Darwinian adaptation, population genetics and the streetcar theory of evolution , 1996, Journal of mathematical biology.

[16]  R. Lande,et al.  RAPID ORIGIN OF SEXUAL ISOLATION AND CHARACTER DIVERGENCE IN A CLINE , 1982, Evolution; international journal of organic evolution.

[17]  Odo Diekmann The Many Facets of Evolutionary Dynamics , 1995 .

[18]  O. Seehausen Lake Victoria rock cichlids : taxonomy, ecology, and distribution , 1996 .

[19]  U Dieckmann,et al.  Evolutionary Cycling of Predator-Prey Interactions : Population Dynamics and the Red Queen , 1999 .

[20]  Mark V. Lomolino,et al.  Species Diversity in Space and Time. , 1996 .

[21]  G. Turner,et al.  A model of sympatric speciation by sexual selection , 1995, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[22]  E. Pasztor Unexploited Dimensions of Optimization Life History Theory , 1988 .

[23]  J. Lythgoe,et al.  Visual pigment polymorphism in the guppy poecilia reticulata , 1987, Vision Research.

[24]  Brian Charlesworth,et al.  Evolution in Age-Structured Populations. , 1983 .

[25]  T. Vincent,et al.  A theory for the evolutionary game , 1987 .

[26]  I. Eshel Evolutionary and continuous stability , 1983 .

[27]  A. Meyer,et al.  Monophyletic origin of Lake Victoria cichlid fishes suggested by mitochondrial DNA sequences , 1990, Nature.

[28]  R. Nisbet,et al.  How should we define 'fitness' for general ecological scenarios? , 1992, Trends in ecology & evolution.

[29]  G. Vermeij Adaptation, Versatility, and Evolution , 1973 .

[30]  J. M. Smith,et al.  The Logic of Animal Conflict , 1973, Nature.

[31]  F. B. Christiansen Hard and Soft Selection in a Subdivided Population , 1975, The American Naturalist.

[32]  L. Kaufman,et al.  Evolution in fast forward: haplochromine fishes of the Lake Victoria region , 1997 .

[33]  É. Kisdi,et al.  Dynamics of Adaptation and Evolutionary Branching , 1997 .

[34]  S. M. Verduyn Lunel,et al.  Stochastic and spatial structures of dynamical systems , 1996 .

[35]  D. E. Matthews Evolution and the Theory of Games , 1977 .

[36]  I. Eshel On the changing concept of evolutionary population stability as a reflection of a changing point of view in the quantitative theory of evolution , 1996, Journal of mathematical biology.

[37]  S. Wright Evolution in mendelian populations , 1931 .

[38]  Odo Diekmann,et al.  When Does Evolution Optimize? On the Relation Between Types of Density Dependence and Evolutionarily Stable Life History Parameters , 1996 .

[39]  R. D. Ricketts,et al.  Late Pleistocene Desiccation of Lake Victoria and Rapid Evolution of Cichlid Fishes , 1996, Science.

[40]  K. Liem Adaptive Significance of Intra- and Interspecific Differences in the Feeding Repertoires of Cichlid Fishes , 1980 .

[41]  W. Getz,et al.  Population dynamics and harvesting of semelparous species with phenotypic and genotypic variability in reproductive age , 1995 .

[42]  Éva Kisdi,et al.  Density dependent life history evolution in fluctuating environments , 1993 .

[43]  R. Macarthur,et al.  COMPETITION, HABITAT SELECTION, AND CHARACTER DISPLACEMENT IN A PATCHY ENVIRONMENT. , 1964, Proceedings of the National Academy of Sciences of the United States of America.

[44]  Thomas L. Vincent,et al.  Evolution via Strategy Dynamics , 1993 .

[45]  John Maynard Smith,et al.  Organizational Constraints on the Dynamics of Evolution , 1990 .

[46]  Richard E. Michod,et al.  Evolution of Life Histories in Response to Age-Specific Mortality Factors , 1979, The American Naturalist.