8/23/2016 Niche ­ ­ Oxford Bibliographies

Niche Evolution Alex Pyron

LAST MODIFIED: 26 MAY 2016 DOI: 10.1093/OBO/9780199941728­0075

Introduction

The evolution of species’ niches is a process that is fundamental to investigations in numerous fields of biology, including , assembly, and long­term regional and global diversification processes. It forms the nexus between ecological and evolutionary questions. Topics as diverse as ecological speciation, niche conservatism, species coexistence, and historical all rely on interpreting patterns and drivers of species’ niches through time and across landscapes. Despite this importance, a distinct research agenda concerning niche evolution as a discrete topic of inquiry has yet to emerge. Niche evolution is often considered as a sidebar or of secondary importance when addressing questions such as “how did two species diverge?” Basic questions such as “what is a niche,” “what is the biological basis of niche evolution,” “at what scale should we evaluate niche evolution,” and “how can we observe niche evolution at different timescales” have rarely been addressed directly, or not at all in some systems. However, various intellectual threads connecting these ideas are evident in a number of recent and historical publications, giving some semblance of form to a framework for interpreting and evaluating niche evolution, and outlining major areas for future research from an evolutionary perspective. There is a reverse perspective from the macroecological scale as well, with questions involving coexistence, distributions and ranges, food webs, and other organismal attributes.

General Overview

Niche evolution has rarely, if ever, been addressed in depth as a standalone topic. Instead, much of the conceptual development in literature is intertwined with that of speciation, , and diversification. Mayr 1942 provided the original background material for recent developments, during the Modern Synthesis. Williams 1966 focuses on at the individual and population level in a theoretical context, providing the necessary background to interpret processes of niche evolution. Schluter 2000 describes the ecological basis of adaptive evolutionary radiations, and how niche evolution drives ecomorphological diversification. Chase and Leibold 2003 is a classic treatise on what a niche is, and how to interpret niche concepts in practice. Odling­Smee, et al. 2003 approaches niche evolution from the perspective of the individual interacting with its environment, and the process of . Coyne and Orr 2004 covers the various mechanisms that drive speciation, which includes heavy focus on ecological isolation and niche evolution. Losos 2009 is a case study of niche evolution in the classic model­system Anolis. Finally, Peterson, et al. 2011 covers a profusion of methodological approaches to understanding niche evolution.

Chase, J. M., and M. A. Leibold. 2003. Ecological niches: Linking classical and contemporary approaches. Chicago: Univ. of Chicago Press. A more in­depth look at some of the issues described in this section, including niche concepts and individual, population, and species­ level considerations both in an ecological and evolutionary context. This is a fundamental starting place for niche evolution.

Coyne, J. A., and H. A. Orr. 2004. Speciation. Sunderland, MA: Sinauer.

http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 1/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies One of the major overviews of speciation processes, and the ecological factors that can underlie lineage divergence leading to speciation. In particular, chapter 5 (“Ecological Isolation”) details niche­related processes such as , pollinator, and temporal isolation. However, these are considered in a more static context, rather than how they, themselves, evolve over time.

Losos, J. B. 2009. Lizards in an evolutionary tree. Berkeley: Univ. of California Press. For empirical case studies of adaptive niche­evolution, few vertebrate systems have been investigated as fully as West Indian Anolis lizards. Losos reviews and synthesizes available studies, with particular emphasis on climatic and ecomorphological these lizards have developed to occupy novel niche­spaces.

Mayr, E. 1942. and the origin of species. New York: Columbia Univ. Press. A landmark in evolutionary biology and the study of speciation. Of particular interest here is the focus on geographic variation in species’ attributes such as habitat preference, and how this can lead to population divergence and speciation over time. Importantly, a genetic basis of ecological characteristics, and the accumulation of related to this, is recognized as a key driver of speciation, implicitly via niche evolution. Lacking, however, is an explicit mechanistic link between these concepts.

Odling­Smee, F. J., K. N. Laland, and M. W. Feldman. 2003. Niche construction: The neglected process in evolution. Princeton, NJ: Princeton Univ. Press. As reviewed empirically in other studies cited here, the ability of species to modify their immediate environments and their overall ecological niches is an underappreciated force in the diversification of lineages. The authors review the evidence and implications of niche construction for a broad range of ecological and evolutionary questions.

Peterson, A. T., J. Soberon, R. G. Pearson, et al. 2011. Ecological niches and geographic distributions. Princeton, NJ: Princeton Univ. Press. A crucial text covering more recent developments in the quantitative analysis of species’ niches, with a particular focus on modeling and predicting species’ niches and distributions using modern computational techniques. Includes background material from many of the references listed in this section with respect to niche concepts, and how niches may change over shorter timescales. In contrast, the authors do not heavily feature a historical perspective, or much address how niche evolution through time affects present­day distributions.

Schluter, D. 2000. The ecology of . Oxford: Oxford Univ. Press. Outlines the ecological basis of rapid speciation in the face of increased ecological opportunity, leading to adaptive radiations. Includes a detailed discussion on the mechanisms of ecological speciation and the basis of niche divergence among species, which is the fundamental unit of niche evolution.

Williams, G. C. 1966. Adaptation and . Princeton, NJ: Princeton Univ. Press. A classic of post­“modern synthesis” evolutionary theory. Describes the epistemological as well as the mechanistic basis of adaptation and natural selection in populations, particularly with respect to ecological aspects. Outlines the multilevel nature of adaptation, which is important when considering niche evolution.

Textbooks

http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 2/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies As a specialized subject, niche evolution has not been the subject of full textbooks. However, the fundamental principles of niche evolution are found in diverse fields in biology, and a number of basic textbooks in ecology and evolution contain the building blocks for studying adaptive niche­evolution. Begon, et al. 2006 is a broadly introductory text on the science of ecology. Futuyma 2013 is similarly a broad, foundational text in evolutionary biology. Pianka 1999 introduced many of the historically important concepts in evolutionary ecology and the study of niche evolution. Mittelbach 2012 places niche evolution and other related processes in a community context. Briggs and Crowther 2001 introduces a paleontological perspective to evolutionary ecology, for long­term analyses of niche evolution.

Begon, M., C. R. Townsend, and J. L. Harper. 2006. Ecology: From individuals to . Malden, MA: Blackwell. A basic ecology textbook, which is a crucial starting point to understanding the ecological processes that affect individuals, populations, species, and communities. The biotic and abiotic factors that comprise ecosystems and affect existence and persistence therein are key to understanding niche evolution, as are the basic definitions of the , both fundamental and realized.

Briggs, D., and P. R. Crowther. 2001. Palaeobiology II. Malden, MA: Blackwell Science. Niche evolution is a difficult subject to place in a historical context, due to the lack of data on traits and ecological niches of extinct organisms. Nevertheless, a historical focus is an important aspect of broad­scale studies of niche evolution, and a text such as this covers processes and empirical advances from a paleontological perspective.

Futuyma, D. J. 2013. Evolution. 3d ed. Sunderland, MA: Sinauer. Just as a basic ecology textbook is needed to understand the ecological basis of the niche from an environmental and perspective, so is an evolution textbook needed. Futuyma covers topics such as biogeography, speciation, and the origin of that are all crucial for integrating niche evolution through time and space.

Mittelbach, G. G. 2012. Community ecology. Sunderland, MA: Sinauer. Building on the more general principles found in broader ecology textbooks, Mittelbach explores community ecology, attempting to understand how co­occurring species can coexist locally and form ecological communities. The interaction of species on small temporal and geographic scales forms the basis for longer­term adaptations, and thus community ecology is an important foundation for niche evolution.

Pianka, E. R. 1999. Evolutionary ecology. 6th ed. San Francisco: Benjamin Cummings. This classic text introduces a number of important macroecological principles in an evolutionary framework, including principles of niche evolution. Importantly, the sixth edition introduces the phylogenetic perspective needed to compare traits across species, using computational methods and phylogenetic trees.

Journals

Much as there are no textbooks on niche evolution per se, there are no journals covering the topic as their sole focus. Instead, recent research on niches, niche conservatism, and niche evolution can be found in a wide breadth of journals focused on ecology and evolutionary biology. American Naturalist is one of the premier outlets for macroecological research with an evolutionary context. Ecography often features empirical and theoretical studies of niche evolution. Ecology is a primary journal for original research in ecology across a variety of subfields. Ecology Letters is a shorter form, high­impact journal for outstandingly novel results in and evolution. Evolution focuses primarily on evolutionary questions, but frequently entertains ecological approaches to evolutionary theory. Global Ecology and Biogeography typically publishes papers dealing with the intersection of evolutionary and macroecological processes in a biogeographic context. Journal of Biogeography typically focuses on biogeographic analyses, but http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 3/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies includes recent, more ecologically focused inquiries, such as phylogeography. Molecular Ecology typically focuses on , speciation, and phylogeography, but commonly integrates ecological perspectives on niche evolution to answer these questions.

American Naturalist. 1867–. Broadly focused in ecology and evolutionary biology, with a great deal of niche­related research.

Ecography. 1978–. A more specialized journal concerned with geographic variation in ecology, which naturally implies a strong focus on the niche and niche evolution.

Ecology. 1920–. A mainstay in ecology, with a strong focus on ecological niches, particularly at the population level.

Ecology Letters. 1998–. A high­impact journal for cutting­edge research at the nexus of ecology and evolution.

Evolution. 1947–. A mainstay in evolutionary biology, but with many papers integrating macroecological perspectives, including niche evolution.

Global Ecology and Biogeography. 1998–. As the name implies, most papers integrate historical biogeography and macroecology, leading to a natural focus on niche evolution in many studies.

Journal of Biogeography. 1996–. The field of biogeography implicitly or explicitly relies on the niche and niche evolution for the majority of its research program.

Molecular Ecology. 1992–. A strong focus on phylogeography and speciation, which includes macroecological perspectives on niche evolution for a significant proportion of papers.

Niche Concepts

A fundamental problem in studying niche evolution is simply definition of terms. What do we mean by “niche”? For that matter, what do we mean by “evolution”? How do we measure niches? Which variables do we include? Are we speaking of abiotic or biotic factors, or a combination thereof? Elith and Leathwick 2009 summarizes available methods for quantifying species’ ecological niches in a climatic framework. Hadly, et al. 2009 discusses how niches can be conserved among higher­level lineages across long evolutionary timescales. Holt 2009 addresses similar topics and outlines how the concept of the niche influences how it can be studied. Kozak, et al. 2008 http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 4/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies summarizes the use of GIS (geographic information system) data and methods to quantify the geographic and climatic space occupied by species, to characterize niches. When comparing niches among species, variable amounts of divergence and conservatism are to be expected. But how do we interpret this? Losos 2008 addresses these issues. The ways in which niche evolution can influence speciation in an ecological context are numerous, and are addressed in detail in Schluter 2001 and Rundle and Nosil 2005. Finally, geographic and ecological isolation can potentially become uncoupled during speciation, which Pyron and Burbrink 2010 outlines in a verbal model.

Elith, J., and J. R. Leathwick. 2009. models: Ecological explanation and prediction across space and time. Annual Review of Ecology Evolution and Systematics 40:677–697. Elith and Leathwick review the profusion of available methods for ecological niche modeling and how these can be applied to understand species’ distributions and climatic adaptations. Such methods form a foundation of evolutionary and ecological studies of niche evolution.

Hadly, E. A., P. A. Spaeth, and C. Li. 2009. Niche conservatism above the species level. Proceedings of the National Academy of Sciences of the United States of America 106:19707–19714. The authors report on empirical evidence for the existence of niche conservatism above the species level, such as fish rarely having left their aquatic niche. Range size in mammals seems to show the signal of higher­level conservatism, and long­term constraints on adaptive niche­evolution at higher levels may have far­reaching impacts on distributions and diversity.

Holt, R. D. 2009. Bringing the Hutchinsonian niche into the 21st century: Ecological and evolutionary perspectives. Proceedings of the National Academy of Sciences of the United States of America 106:19659–19665. Holt gives an extensive and insightful review of niche concepts and methods for quantifying and comparing niches. Importantly, he discusses how niche evolution and conservatism may play out across phylogenetic scales. He offers a variety of forward­looking questions to guide future research programs in evolutionary ecology.

Kozak, K. H., C. H. Graham, and J. J. Wiens. 2008. Integrating GIS­based environmental data into evolutionary biology. Trends in Ecology & Evolution 23:141–148. A detailed review of the available techniques from geographic information systems to extract data on the climatic niches of organisms. These are prerequisite for almost any study investigating niches or adaptive niche­evolution.

Losos, J. B. 2008. Phylogenetic niche conservatism, phylogenetic signal and the relationship between phylogenetic relatedness and ecological similarity among species. Ecology Letters 11:995–1003. In this brief but influential review, Losos discusses the nature of niche conservatism and niche evolution (phylogenetic signal) within lineages. He discusses case studies, tests, and implications for future research. He concludes that the issues involved are highly dependent on the scope and scale of the question, and merit a detailed analysis, rather than blanket assumptions of process.

Pyron, R. A., and F. T. Burbrink. 2010. Hard and soft allopatry: Physically and ecologically mediated modes of geographic speciation. Journal of Biogeography 37:2005–2015. The authors present a verbal model for the interaction of niche conservatism, adaptive niche­evolution, and geographic and climatic variation to affect ecological speciation. They describe several potential outcomes for geographic versus climatic modes of speciation, and suggest how they might be tested in future studies at the landscape level.

Rundle, H. D., and P. Nosil. 2005. Ecological speciation. Ecology Letters 8:336–352. http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 5/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies In this foundational paper, the authors review the mechanistic basis for ecological speciation, including divergent ecological selection, reproductive isolation, and the genetic basis of the two. The implications for studying adaptive niche­evolution are numerous.

Schluter, D. 2001. Ecology and the origin of species. Trends in Ecology & Evolution 16:372–380. In this foundational review, Schluter outlines the evolutionary basis of ecological divergence among populations and the interplay between divergent ecological selection, adaptive niche­evolution, and speciation. This forms the basis for understanding adaptive differentiation of populations along climatic axes.

Demes, Populations, and Species

In most niche concepts, the variable of interest is an emergent property of a larger group, deriving from preferences or tolerances of individuals. Thus, diversity within species may be an important source of variation for niche evolution. Fine­scale geographic patterns may contain a great deal of information for understanding ecological adaptations. For instance, Gee 2004 shows how ecological limitations on dispersal limit genetic contact between recently diverged populations. Fisher­Reid, et al. 2012 asks whether or not species with narrower niches have higher rates of niche evolution, which does not seem to be the case. Ecological constraints may influence the evolution of life­history traits, and thus the prevalence of behavioral adaptations for coexistence, as reported in Hatchwell and Komdeur 2000. Holt, et al. 2004 describes a mathematical model showing how source­sink dynamics can be maintained by temporal variation in the adaptive landscape. Pulliam 2000 discusses how demographic factors can, in part, explain mismatches between potential and actual distributions. Rangel, et al. 2007 uses a simulation approach to show how spatial variation in conservatism and divergence among populations can determine range limits. Finally, genetic variability among populations can influence ecological responses to speciation, either through divergence as described in Schluter and Conte 2009 or by canalization as described in Wagner, et al. 1997.

Fisher­Reid, M. C., K. H. Kozak, and J. J. Wiens. 2012. How is the rate of climatic­niche evolution related to climatic­niche breadth? Evolution 66:3836–3851. Fisher­Reid and collaborators first quantify the climatic niches of a large clade, and then use phylogenetic methods to estimate rates of climatic­niche evolution. They test the intriguing hypothesis that niche breadths are related to the speed at which niches evolve. Generally, they find little relationship, and species with narrower niches do not have higher rates of niche evolution. However, certain variables do evolve more quickly in species with broader niches.

Gee, J. M. 2004. across a climatic barrier between hybridizing avian species, California and Gambel’s quail (Callipepla californica and C. gambelii). Evolution 58:1108–1121. An interesting empirical example of how climatic­niche differences between recently diverged populations can act to limit hybridization and gene flow. Morphological traits are closely linked to the climatic variables that vary across the hybrid zone, yielding a complex suite of isolating mechanisms. This divergence is almost solely ecological, as there are no major dispersal barriers.

Hatchwell, B. J., and J. Komdeur. 2000. Ecological constraints, life history traits and the evolution of cooperative breeding. Animal Behaviour 59:1079–1086. Niche evolution results from a complex interplay of processes including ecological constraints, demographics, and biogeography. Often­ overlooked aspects are behavioral processes such as individual interactions. Hatchwell and Komdeur review how life­history traits and ecological factors interact to facilitate the evolution of complex behaviors such as .

Holt, R. D., M. Barfield, and R. Gomulkiewicz. 2004. Temporal variation can facilitate niche evolution in harsh sink environments. American Naturalist 164:187–200. http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 6/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies The authors evaluate several models for niche­evolution dynamics in sink . They find that temporal variation in the adaptive landscape of the sink can facilitate its persistence and permit niche evolution in marginal populations. This may explain dynamics observed in encountering conditions outside their core range.

Pulliam, H. R. 2000. On the relationship between niche and distribution. Ecology Letters 3:349–361. Several referenced papers in this article deal with empirical mismatches between a species’ fundamental niche (and thus its potential geographic range) and its realized niche (and thus its occupied geographic range). Pulliam reviews niche concepts related to occupancy and introduces a verbal and mathematical model for how biogeography, demography, and ecology can interact to affect species’ distributions in space and time.

Rangel, T. F. L. V. B., J. A. F. Diniz­Filho, and R. K. Colwell. 2007. and evolutionary niche dynamics: A spatial pattern­oriented simulation experiment. American Naturalist 170:602–616. Using a simulation approach, the authors show that niche dynamics (conservatism and adaptive evolution) exert a strong effect on species’ ranges, explaining a large proportion of variance in observed species­richness patterns. This gives a framework for interpreting the spatial drivers of diversity in an ecological and biogeographic framework.

Schluter, D., and G. L. Conte. 2009. Genetics and ecological speciation. Proceedings of the National Academy of Sciences of the United States of America 106:9955–9962. Schluter and Conte build on previous work to illustrate the genetic basis of ecological speciation, which in many cases acts on standing genetic variation related to ecological variability. This preexisting diversity may explain the frequency and rapidity of ecological speciation in some groups.

Wagner, G. P., G. Booth, and H. C. Bagher. 1997. A population genetic theory of canalization. Evolution 51:329–347. The authors provide a review and model of canalization, which is the suppression of phenotypic variation. This can occur when traits are insensitive to mutations (genetic) or ecological (environmental) factors. This phenomenon has broad implications for studies of adaptive niche­evolution.

Niche Plasticity

Niches may be extremely variable within and among lineages, species, and populations. This may result from lability, or plasticity in the underlying mechanisms regulating the realized niche. Sometimes this variation may be indicative of real biological processes, but it may also result from methodological biases. Boucher, et al. 2014 shows via simulation that strong artifacts of niche conservatism may be driven by geographic variation in available climatic spaces. Peterson and Holt 2003 uses GIS (geographic information system) data to examine at the landscape level in incipient species. However, Anderson and Raza 2010 shows that patterns observed using this kind of analysis may be strongly influenced by the parameters of the modeling effort. Quintero and Wiens 2013 shows that climatic variation within sites usually contains most of the climatic variation across sites, and that species with wider niches may occur in more climatically variable sites. This may explain the prevalence and frequency of ecological speciation in allopatry and the rapid divergence of climatic niches among closely related species, as discussed in Knouft, et al. 2006 and Nosil, et al. 2009. Species may also influence their own niche, whereby niche dynamics may shift rapidly and unstably over time, an underappreciated evolutionary mechanism discussed in Saltz and Nuzhdin 2014. Finally, niche variation within species has strong implications for modeling and forecasting efforts under climate­change scenarios, as outlined in Jackson, et al. 2009.

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Anderson, R. P., and A. Raza. 2010. The effect of the extent of the study region on GIS models of species geographic distributions and estimates of niche evolution: Preliminary tests with montane rodents (genus Nephelomys) in Venezuela. Journal of Biogeography 37:1378–1393. Methods for species­distribution modeling often contain crucial assumptions that are overlooked or downplayed in empirical studies. Here, the authors show that the background (the total geographic space considered potentially habitable) has a strong effect on distribution models, and potentially on estimated rates of niche evolution. They suggest that distribution models be conditioned on more limited areas containing only the known, realized distribution, and that more accurate estimates can then be obtained by projecting these into other areas.

Boucher, F. C., W. Thuiller, T. J. Davies, and S. Lavergne. 2014. Neutral biogeography and the evolution of climatic niches. American Naturalist 183:573–584. Simulation results from a spatially explicit model suggest that the geographic dimension of speciation exerts a strong influence on observed patterns of “niche conservatism,” and that tests of niche evolution must consider physiological traits and spatial contexts to be relevant.

Jackson, S. T., J. L. Betancourt, R. K. Booth, and S. T. Gray. 2009. Ecology and the ratchet of events: Climate variability, niche dimensions, and species distributions. Proceedings of the National Academy of Sciences of the United States of America 106:19685–19692. The authors review a variety of mechanisms and processes related to understanding niche evolution in the face of natural and anthropogenic climate change. They outline a broad research program for future studies of climatic variability and niche evolution, particularly with respect to predictive efforts in the face of changing global climates.

Knouft, J. H., J. B. Losos, R. E. Glor, and J. J. Kolbe. 2006. Phylogenetic analysis of the evolution of the niche in lizards of the Anolis sagrei group. Ecology 87:S29–S38. Using phylogenetic and GIS data, the authors test whether niche similarity is related to phylogenetic similarity. Surprisingly (or perhaps not), they find little relationship, suggesting that rates of climatic­niche evolution can be very high. Species pairs may have conserved, divergent, or nested climatic niches. They suggest that more niche factors should be examined.

Nosil, P., L. J. Harmon, and O. Seehause. 2009. Ecological explanations for (incomplete) speciation. Trends in Ecology & Evolution 24:145–156. An influential review of the ecological mechanisms of speciation. The authors suggest that the completeness of speciation can be influenced by the strength of selection and the number of traits selected. However, reducing speciation to one or two primary axes may be overreduction in many cases, and the geographic and ecological drivers of divergence may be complex.

Peterson, A. T., and R. D. Holt. 2003. Niche differentiation in Mexican birds: Using point occurrences to detect ecological innovation. Ecology Letters 6:774–782. The authors note that evaluation of intraspecific variation in climatic niches is often lacking in empirical studies. They develop a protocol for comparing subspecific niche differences, and find that this can reveal conservatism or adaptive divergence within species.

Quintero, I., and J. J. Wiens. 2013. What determines the climatic niche width of species? The role of spatial and temporal climatic variation in three vertebrate clades. Global Ecology and Biogeography 22:422–432.

http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 8/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies These authors examine climatic­niche widths of species, the basis of variation for adaptive niche­evolution. They find that the climatic variation experienced at sites within a species’ range typically encompasses most of the variation found throughout the species’ range. Species with wider niches also show greater climatic divergence between localities. This provides a key insight into how ecological selection may act on geographically separated populations.

Saltz, J. B., and S. V. Nuzhdin. 2014. Genetic variation in niche construction: Implications for development and evolutionary genetics. Trends in Ecology & Evolution 29:8–14. The authors review the evidence and mechanisms for an underappreciated process of adaptive niche­evolution, the ability of organisms to influence their environment via niche construction. This has a variety of effects on the phenotype­environment correlation and influences diversification and niche dynamics in a variety of ways.

Environment and Geography

Niche evolution involves numerous disparate factors at the intersection of geographic and climatic variation, , and physiological and life­history variability. The geographic and environmental context of species’ distributions influences the paths that niche evolution can take, as do other physiological constraints related to life­history traits. For instance, the longer generation time of woody plants decreases both the rate of molecular substitutions and the rate of climatic­niche evolution and volume of climatic space explored over time, as described in Smith and Beaulieu 2009. Adaptive niche­evolution on the periphery of ranges may be a particularly common mode of speciation for more sessile organisms such as plants, as discussed in Anacker and Strauss 2014. Carnicer, et al. 2012 outlines a verbal model for speciation dynamics based on the interplay between geography, demography, and trait evolution in climatic­ niche space. On short timescales, geographic variation in climatic variables may act to separate populations even in the absence of geographic barriers to gene flow, as shown in Costa, et al. 2008. Dynesius and Jansson 2000 presents a model for global dynamics over longer timescales where geographic ranges are fractured and reinforced by repeated glacial cycles. Donoghue 2008 outlines these patterns in plants from a phylogenetic perspective. Glor and Warren 2011 introduces a suite of statistical tools for comparing climatic niches and testing evolutionary and biogeographic hypotheses. Similar versions of these tests are used in Pyron and Burbrink 2009 to show that both conservatism and divergence in climatic niches can drive speciation over short timescales, dependent on the geographic space inhabited by the lineages. Thompson 1999 discusses the geographic mosaic theory of , describing how a geographic perspective on species interactions can explain many patterns in macroecology and niche evolution. Urban, et al. 2008 introduces a community­centered view of evolutionary ecology, emphasizing the importance of spatial dimensions and the influence of dispersal.

Anacker, B. L., and S. Y. Strauss. 2014. The geography and ecology of plant speciation: Range overlap and niche divergence in sister species. Proceedings of the Royal Society B: Biological Sciences 281:20132980. Results from sister­species comparisons in the California Floristic Province suggest that “budding” speciation (isolation of peripheral populations) is common in plants and heavily driven by adaptive niche­evolution, particularly via habitat and soil selection.

Carnicer, J., L. Brotons, C. Stefanescu, and J. Penuelas. 2012. Biogeography of species richness gradients: Linking adaptive traits, demography and diversification. Biological Reviews 87:457–479. Develops primarily verbal models that link adaptive trait evolution to demography and diversification, suggesting that these processes may have broad explanatory power for large­scale biogeographic patterns. They outline a research program for testing these hypotheses.

Costa, G. C., C. Wolfe, D. B. Shepard, J. P. Caldwell, and L. J. Vitt. 2008. Detecting the influence of climatic variables on species distributions: A test using GIS niche­based models along a steep longitudinal environmental gradient. Journal of Biogeography 35:637–646.

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Donoghue, M. J. 2008. A phylogenetic perspective on the distribution of plant diversity. Proceedings of the National Academy of Sciences of the United States of America 105:11549–11555. Donoghue suggests that limited adaptive­niche evolution in climate­related traits, manifested as phylogenetic niche conservatism, explains major distributional patterns in plants. Particularly strong mechanisms include environmental filtering in local and regional assemblages, and limitations on intercontinental dispersal between climatically dissimilar regions.

Dynesius, M., and R. Jansson. 2000. Evolutionary consequences of changes in species’ geographical distributions driven by Milankovitch climate oscillations. Proceedings of the National Academy of Sciences of the United States of America 97:9115– 9120. In an influential paper, Dynesius and Jansson address the perplexing of how the apparently delicate climatic­organismal relationships observed in most species translate across the drastic arrangements of climatic space that have occurred frequently in geological history. They suggest that adaptations to climatic oscillations produce resistance to and quicker speciation, and that this may have far­ reaching explanatory power for biogeographic patterns.

Glor, R. E., and D. Warren. 2011. Testing ecological explanations for biogeographic boundaries. Evolution 65:673–683. Glor and Warren describe an influential set of techniques for assessing ecological hypotheses for biogeographic boundaries. For instance, two species’ ranges could be divided by a sharp ecological transition, or by a narrow ribbon of unsuitable habitat. This is facilitated by new metrics for quantitatively comparing ecological niche models.

Pyron, R. A., and F. T. Burbrink. 2009. Lineage diversification in a widespread species: Roles for niche divergence and conservatism in the common kingsnake, Lampropeltis getula. Molecular Ecology 18:3443–3457. The authors present empirical evidence at the phylogeographic scale that recently diverged species­pairs can exhibit both niche conservatism and niche divergence across climatic and geographic space. They suggest that while niche evolution may not be constrained per se, conservatism may simply represent the path of least resistance in many cases.

Smith, S. A., and J. M. Beaulieu. 2009. Life history influences rates of climatic niche evolution in flowering plants. Proceedings of the Royal Society B: Biological Sciences 279:4345–4352. Adaptive niche­evolution involves concerted changes in physiological tolerances, phenotypic traits, and genetic variability, arranged in a hierarchical manner. This is often difficult to analyze empirically. It is known that rates of molecular substitution differ between woody and herbaceous plants, linked to generation time. Here, the authors show that woody plants have lower rates of climatic­niche evolution and explore less climatic space over time. Thus, evolutionary dynamics differ significantly among life­history categories.

Thompson, J. N. 1999. Specific hypotheses on the geographic mosaic of coevolution. American Naturalist 155:S1–S14. The coevolution of multiple lineages simultaneously (including symbiotic and antagonistic interactions) has historically been an important driving factor in the diversity of life on earth. Coevolution is also structured at large geographic scales, oftentimes much differently than it is at local scales. The author outlines the geographic mosaic theory, offering specific hypotheses to test and suggesting future research areas and questions that might be addressed and patterns that might be explained in this geographic framework.

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Urban, M. C., M. A. Leibold, P. Amarasekare, et al. 2008. The evolutionary ecology of metacommunities. Trends in Ecology and Evolution 23:311–317. The authors discuss a dispersal­centered model of evolutionary and ecological dynamics, extending models from local spatial scales to the community level. They argue that this predicts outcomes and patterns not accounted for by previous models. Furthermore, they outline a future research program to apply these ideas to an integrative evolutionary, ecological, and genetic framework for studying species interactions in an explicitly spatial context.

Niche Conservatism

Any discussion of niche evolution must focus heavily on niche conservatism: the observed degree or similarity in ecological niches between species. Theoretical and empirical reviews are given in Crisp and Cook 2012; Peterson 2011; Pyron, et al. 2015; and Wiens, et al. 2010. Empirical investigations of the varied effects of niche conservatism on speciation and diversity patterns are outlined in Buckley, et al. 2010 and Peterson, et al. 1999. Etterson and Shaw 2001 and Futuyma 2010 discuss the genetic basis of constraint and conservatism of niches, and niche evolution by extension. Leibold and McPeek 2005 discusses how niche equivalence may arise, how neutral theory relates to older notions of niche differentiation, and how niche theory and neutral theory can be reconciled.

Buckley, L. B., T. J. Davies, D. D. Ackerly, N. J. B. Kraft, S. P. Harrison, and B. L. Anacker. 2010. Phylogeny, niche conservatism and the latitudinal diversity gradient in mammals. Proceedings of the Royal Society B: Biological Sciences 277:2131–2138. Uses a large­scale phylogenetic analysis to show that climate­richness relationships in mammals seem to result from long­term conservatism in climatic­niche space, and presumably low lability in niche evolution. They suggest that, rather than climate driving diversification, these relationships are generated by the geographic and climatic origins of clades through time, and that richness and climate are thus covariates of other underlying evolutionary mechanisms.

Crisp, M. D., and L. G. Cook. 2012. Phylogenetic niche conservatism: What are the underlying evolutionary and ecological causes? New Phytologist 196:681–694. One of many recent reviews of niche conservatism and evolution. Crisp and Cook argue that niche conservatism is a pattern only, and found only in some traits and lineages. They suggest a pattern­based ecological research program for niche evolution as it relates to speciation.

Etterson, J. R., and R. G. Shaw. 2001. Constraint to adaptive evolution in response to global warming. Science 294:151–154. Etterson and Shaw directly quantify traits related to selection and adaptive niche­evolution in multiple populations. They find that while these traits exhibit standing genetic variation, strong correlations among traits cause resistance to selection and limit adaptive evolution. Thus, rates of adaptation to changing climates are much lower than those of the climates themselves.

Futuyma, D. J. 2010. Evolutionary constraint and ecological consequences. Evolution 64:1865–1884. Futuyma offers a broad review of the mechanisms that can halt adaptive evolution and engender evolutionary constraints on traits. In particular, this may underlie a large part of phylogenetic niche conservatism, limited climatic­niche evolution, and community assembly.

Leibold, M. A., and M. A. McPeek. 2005. Coexistence of the niche and neutral perspectives in community ecology. Ecology 87:1399–1410.

http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 11/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies The authors review the relationship between neutral theory, which assumes ecological equivalence, and existing ideas about niche differentiation. They provide a verbal and mathematical framework explaining how these ideas can be reconciled. They explain how future researchers might interpret these ideas for assessing species coexistence.

Peterson, A. T. 2011. Ecological niche conservatism: A time­structured review of evidence. Journal of Biogeography 38:817–827. Peterson provides a review of the evidence and mechanisms for conservatism and adaptive niche­evolution in coarsely defined ecological niches. He finds evidence for strong conservatism generally, but warns of various methodological problems that can complicate interpretations.

Peterson, A. T., J. Soberon, and V. Sanchez­Cordero. 1999. Conservatism of ecological niches in evolutionary time. Science 285:1265–1267. An influential empirical paper showing evidence that ecological niches are commonly conserved in evolutionary time, with limited niche divergence in species pairs separated geographically.

Pyron, R. A., G. C. Costa, M. A. Patten, and F. T. Burbrink. 2015. Phylogenetic niche conservatism and the evolutionary basis of ecological speciation. Biological Reviews 90:1248–1262. The authors review niche conservatism from a ground­up perspective, integrating mechanisms of genetic constraint, canalization, physiological tolerances, and ecological selection. They suggest that niche conservatism is a specific set of processes that can lead to a variety of patterns, not simply niche similarity. This is in contrast to the other reviews in this section.

Wiens, J. J., D. D. Ackerly, A. P. Allen, et al. 2010. Niche conservatism as an emerging principle in ecology and conservation biology. Ecology Letters 13:1310–1324. A very detailed review of the mechanisms, causes, and tests of niche conservatism, and by extension, aspects of adaptive niche­ evolution.

Range Limits

Species have a fundamental and realized niche, and thus a potential and occupied geographic range. Various factors influence the extent of the latter relative to the former, and understanding the mechanisms that govern distributions can offer great insight into the processes of niche evolution. Broenniman, et al. 2012 describes a suite of tools for accurately quantifying distribution and the extent of overlap in ecological and geographic space. In a meta­analysis, Hargreaves, et al. 2014 shows that niche limits typically exceed range limits, suggesting that dispersal constraints or abiotic interactions commonly limit ranges. Hurlbert and White 2007 outlines some of the ecological correlates of these limits, while McCormack, et al. 2010 and Tingley, et al. 2009 describe how these limits can change during speciation to reinforce divergence. Valladeres, et al. 2014 and Yackulic, et al. 2015 describe mathematical models for understanding range limits with respect to across the range. Finally, Sexton, et al. 2009 offers a broad­scale review of the factors and processes that can affect range limits, distributions, and abundance.

Broenniman, O., M. C. Fitzpatrick, P. B. Pearman, B. Petitpierre, L. Pellissier, N. G. Yoccoz, et al. 2012. Measuring ecological niche overlap from occurrence and spatial environmental data. Global Ecology and Biogeography 21:481–497. Measuring even relatively simple proxies of species’ niches, such as realized climatic niche, can be difficult in practice. The authors present a suite of statistical methods leveraging occurrence and spatial environmental layers to provide comparable estimates across species as well as measure how much those estimates have changed across time.

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Hargreaves, A. L., K. E. Samis, and C. G. Eckert. 2014. Are species’ range limits simply niche limits writ large? A review of transplant experiments beyond the range. American Naturalist 183:157–173. The authors report meta­analysis of beyond­range transplant experiments, to test whether or not geographic ranges simply reflect niche limits. This is a general pattern, but a significant proportion of species exhibited niche limits greater than their realized geographic range. This suggests that dispersal limitations may play a large part in determining species’ ranges.

Hurlbert, A. H., and E. P. White. 2007. Ecological correlates of geographical range occupancy in North American birds. Global Ecology and Biogeography 16:764–773. Hurlbert and White attempt to address the ecological and trait­based factors that limit the realized occupancy of potential geographic ranges. They find that abundance at sites and wide climatic tolerances positively predict occupancy, while species with specialized niches that differ from geographic norms in climate have lower occupancy. These findings suggest ways in which species’ traits and their preferred climatic spaces may act to shape adaptive niche­evolution with respect to geographic distribution and local abundance within populations.

McCormack, J. E., A. J. Zellmer, and L. L. Knowles. 2010. Does niche divergence accompany allopatric divergence in Aphelocoma jays as predicted under ecological speciation? Insights from tests with niche models. Evolution 64:1231–1244. The authors use a model system for niche evolution, Aphelocoma jays, to test whether niche divergence is associated with speciation. They find varying levels of divergence, including low divergence in speciating lineages, but high divergence in secondary contact. The results have broad applicability for interpreting the ecospatial context of speciation.

Sexton, J. P., P. J. McIntyre, A. L. Angert, and K. J. Rice. 2009. Evolution and ecology of species range limits. Annual Review of Ecology Evolution and Systematics 40:415–436. Species’ range limits represent the nexus of numerous disparate aspects of organismal biology, including niche evolution, climatic tolerances, population demographics, and genetic variability and plasticity. The authors review these mechanisms and others in detail, forming a starting point for investigations of the role of niche evolution in geographic distribution.

Tingley, M. W., W. B. Monahan, S. R. Beissinger, and C. Moritz. 2009. Birds track their Grinnellian niche through a century of climate change. Proceedings of the National Academy of Sciences of the United States of America 106:19637–19643. In this extremely fine­scale longitudinal study, the authors track climatic niches and site occupancy for fifty­three species sampled for approximately 100 years in four transects. They find limited evidence for adaptive niche­evolution on this timescale. Instead, distributions strongly track climatic changes, indicative of niche conservatism.

Valladeres, F., S. Matesanz, F. Guilhanmon, et al. 2014. The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change. Ecology Letters 17:1351–1364. The authors develop a conceptual model and present empirical data showing how phenotypic plasticity and adaptive niche­evolution at the population level can interact to mediate responses to climate change. The results are pessimistic for the species examined, but suitable data are only available for a small number of taxa. Literature reviews reveal contrasting patterns, so more in­depth investigations are needed.

Yackulic, C. B., J. D. Nichols, J. Reid, and R. Der. 2015. To predict the niche, model colonization and extinction. Ecology 96:16–23. Several of the referenced papers in this collection address the potential versus realized niche, and occupancy versus extent in species’ ranges. Here, the authors develop a mathematical model for how occupancy­environment relationships may fluctuate over time as a http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 13/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies result of demographic processes such as colonization and extinction. Temporal variation in occupancy­environment relationships is expected when distributions are not at equilibrium. This may have strong impacts on ecological niche modeling, and the estimation and prediction of species’ niches.

Speciation Processes

Speciation and niche evolution are often tightly linked, particularly when speciation has a strong ecological component. Untangling the adaptive component of niche evolution in this process can be difficult, however. Belliure, et al. 2000 presents empirical data suggesting that reduced dispersal abilities increase local population differentiation, but increased niche plasticity is not associated with long­ distance dispersal. For less vagile organisms such as plants, Evans, et al. 2009 shows that high rates of niche evolution facilitated geographic movement and diversification. Similar patterns can be seen in birds from data presented in Cicero and Koo 2012, showing that climatic­niche divergence serves to reinforce speciation. Janzen 1967 presents what is now a classic model for how montane species­pumps occur in tropical regions, and Graham, et al. 2004 shows empirical support for climatic divergence accompanying speciation in Andean frogs. Knowles and Alvarado­Serrano 2010 presents a mathematically explicit modeling framework for studying speciation while incorporating spatial, demographic, and ecological data. Kozak and Wiens 2010 presents data suggesting that clades with higher rates of niche evolution have higher rates of speciation. Maan and Seehausen 2011 reviews the mechanisms of ecological and sexual selection on speciation. Gillespie and Roderick 2002 reviews speciation on islands, incorporating ecological and biogeographic processes.

Belliure, J., G. Sorci, A. P. Moller, and J. Clobert. 2000. Dispersal distances predict subspecies richness in birds. Journal of Evolutionary Biology 13:480–487. Reduced dispersal abilities are associated with greater subspecific diversity, suggesting greater propensity for local adaptation in less vagile lineages. However, greater dispersal ability is not associated with ecological plasticity in habitat use, suggesting that either colonization of new habitats is not significantly facilitated by long­distance dispersal, or that decreased propensity for adaptive­niche evolution limits these potential invasions.

Cicero, C., and M. S. Koo. 2012. The role of niche divergence and phenotypic adaptation in promoting lineage diversification in the Sage Sparrow (Artemisiospiza belli, Aves: Emberizidae). Biological Journal of the Linnean Society 107:332–354. A detailed, population­level study of speciation in an ecological context. Cicero and Koo show the effects of barriers in dividing populations and promoting divergence in climatic niches. Paleoclimatic approaches show how climatic niches changed through time and space to isolate populations; even lineages in secondary geographic contact are climatically isolated.

Evans, M. E. K., S. A. Smith, R. S. Flynn, and M. J. Donoghue. 2009. Climate, niche evolution, and diversification of the “bird­cage” evening primroses (Oenothera, Sections Anogra and Kleinia). American Naturalist 173:225–240. Evans and coauthors use detailed presence­locality datasets to characterize the climatic niche of species. They then reconstruct ancestral climatic niches using phylogenetic analyses to quantify rates of adaptive niche­evolution and disparity through time and space. They suggest strong relevance of these methods to detailed studies of speciation and diversification.

Gillespie, R. G., and G. K. Roderick. 2002. Arthropods on islands: Colonization, speciation, and conservation. Annual Review of 47:595–632. This influential review discusses speciation processes on island systems, ranging from islands generated de novo to those that originate from a nearby continental species pool. These various assembly processes yield different patterns of community assembly. As a result, processes of niche evolution and saturation can vary markedly.

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Graham, C. H., S. R. Ron, J. C. Santos, C. J. Schneider, and C. Mortiz. 2004. Integrating and environmental niche models to explore speciation mechanisms in dendrobatid frogs. Evolution 58:1781–1793. The authors describe an integrative framework for investigating speciation processes in an ecological and biogeographic context. Some of the methods have been supplanted (e.g., Glor and Warren 2011, cited under Environment and Geography), but many of the hypotheses and logical frameworks are still relevant. Empirically, they report that sister­species pairs exhibit common axes of ecological divergence, such as temperature and seasonality along elevational axes.

Janzen, D. H. 1967. Why mountain passes are higher in the tropics. American Naturalist 101:233–249. In an influential paper for ecology and evolution, Janzen argues that greater niche specificity in tropical species versus temperate species increases the action of barriers involving climatic gradients. Temperate species must be adapted to a wider range of climates, and can thus more easily pass through colder, drier areas at the tops of mountains. This also implies a mechanism for changing rates of niche evolution, as seen in other papers in this section.

Knowles, L. L., and D. F. Alvarado­Serrano. 2010. Exploring the population genetic consequences of the colonization process with spatio­temporally explicit models: Insights from coupled ecological, demographic and genetic models in montane grasshoppers. Molecular Ecology 19:3727–3745. Knowles and Alvarado­Serrano propose a modeling framework that includes ecological niche predictions with demographic and landscape­genetic parameters to model the population­level processes involved in speciation. This has the benefit of merging population­genetic and phylogenetic processes in a single framework with ecological niches. Future improvements could include a model of adaptive niche­evolution, which is absent here.

Kozak, K. H., and J. J. Wiens. 2010. Accelerated rates of climatic­niche evolution underlie rapid species diversification. Ecology Letters 13:1378–1389. Kozak and Wiens uses models of character evolution to quantify rates of adaptive niche­evolution in a phylogenetic context. They find that climatic­niche rates are related to net­diversification rates, that groups with faster niche evolution have faster diversification. They suggest that this is causative: the increased exploration of novel niches drives speciation.

Maan, M. E., and O. Seehausen. 2011. Ecology, sexual selection and speciation. Ecology Letters 14:591–602. Maan and Seehausen review the available evidence for interactions between sexual and natural selection on traits during speciation. They find little evidence for such interactions. However, sexual selection and environmental heterogeneity interact strongly. Traits related both to adaptive niche­evolution and sexual signaling may have strong impacts on diversification.

Invasive Species

Invasive species offer an unparalleled opportunity to study niche evolution in action over short timescales. Alien invasions allow researchers to compare native realized niches to the potentially novel climatic spaces occupied by invaders and measure plasticity and lability in physiological tolerances. Broenniman, et al. 2007 and Gallagher, et al. 2010 show evidence for adaptive niche shifts in invasive plants, beyond their native ranges. Liu, et al. 2014 shows that anthropogenic effects can strongly influence the spread of invasive species when niches are suitable, and that topographic heterogeneity and decreased congener diversity acts to limit invasions. In contrast, Petitpierre, et al. 2012 reports limited evidence for niche shifts in terrestrial plant invaders, and instead suggests that many modeling techniques may not accurately capture niche dynamics in invasions. Rodder and Lotters 2009 and Schulte, et al. 2012 report on terrestrial vertebrate invaders, finding similar evidence that apparent niche­shifts may actually represent movements within the fundamental niche, while the realized niches are constrained by different forces in the native and invasive ranges. Svensson 2012 and

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Broenniman, O., U. A. Treier, H. Muller­Scharer, W. Thuller, A. T. Peterson, and A. Guisan. 2007. Evidence of climatic niche shift during biological invasion. Ecology Letters 10:701–709. Ecological niche models are commonly treated as representing a roughly accurate picture of the potential climatic space that species can occupy and are often used to predict invasion dynamics. The authors show evidence for significant niche shifts in invasive plants that are not predicted by ordinary modeling techniques. This illustrates the potential outcomes of adaptive niche evolution and realized niches that are relatively small subsets of fundamental niches.

Gallagher, R. V., L. J. Beaumont, L. Hughes, and M. R. Leishman. 2010. Evidence for climatic niche and biome shifts between native and novel ranges in plant species introduced to Australia. Journal of Ecology 98:790–799. The authors compare the climatic niches of the native and introduced range of twenty­six plants in Australia. They find very little overlap in most cases, and that alien species commonly inhabit climatic niches far outside their native habitats. Climatic differences may not present a strong barrier to colonization, particularly if mechanisms such as ecological release, phenotypic plasticity, or rapid adaptive niche­evolution come into play.

Liu, X., X. Li, Z. Liu, et al. 2014. Congener diversity, topographic heterogeneity and human­assisted dispersal predict spread rates of alien herpetofauna at a global scale. Ecology Letters 17:821–829. As with several other studies cited here, Liu and others find that alien invasions are unpredictable, and often occur far outside a species’ native niche. Richness of native congeners and human activities both increase the rate of spread, while expansion is limited by topographic heterogeneity. The influence of plasticity versus rapid niche­evolution is unclear.

Petitpierre, B., C. Kueffer, O. Broenniman, C. Randin, C. Daehler, and A. Guisan. 2012. Climatic niche shifts are rare among terrestrial plant invaders. Science 335:1344–1348. In contrast to some of the other referenced papers in this article, these authors report limited climatic­niche shifts in invasive terrestrial plants. They suggest that niche models from species’ native ranges may be good predictors of potential invasion spaces.

Rodder, D., and S. Lotters. 2009. Niche shift versus niche conservatism? Climatic characteristics of the native and invasive ranges of the Mediterranean house gecko (Hemidactylus turcicus). Global Ecology and Biogeography 18:674–687. The authors examine one of the most widely distributed invasive species in the world, to examine the degree of niche shifts and conservatism in introduced lineages. They find variable evidence for shifts and conservatism based on input variables. This suggests that habitat selection, background effects, and predictor selection may influence conclusions. The use of null models is strongly indicated to counteract these variables.

Schulte, U., A. Hochkich, S. Lotters, et al. 2012. Cryptic niche conservatism among evolutionary lineages of an invasive lizard. Global Ecology and Biogeography 21:198–211. In a very detailed population­level study, the authors report the existence of cryptic niche conservatism in invasive lineages, which may confound distribution modeling for alien species. Invasive lineages often had invasive niches outside their native range, suggesting that realized niches might be constrained by stochastic invasion dynamics, but that fundamental niches might be much broader. Thus, niches may be cryptically conserved, confounding modeling attempts.

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Sexton, J. P., S. B. Hangartner, and A. A. Hoffman. 2014. Genetic isolation by environment or distance: Which pattern of gene flow is most common? Evolution 68:1–15. Adaptive niche­evolution may be constrained or promoted by gene flow from surrounding populations. The authors review the literature for the prevalence of isolation by distance and environment, finding strong support for isolation by environment in a majority (74.3 percent) of cases. This suggests that ecological divergence is a primary factor limiting gene flow.

Svensson, E. I. 2012. Non­ecological speciation, niche conservatism and thermal adaptation: How are they connected? Organisms Diversity & Evolution 12:229–240. Svensson points out that reproductive isolation is often observed in concert with niche conservatism, potentially arguing against a widespread role for ecological speciation. He reviews ways in which reproductive isolation might be decoupled from adaptive niche evolution.

Community Perspectives

While niche evolution is commonly studied within lineages or single species, the interaction of species in a community likely represents a strong driver of evolutionary dynamics in many cases. Thus, a community perspective on adaptive niche­evolution is desirable both to understand how species interact and how these interactions can affect niche shifts. Ackerly 2003 tackles many of these issues in a broad­scale review, outlining scenarios that may promote adaptive niche evolution. The influential review Chesson and Huntly 1997 provides models for how fluctuating conditions of environmental can promote species coexistence and select for certain traits at the population level, leading to niche shifts. Araya, et al. 2012 suggests that traits related to harsh conditions, such as drought tolerance, might be highly labile and heavily convergent in different areas where they are observed. Pillar and Duarte 2010 introduces a phylogenetic framework for assessing niche conservatism within phylogenies. Ackerly, et al. 2006 presents data suggesting that within­ community niche traits are likely to diverge first during adaptive radiation, followed by climatic­niche preferences at higher levels. Soberon 2007 and Zink 2014 review niche concepts more generally, with applications to how niche differences allow species to coexist locally. Finally, Sinervo, et al. 2010 presents sobering evidence that minor changes in climate at the upper end of lizards’ thermal tolerances have drastically eroded diversity at many sites, and are predicted to do so increasingly during global climate change.

Ackerly, D. D. 2003. Community assembly, niche conservatism, and adaptive evolution in changing environments. International Journal of Plant Sciences 164:S165–S184. Ackerly outlines the multiscale nature of organism­environment interactions, comprising plasticity, ecological sorting, and adaptive evolution of functional traits. Three broad scenarios are outlined that may promote adaptive niche­evolution: (i) colonization of novel habitats, (ii) trailing edges during climate change, and (iii) shifts in realized environmental space within the species’ fundamental niche, changing the potential niche with respect to the niche optimum.

Ackerly, D. D., D. W. Schwilk, and C. O. Webb. 2006. Niche evolution and adaptive radiation: Testing the order of trait divergence. Ecology 87:S50–S61. Niches are constantly evolving, but the traits under strongest selection during divergence, and the rate at which they diverge, may vary drastically among lineages and ecological scenarios. Ackerly and colleagues report evidence that within­community niche differences are most likely to evolve first during adaptive radiation, followed by climatic­niche differences, though high rates of climatic­niche divergence may obscure these patterns.

Araya, Y. N., J. Silvertown, D. J. Gowing, K. J. McConway, H. P. Linder, and G. Midgley. 2012. Do niche­structured plant communities exhibit phylogenetic conservatism? A test case in an endemic clade. Journal of Ecology 100:1434–1439. http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 17/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies Combined ecological and phylogenetic analyses suggest that crucial traits related to hydrological niche responses (such as tolerance to drought) structuring communities are evolutionarily labile, and are heavily convergent within assemblages.

Chesson, P., and N. Huntly. 1997. The roles of harsh and fluctuating conditions in the dynamics of ecological communities. American Naturalist 150:519–553. Fundamental paper on the ecological processes affecting species interactions, and the impact of disturbance on niche availability and species coexistence. The mechanisms discussed have far­reaching implications for mechanisms underlying adaptive niche evolution at the population level.

Pillar, V. D., and L. D. S. Duarte. 2010. A framework for metacommunity analysis of phylogenetic structure. Ecology Letters 13:587–596. The authors introduce a phylogenetic framework for analyzing community assembly patterns. In particular, they address how incorporating species’ average trait values may provide evidence for conservatism or ecological filtering along gradients.

Sinervo, B., F. Méndez­de­la­Cruz, D. B. Miles, et al. 2010. Erosion of lizard diversity by climate change and altered thermal niches. Science 328:894–899. The authors use historical data and physiological models to assess historical and future risks. Under even moderate climate­ change scenarios, they predict that adaptive niche­evolution will not keep pace in a significant proportion of lizard diversity, which will be at high risk for extinction.

Soberon, J. 2007. Grinnellian and Eltonian niches and geographic distributions of species. Ecology Letters 10:1115–1211. Species’ ranges are affected both by broad­scale and primarily abiotic variables (Grinnellian niche) and local­scale and primarily biotic variables (Eltonian niche). Soberon reviews the theoretical basis of these dynamics and clarifies major questions related to geography, macroecology, and adaptive niche­evolution.

Zink, R. M. 2014. Homage to Hutchinson, and the role of ecology in lineage divergence and speciation. Journal of Biogeography 41:999–1006. Zink reviews mechanisms of adaptive niche­evolution influenced by ecological selection and finds a strong role therein for promoting speciation. However, niche conservatism also affects species’ traits, and morphological divergence may often be a mechanism to facilitate co­occurrence, rather than a byproduct of divergent ecological selection.

Historical Perspectives

Studying niche evolution from palaeontological or historical data can be difficult, as relevant variables often can’t be measured directly. However, detailed datasets have been created to evaluate niche dynamics in deep and recent time by numerous authors. Eldredge, et al. 2005 presents a verbal model for how high rates of short­term niche evolution may be homogenized by geographic variability to produce the long­term stasis often seen in the fossil record. Ricklefs 2004 discusses the integration of evolution, including niches, over temporal and spatial scales to understand diversity. Holland and Zaffos 2011 presents a dataset for benthic marine invertebrates that shows strong conservatism in microhabitat and abundance, but high rates of niche evolution in environmental tolerances. Pearman, et al. 2008; Lavergne, et al. 2013; and Araujo, et al. 2013 use recent­occurrence datasets to document niche shifts over short timescales and evaluate the performance of predictive modeling approaches. Roberts and Hamann 2012 and Stigall 2012 take a similar approach with paleontological data. Prinzing, et al. 2001 suggests that paleo­adaptations to climatic instability are reflected in present­day

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Araujo, M. B., F. Ferri­Yanez, F. Bozinovic, P. A. Marquet, F. Valladeres, and S. L. Chown. 2013. Heat freezes niche evolution. Ecology Letters 16:1206–1219. Using a broad comparative dataset, the authors show that heat tolerance is generally conserved, while cold tolerance shows large shifts among and between lineages. They suggest that physiological limits constrain heat tolerances, while cold tolerances should be more labile. Thus, models of extinction risk under climate change may overestimate sensitivity of cold­adapted lineages, while responses of lineages near their upper thermal tolerances may be more predictable. Niche­evolution estimates should take these differences into account.

Eldredge, N., J. N. Thompson, P. M. Brakefield, et al. 2005. The dynamics of evolutionary stasis. Paleobiology 31:133–145. Reviews another historical conundrum related to adaptive niche­evolution: why is stasis so common in the fossil record when extant populations are so variable? They suggest a primarily verbal model based on ecological and demographic parameters that links geographical complexity to short­term variability and long­term stasis.

Holland, S. M., and A. Zaffos. 2011. Niche conservatism along an onshore­offshore gradient. Paleobiology 37:270–286. Holland and Zaffos use a detailed stratigraphic and paleomorphological dataset to quantify rates of adaptive niche­evolution in benthic marine invertebrates. They find strong conservatism in abundance and microhabitat, but little in environmental tolerance. Again, ecological constraint seems to be a primary factor in limiting niche evolution and governing geographic distributions.

Lavergne, S., M. K. E. Evans, I. J. Burfield, F. Jiguet, and W. Thuiller. 2013. Are species’ responses to global change predicted by past niche evolution? Philosophical Transactions of the Royal Society B 368:20120091. Using a large­scale comparative dataset for birds, the authors show that propensity for demographic decline is linked to lower rates of climatic­niche evolution. This is true at the species level, and for deeper lineages. This may allow for better prediction of future dynamics based on past responses to climate change.

Pearman, P. B., A. Guisan, O. Broenniman, and C. F. Randin. 2008. Niche dynamics in space and time. Trends in Ecology & Evolution 23:149–158. The authors review evidence for short­term niche dynamics within the last hundred years. They identify cases of both conservatism and adaptive niche­evolution. They suggest that combining ecological niche modeling with phylogenetic methods can help improve the accuracy and predictive power of niche models, particularly with respect to assumptions of conservatism.

Prinzing, A., W. Durka, S. Klotz, and R. Brandl. 2001. The niche of higher plants: Evidence for phylogenetic conservatism. Proceedings of the Royal Society B: Biological Sciences 268:2383–2389. The authors present evidence that significant niche conservatism is present in higher­level plant lineages, and that adaptive niche­ evolution has been constrained. They suggest that this represents the present­day persistence of adaptations to paleo­environmental trends.

Ricklefs, R. E. 2004. A comprehensive framework for global patterns in . Ecology Letters 7:1–15.

http://www.oxfordbibliographies.com/view/document/obo­9780199941728/obo­9780199941728­0075.xml?rskey=WDTkgc&result=51&print 19/20 8/23/2016 Niche Evolution ­ Evolutionary Biology ­ Oxford Bibliographies Ricklefs gives a broad overview of the range of temporal and spatial scales that need to be considered to understand diversity. This includes niche evolution at local scales and over longer time periods. He lays out eight distinct research tracks that need to be considered in an integrative ecological and evolutionary explanation of diversity.

Roberts, D. R., and A. Hamann. 2012. Predicting potential climate change impacts with bioclimate envelope models: A palaeoecological perspective. Global Ecology and Biogeography 21:121–133. The authors take an unorthodox approach to analyzing the predictive power of ecological niche models by retrospectively validating models using palynological and fossil data. They find moderate performance in most cases and suggest that no­analogue climates may not confound future projections to an unacceptable degree.

Stigall, A. L. 2012. Using ecological niche modelling to evaluate niche stability in deep time. Journal of Biogeography 39:772–781. Stigall uses a detailed paleocological dataset to assess morphological and ecological stasis in marine brachiopods. She finds high morphological stasis through time, but high ecological stasis (niche conservatism) only during periods of gradual climatic change. In contrast, rates of adaptive niche­evolution keep the pace with the environment during rapid climatic changes.

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