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The Community Context of Species' Borders: Ecological And OIKOS 108: 28Á/46, 2005 The community context of species’ borders: ecological and evolutionary perspectives Ted J. Case, Robert D. Holt, Mark A. McPeek and Timothy H. Keitt Case, T. J., Holt, R. D., McPeek, M. A. and Keitt, T. H. 2005. The community context of species’ borders: ecological and evolutionary perspectives. Á/ Oikos 108: 28Á/46. Species distributional limits may coincide with hard dispersal barriers or physiological thresholds along environmental gradients, but they may also be influenced by species interactions. We explore a number of models of interspecific interactions that lead to (sometimes abrupt) distribution limits in the presence and absence of environmental gradients. We find that gradients in competitive ability can lead to spatial segregation of competitors into distinct ranges, but that spatial movement tends to broaden the region of sympatry between the two species, and that Allee effects tend to sharpen these boundaries. We generalize these simple models to include metapopulation dynamics and other types of interactions including predatorÁ/prey and host-parasite interactions. We derive conditions for range limits in each case. We also consider models that include coevolution and gene flow and find that character displacement along environmental gradients can lead to stable parapatric distributions. We conclude that it is essential to consider coevolved species interactions as a potential mechanism limiting species distributions, particularly when barriers to dispersal are weak and environmental gradients are gradual. T. J. Case, Dept of Biology, Univ. of California at San Diego, La Jolla, CA 92093-0116, USA ([email protected]). Á/ R. D. Holt, Dept of Zoology, Univ. of Florida, Gaines- ville, FL 32611, USA. Á/ M. A. McPeek, Dept of Biological Sciences, Dartmouth College, Hanover, NH 03755, USA. Á/ T. H. Keitt, Integrative Biology, Univ. of Texas, Austin, TX 78712, USA. A common observation about life is that species have fact that South Africa has a fauna that is different from, geographical distributions that are more or less spatially say, Argentina, depends on the important role of barriers confined. Biogeography attempts to make sense of these in preventing colonization of species. The common distributional patterns by studying both present day observation that introduced species can sometimes thrive environmental and ecological factors determining fitness in foreign terrain, once transported there, indicates that and populational persistence, and the historical origins present day distributions are not determined solely by and subsequent movements of individuals. One obvious environmental conditions, but that barriers to dispersal, feature that prevents species from spreading uniformly both past and present, are responsible for some degree of over space is the existence of physical barriers to the regional and global patterning of the biota. dispersal. The association between broad scale biogeo- Over time, however, even hard barriers to dispersal graphic provinces and major barriers to dispersal, (e.g. may be breached. Oceanic volcanic islands (e.g. Hawaii, oceans for terrestrial forms, or strong cross currents for the Galapagos, or the Canary Islands) that have never ocean forms with pelagic larval dispersal, Gaylord and been connected to another region are eventually colo- Gaines 2000) is a persistent reminder of the role of nized by a wide range of terrestrial species. Furthermore, barriers in determining community composition. The given the ability of populations to adapt to local Accepted 8 June 2004 Copyright # OIKOS 2005 ISSN 0030-1299 28 OIKOS 108:1 (2005) conditions by natural selection, it is not immediately scape and the transport of organisms allows unique obvious why some hard barriers to dispersal could not mixing of faunas that may not have existed even just a eventually be breached. A mountain range may form a century ago. For instance, Nowak (1971) cataloged 28 barrier to dispersal for a lowland species. However, examples of recent range expansions of plants and populations at elevations at the base of the mountain animals primarily in Europe. In a majority of cases, may adapt to survive there. What prevents a sequence of the range expansion could be attributed to some human- such episodes of local adaptation from eventually caused alteration of the landscape creating new habitat allowing the populations to march across the mountain? or avenues for dispersal. Finally, habitat loss through One explanation is that there may not be sufficient human alteration has caused many species’ ranges to genetic variation in the population for natural selection, retract to selected corners of their former range where given the intensity of selection. As shown by Holt and human impacts have been less (Lomolino and Channell Gomulkiewicz (1997), adaptation in marginal popula- 1995). A study of these present day ranges, in the absence tions requires the availability of genetic variation simply of historical information about their former extents for population persistence in the absence of immigration. would provide a misleading and biased impression of Sometimes such variation will be absent (Bradshaw the role of climate, interspecific interactions, and evolu- 1991). It is hard to imagine that sufficient genetic tionary dynamics in determining range limits. variation exists to allow a mouse to evolve to both live in the ocean and then to colonize the next continent on the other side. A barrier to dispersal is one that must be defined in terms of the existing genetic variability of the Nonevolutionary models focal species, the severity of the selection gradient over space, and the time scale in question. Two competitors Another reason that local adaptation may fail to Range limits with an environmental gradient advance the spatial spread of a population is because it is Interspecific competition has the capacity to produce opposed by net gene flow from a more populous abrupt range limits in either homogeneous or hetero- distributional center where different genotypes may be geneous space, although by different scenarios. Rough- favored. Theoretical studies suggest that this antagonism garden (1979) was one of the first to model range limits between local adaptation and the detrimental effects of in the context of the LotkaÁ/Volterra competition gene flow can produce a stable limited geographic range equations. The carrying capacity of the two competitors even in the absence of any barriers to dispersal (Kirkpa- was assumed to vary inversely over one-dimensional trick and Barton 1997, Case and Taper 2000, Antonovics space x; competitor A’s carrying capacity KA(x) mono- et al. 2001, but see Barton 2001 and Gomulkiewicz et al. tonically declines going from left to right, while compe- 1999 for analyses of the countervailing positive effect of titor B’s carrying capacity monotonically increases over gene flow upon local adaptation via the infusion of the same range. In the absence of the other competitor, genetic variation). If the environment varies smoothly each species’ equilibrium density over space would and gradually over space, and dispersal coefficients are follow its K curve. Movements were not explicitly not too high (which would cause high gene flow), and considered and interspecific competition coefficients (a) given adequate genetic variation, these models predict were assumed to be constants independent of spatial that a species range could eventually expand across position or population density. Even if interspecific space. As we discuss further below, however, the capacity competition is less than intraspecific competition, one for unlimited spread is greatly diminished in the presence species can competitively exclude its competitor in some of interspecific interactions. portion of space where it has a sufficiently higher In this paper, we review and synthesize the ways that carrying capacity, giving rise to more or less sharp range biotic interactions may produce stable range limits. We limits depending on the magnitude of the interspecific focus on situations where the species boundary is stable effects and the steepness of the K gradients. or quasi-stable but not associated with obvious barriers As a simple example, assume that two LotkaÁ/Volterra to dispersal. This restriction on ‘‘stable’’ is important competitors have equal competition coefficients and that because the geographic ranges of species with low their carrying capacities vary in opposite directions movement rates and slow population growth may not along a gradient. Coexistence at any point along the yet be at equilibrium with respect to past climate change gradient requires that 1/a/K1(x)/ K2(x)/a, where a is and continuing habitat alteration. Fossil records over the the competition coefficient, and Ki(x) is the carrying late Pleistocene and Holocene for tree and mammal capacity of competitor i at gradient position x. species show slow spatial dynamics as they respond to For illustrative purposes, assume that a5/1, and that glacial cycles (Davis 1986, Graham 1986, Woods and the two competitors are specialized to opposite ends Davis 1989, Gear and Huntley 1991, Martin and Fair- of the gradient, with linear but opposite relations banks 1999). Recent human modifications to the land- of spatial position to carrying capacity, as follows: OIKOS 108:1 (2005) 29 2 K (x)/K?/gx, and K (x)/gx, for 05/x5/K?/g. The 1N (x; t) 1 N (x; t) r N (x; t) 1 2 i D i i i gradient has length K?/g, and in the absence of i 2 1t 1x Ki(x) competition each species can occupy the entire gradient. Â [K (x)N (x; t)a N (x; t)] d N (x; t) If both species are present, competition can restrict each i i ij j i i species’ range size, with a zone of overlap in the middle (2) of the gradient. After substitution into the expression for coexistence, the width W of the overlap zone is Here, Di is the diffusion coefficient and di is density- W/(K?/g) (1/a)/(1/a). Competition shrinks the range independent mortality. Compared to nonspatial models, edge of, say, species 1, from K?/g to K?/(g(1/a)).
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