niche evolution - george washington universityrpyron/publications/pyron_2016a.pdf · 2016-09-12 ·...

8/23/2016 Niche Evolution Evolutionary Biology Oxford Bibliographies

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Niche EvolutionAlex Pyron

LAST MODIFIED: 26 MAY 2016DOI: 10.1093/OBO/97801999417280075

Introduction

The evolution of species’ niches is a process that is fundamental to investigations in numerous fields of biology, including speciation,community assembly, and longterm regional and global diversification processes. It forms the nexus between ecological andevolutionary questions. Topics as diverse as ecological speciation, niche conservatism, species coexistence, and historicalbiogeography all rely on interpreting patterns and drivers of species’ niches through time and across landscapes. Despite thisimportance, a distinct research agenda concerning niche evolution as a discrete topic of inquiry has yet to emerge. Niche evolution isoften considered as a sidebar or of secondary importance when addressing questions such as “how did two species diverge?” Basicquestions 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 somesemblance of form to a framework for interpreting and evaluating niche evolution, and outlining major areas for future research from anevolutionary 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 inliterature is intertwined with that of speciation, ecology, and diversification. Mayr 1942 provided the original background material forrecent developments, during the Modern Synthesis. Williams 1966 focuses on adaptation at the individual and population level in atheoretical context, providing the necessary background to interpret processes of niche evolution. Schluter 2000 describes the ecologicalbasis of adaptive evolutionary radiations, and how niche evolution drives ecomorphological diversification. Chase and Leibold 2003 is aclassic treatise on what a niche is, and how to interpret niche concepts in practice. OdlingSmee, et al. 2003 approaches niche evolutionfrom the perspective of the individual interacting with its environment, and the process of niche construction. Coyne and Orr 2004 coversthe various mechanisms that drive speciation, which includes heavy focus on ecological isolation and niche evolution. Losos 2009 is acase study of niche evolution in the classic modelsystem Anolis. Finally, Peterson, et al. 2011 covers a profusion of methodologicalapproaches to understanding niche evolution.

Chase, J. M., and M. A. Leibold. 2003. Ecological niches: Linking classical and contemporary approaches. Chicago: Univ. ofChicago Press.

A more indepth look at some of the issues described in this section, including niche concepts and individual, population, and specieslevel 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.

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One of the major overviews of speciation processes, and the ecological factors that can underlie lineage divergence leading tospeciation. In particular, chapter 5 (“Ecological Isolation”) details nicherelated processes such as habitat, pollinator, and temporalisolation. 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 Anolislizards. Losos reviews and synthesizes available studies, with particular emphasis on climatic and ecomorphological adaptations theselizards have developed to occupy novel nichespaces.

Mayr, E. 1942. Systematics 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 geneticbasis of ecological characteristics, and the accumulation of mutations related to this, is recognized as a key driver of speciation, implicitlyvia 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 overallecological niches is an underappreciated force in the diversification of lineages. The authors review the evidence and implications ofniche 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: PrincetonUniv. Press.

A crucial text covering more recent developments in the quantitative analysis of species’ niches, with a particular focus on modeling andpredicting species’ niches and distributions using modern computational techniques. Includes background material from many of thereferences listed in this section with respect to niche concepts, and how niches may change over shorter timescales. In contrast, theauthors do not heavily feature a historical perspective, or much address how niche evolution through time affects presentdaydistributions.

Schluter, D. 2000. The ecology of adaptive radiation. Oxford: Oxford Univ. Press.

Outlines the ecological basis of rapid speciation in the face of increased ecological opportunity, leading to adaptive radiations. Includesa detailed discussion on the mechanisms of ecological speciation and the basis of niche divergence among species, which is thefundamental unit of niche evolution.

Williams, G. C. 1966. Adaptation and natural selection. Princeton, NJ: Princeton Univ. Press.

A classic of post“modern synthesis” evolutionary theory. Describes the epistemological as well as the mechanistic basis of adaptationand natural selection in populations, particularly with respect to ecological aspects. Outlines the multilevel nature of adaptation, which isimportant when considering niche evolution.

Textbooks



As a specialized subject, niche evolution has not been the subject of full textbooks. However, the fundamental principles of nicheevolution are found in diverse fields in biology, and a number of basic textbooks in ecology and evolution contain the building blocks forstudying adaptive nicheevolution. Begon, et al. 2006 is a broadly introductory text on the science of ecology. Futuyma 2013 is similarly abroad, foundational text in evolutionary biology. Pianka 1999 introduced many of the historically important concepts in evolutionaryecology 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 ecosystems. 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 keyto understanding niche evolution, as are the basic definitions of the ecological niche, 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 extinctorganisms. Nevertheless, a historical focus is an important aspect of broadscale studies of niche evolution, and a text such as thiscovers 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 ecosystemperspective, so is an evolution textbook needed. Futuyma covers topics such as biogeography, speciation, and the origin of geneticvariation 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 tounderstand how cooccurring species can coexist locally and form ecological communities. The interaction of species on small temporaland geographic scales forms the basis for longerterm adaptations, and thus community ecology is an important foundation for nicheevolution.

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 nicheevolution. Importantly, the sixth edition introduces the phylogenetic perspective needed to compare traits across species, usingcomputational 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, recentresearch on niches, niche conservatism, and niche evolution can be found in a wide breadth of journals focused on ecology andevolutionary 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 inecology across a variety of subfields. Ecology Letters is a shorter form, highimpact journal for outstandingly novel results inmacroecology and evolution. Evolution focuses primarily on evolutionary questions, but frequently entertains ecological approaches toevolutionary theory. Global Ecology and Biogeography typically publishes papers dealing with the intersection of evolutionary andmacroecological processes in a biogeographic context. Journal of Biogeography typically focuses on biogeographic analyses, but















includes recent, more ecologically focused inquiries, such as phylogeography. Molecular Ecology typically focuses on populationgenetics, speciation, and phylogeography, but commonly integrates ecological perspectives on niche evolution to answer thesequestions.

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 andniche 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 inmany 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 significantproportion 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 dowe mean by “evolution”? How do we measure niches? Which variables do we include? Are we speaking of abiotic or biotic factors, or acombination thereof? Elith and Leathwick 2009 summarizes available methods for quantifying species’ ecological niches in a climaticframework. 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


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summarizes the use of GIS (geographic information system) data and methods to quantify the geographic and climatic space occupiedby species, to characterize niches. When comparing niches among species, variable amounts of divergence and conservatism are to beexpected. But how do we interpret this? Losos 2008 addresses these issues. The ways in which niche evolution can influence speciationin an ecological context are numerous, and are addressed in detail in Schluter 2001 and Rundle and Nosil 2005. Finally, geographicand 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. Species distribution 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 tounderstand species’ distributions and climatic adaptations. Such methods form a foundation of evolutionary and ecological studies ofniche evolution.

Hadly, E. A., P. A. Spaeth, and C. Li. 2009. Niche conservatism above the species level. Proceedings of the National Academy ofSciences 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 lefttheir aquatic niche. Range size in mammals seems to show the signal of higherlevel conservatism, and longterm constraints onadaptive 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 ofthe 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, hediscusses how niche evolution and conservatism may play out across phylogenetic scales. He offers a variety of forwardlookingquestions 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 inEcology & 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 relatednessand 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) withinlineages. He discusses case studies, tests, and implications for future research. He concludes that the issues involved are highlydependent 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 geographicspeciation. Journal of Biogeography 37:2005–2015.

The authors present a verbal model for the interaction of niche conservatism, adaptive nicheevolution, and geographic and climaticvariation 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.







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 interplaybetween divergent ecological selection, adaptive nicheevolution, and speciation. This forms the basis for understanding adaptivedifferentiation 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 ofindividuals. Thus, diversity within species may be an important source of variation for niche evolution. Finescale geographic patternsmay contain a great deal of information for understanding ecological adaptations. For instance, Gee 2004 shows how ecologicallimitations on dispersal limit genetic contact between recently diverged populations. FisherReid, et al. 2012 asks whether or not specieswith narrower niches have higher rates of niche evolution, which does not seem to be the case. Ecological constraints may influence theevolution of lifehistory traits, and thus the prevalence of behavioral adaptations for coexistence, as reported in Hatchwell and Komdeur2000. Holt, et al. 2004 describes a mathematical model showing how sourcesink dynamics can be maintained by temporal variation inthe adaptive landscape. Pulliam 2000 discusses how demographic factors can, in part, explain mismatches between potential and actualdistributions. Rangel, et al. 2007 uses a simulation approach to show how spatial variation in conservatism and divergence amongpopulations 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 ofclimaticniche 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, certainvariables do evolve more quickly in species with broader niches.

Gee, J. M. 2004. Gene flow across a climatic barrier between hybridizing avian species, California and Gambel’s quail (Callipeplacalifornica and C. gambelii). Evolution 58:1108–1121.

An interesting empirical example of how climaticniche differences between recently diverged populations can act to limit hybridizationand gene flow. Morphological traits are closely linked to the climatic variables that vary across the hybrid zone, yielding a complex suiteof 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. AnimalBehaviour 59:1079–1086.

Niche evolution results from a complex interplay of processes including ecological constraints, demographics, and biogeography. Oftenoverlooked aspects are behavioral processes such as individual interactions. Hatchwell and Komdeur review how lifehistory traits andecological factors interact to facilitate the evolution of complex behaviors such as cooperation.

Holt, R. D., M. Barfield, and R. Gomulkiewicz. 2004. Temporal variation can facilitate niche evolution in harsh sink environments.American Naturalist 164:187–200.











The authors evaluate several models for nicheevolution dynamics in sink habitats. They find that temporal variation in the adaptivelandscape of the sink can facilitate its persistence and permit niche evolution in marginal populations. This may explain dynamicsobserved in invasive species 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 potentialgeographic range) and its realized niche (and thus its occupied geographic range). Pulliam reviews niche concepts related to occupancyand 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. Species richness and evolutionary niche dynamics: A spatialpatternoriented 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 onspecies’ ranges, explaining a large proportion of variance in observed speciesrichness patterns. This gives a framework for interpretingthe 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 theUnited 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 standinggenetic variation related to ecological variability. This preexisting diversity may explain the frequency and rapidity of ecologicalspeciation 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 areinsensitive to mutations (genetic) or ecological (environmental) factors. This phenomenon has broad implications for studies of adaptivenicheevolution.

Niche Plasticity

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











Anderson, R. P., and A. Raza. 2010. The effect of the extent of the study region on GIS models of species geographic distributionsand estimates of niche evolution: Preliminary tests with montane rodents (genus Nephelomys) in Venezuela. Journal ofBiogeography 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 ondistribution models, and potentially on estimated rates of niche evolution. They suggest that distribution models be conditioned on morelimited areas containing only the known, realized distribution, and that more accurate estimates can then be obtained by projecting theseinto other areas.

Boucher, F. C., W. Thuiller, T. J. Davies, and S. Lavergne. 2014. Neutral biogeography and the evolution of climatic niches. AmericanNaturalist 183:573–584.

Simulation results from a spatially explicit model suggest that the geographic dimension of speciation exerts a strong influence onobserved patterns of “niche conservatism,” and that tests of niche evolution must consider physiological traits and spatial contexts to berelevant.

Jackson, S. T., J. L. Betancourt, R. K. Booth, and S. T. Gray. 2009. Ecology and the ratchet of events: Climate variability, nichedimensions, and species distributions. Proceedings of the National Academy of Sciences of the United States of America106:19685–19692.

The authors review a variety of mechanisms and processes related to understanding niche evolution in the face of natural andanthropogenic 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 Anolissagrei group. Ecology 87:S29–S38.

Using phylogenetic and GIS data, the authors test whether niche similarity is related to phylogenetic similarity. Surprisingly (or perhapsnot), 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 & Evolution24:145–156.

An influential review of the ecological mechanisms of speciation. The authors suggest that the completeness of speciation can beinfluenced by the strength of selection and the number of traits selected. However, reducing speciation to one or two primary axes maybe 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 ecologicalinnovation. 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 protocolfor 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 climaticvariation in three vertebrate clades. Global Ecology and Biogeography 22:422–432.



These authors examine climaticniche widths of species, the basis of variation for adaptive nicheevolution. They find that the climaticvariation 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 ecologicalselection may act on geographically separated populations.

Saltz, J. B., and S. V. Nuzhdin. 2014. Genetic variation in niche construction: Implications for development and evolutionarygenetics. Trends in Ecology & Evolution 29:8–14.

The authors review the evidence and mechanisms for an underappreciated process of adaptive nicheevolution, the ability of organismsto influence their environment via niche construction. This has a variety of effects on the phenotypeenvironment correlation andinfluences 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, genetic diversity, andphysiological and lifehistory variability. The geographic and environmental context of species’ distributions influences the paths thatniche evolution can take, as do other physiological constraints related to lifehistory traits. For instance, the longer generation time ofwoody plants decreases both the rate of molecular substitutions and the rate of climaticniche evolution and volume of climatic spaceexplored over time, as described in Smith and Beaulieu 2009. Adaptive nicheevolution on the periphery of ranges may be a particularlycommon mode of speciation for more sessile organisms such as plants, as discussed in Anacker and Strauss 2014. Carnicer, et al. 2012outlines a verbal model for speciation dynamics based on the interplay between geography, demography, and trait evolution in climaticniche space. On short timescales, geographic variation in climatic variables may act to separate populations even in the absence ofgeographic barriers to gene flow, as shown in Costa, et al. 2008. Dynesius and Jansson 2000 presents a model for global dynamics overlonger timescales where geographic ranges are fractured and reinforced by repeated glacial cycles. Donoghue 2008 outlines thesepatterns in plants from a phylogenetic perspective. Glor and Warren 2011 introduces a suite of statistical tools for comparing climaticniches and testing evolutionary and biogeographic hypotheses. Similar versions of these tests are used in Pyron and Burbrink 2009 toshow that both conservatism and divergence in climatic niches can drive speciation over short timescales, dependent on the geographicspace inhabited by the lineages. Thompson 1999 discusses the geographic mosaic theory of coevolution, describing how a geographicperspective on species interactions can explain many patterns in macroecology and niche evolution. Urban, et al. 2008 introduces acommunitycentered 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 insister 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 peripheralpopulations) 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 adaptivetraits, demography and diversification. Biological Reviews 87:457–479.

Develops primarily verbal models that link adaptive trait evolution to demography and diversification, suggesting that these processesmay have broad explanatory power for largescale biogeographic patterns. They outline a research program for testing thesehypotheses.

Costa, G. C., C. Wolfe, D. B. Shepard, J. P. Caldwell, and L. J. Vitt. 2008. Detecting the influence of climatic variables on speciesdistributions: A test using GIS nichebased models along a steep longitudinal environmental gradient. Journal of Biogeography35:637–646.













Using ecological niche modeling techniques, Costa and others suggest that adaptive niche evolution across climatic axes can act toseparate populations, even in regions that lack strong geographic barriers to dispersal. Multiple speciespairs are in contact acrossecological gradients in the central United States, but not divided allopatrically by features such as mountains or rivers.

Donoghue, M. J. 2008. A phylogenetic perspective on the distribution of plant diversity. Proceedings of the National Academy ofSciences 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 regionalassemblages, and limitations on intercontinental dispersal between climatically dissimilar regions.

Dynesius, M., and R. Jansson. 2000. Evolutionary consequences of changes in species’ geographical distributions driven byMilankovitch 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 relationshipsobserved 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 extinction and quicker speciation, and that this may have farreaching 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. Forinstance, two species’ ranges could be divided by a sharp ecological transition, or by a narrow ribbon of unsuitable habitat. This isfacilitated 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 andconservatism 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 nicheconservatism and niche divergence across climatic and geographic space. They suggest that while niche evolution may not beconstrained 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 ofthe Royal Society B: Biological Sciences 279:4345–4352.

Adaptive nicheevolution involves concerted changes in physiological tolerances, phenotypic traits, and genetic variability, arranged in ahierarchical manner. This is often difficult to analyze empirically. It is known that rates of molecular substitution differ between woody andherbaceous plants, linked to generation time. Here, the authors show that woody plants have lower rates of climaticniche evolution andexplore 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 animportant driving factor in the diversity of life on earth. Coevolution is also structured at large geographic scales, oftentimes muchdifferently than it is at local scales. The author outlines the geographic mosaic theory, offering specific hypotheses to test and suggestingfuture research areas and questions that might be addressed and patterns that might be explained in this geographic framework.



Urban, M. C., M. A. Leibold, P. Amarasekare, et al. 2008. The evolutionary ecology of metacommunities. Trends in Ecology andEvolution 23:311–317.

The authors discuss a dispersalcentered model of evolutionary and ecological dynamics, extending models from local spatial scales tothe community level. They argue that this predicts outcomes and patterns not accounted for by previous models. Furthermore, theyoutline a future research program to apply these ideas to an integrative evolutionary, ecological, and genetic framework for studyingspecies 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 nichesbetween species. Theoretical and empirical reviews are given in Crisp and Cook 2012; Peterson 2011; Pyron, et al. 2015; and Wiens, etal. 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 andconservatism of niches, and niche evolution by extension. Leibold and McPeek 2005 discusses how niche equivalence may arise, howneutral 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 andthe 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 longtermconservatism in climaticniche space, and presumably low lability in niche evolution. They suggest that, rather than climate drivingdiversification, these relationships are generated by the geographic and climatic origins of clades through time, and that richness andclimate 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, andfound only in some traits and lineages. They suggest a patternbased ecological research program for niche evolution as it relates tospeciation.

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 whilethese 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. Inparticular, 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. Ecology87:1399–1410.












The authors review the relationship between neutral theory, which assumes ecological equivalence, and existing ideas about nichedifferentiation. They provide a verbal and mathematical framework explaining how these ideas can be reconciled. They explain howfuture 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 definedecological niches. He finds evidence for strong conservatism generally, but warns of various methodological problems that cancomplicate interpretations.

Peterson, A. T., J. Soberon, and V. SanchezCordero. 1999. Conservatism of ecological niches in evolutionary time. Science285:1265–1267.

An influential empirical paper showing evidence that ecological niches are commonly conserved in evolutionary time, with limited nichedivergence 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 ofecological 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 avariety 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 conservationbiology. Ecology Letters 13:1310–1324.

A very detailed review of the mechanisms, causes, and tests of niche conservatism, and by extension, aspects of adaptive nicheevolution.

Range Limits

Species have a fundamental and realized niche, and thus a potential and occupied geographic range. Various factors influence theextent of the latter relative to the former, and understanding the mechanisms that govern distributions can offer great insight into theprocesses of niche evolution. Broenniman, et al. 2012 describes a suite of tools for accurately quantifying distribution and the extent ofoverlap in ecological and geographic space. In a metaanalysis, Hargreaves, et al. 2014 shows that niche limits typically exceed rangelimits, suggesting that dispersal constraints or abiotic interactions commonly limit ranges. Hurlbert and White 2007 outlines some of theecological correlates of these limits, while McCormack, et al. 2010 and Tingley, et al. 2009 describe how these limits can change duringspeciation to reinforce divergence. Valladeres, et al. 2014 and Yackulic, et al. 2015 describe mathematical models for understandingrange limits with respect to abundance across the range. Finally, Sexton, et al. 2009 offers a broadscale review of the factors andprocesses 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 nicheoverlap 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 authorspresent a suite of statistical methods leveraging occurrence and spatial environmental layers to provide comparable estimates acrossspecies as well as measure how much those estimates have changed across time.











Hargreaves, A. L., K. E. Samis, and C. G. Eckert. 2014. Are species’ range limits simply niche limits writ large? A review oftransplant 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 nichelimits. 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. GlobalEcology and Biogeography 16:764–773.

Hurlbert and White attempt to address the ecological and traitbased factors that limit the realized occupancy of potential geographicranges. They find that abundance at sites and wide climatic tolerances positively predict occupancy, while species with specializedniches that differ from geographic norms in climate have lower occupancy. These findings suggest ways in which species’ traits and theirpreferred climatic spaces may act to shape adaptive nicheevolution with respect to geographic distribution and local abundance withinpopulations.

McCormack, J. E., A. J. Zellmer, and L. L. Knowles. 2010. Does niche divergence accompany allopatric divergence in Aphelocomajays 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. Theresults 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 ofEcology Evolution and Systematics 40:415–436.

Species’ range limits represent the nexus of numerous disparate aspects of organismal biology, including niche evolution, climatictolerances, 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 climatechange. 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 forapproximately 100 years in four transects. They find limited evidence for adaptive nicheevolution on this timescale. Instead, distributionsstrongly 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 ofspecies 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 atthe population level can interact to mediate responses to climate change. The results are pessimistic for the species examined, butsuitable data are only available for a small number of taxa. Literature reviews reveal contrasting patterns, so more indepth investigationsare 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



result of demographic processes such as colonization and extinction. Temporal variation in occupancyenvironment relationships isexpected when distributions are not at equilibrium. This may have strong impacts on ecological niche modeling, and the estimation andprediction of species’ niches.

Speciation Processes

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

Belliure, J., G. Sorci, A. P. Moller, and J. Clobert. 2000. Dispersal distances predict subspecies richness in birds. Journal ofEvolutionary Biology 13:480–487.

Reduced dispersal abilities are associated with greater subspecific diversity, suggesting greater propensity for local adaptation in lessvagile lineages. However, greater dispersal ability is not associated with ecological plasticity in habitat use, suggesting that eithercolonization of new habitats is not significantly facilitated by longdistance dispersal, or that decreased propensity for adaptivenicheevolution limits these potential invasions.

Cicero, C., and M. S. Koo. 2012. The role of niche divergence and phenotypic adaptation in promoting lineage diversification in theSage 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 dividingpopulations and promoting divergence in climatic niches. Paleoclimatic approaches show how climatic niches changed through timeand 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 reconstructancestral 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 ofEntomology 47:595–632.

This influential review discusses speciation processes on island systems, ranging from islands generated de novo to those that originatefrom 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.












Graham, C. H., S. R. Ron, J. C. Santos, C. J. Schneider, and C. Mortiz. 2004. Integrating phylogenetics and environmental nichemodels 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. Someof the methods have been supplanted (e.g., Glor and Warren 2011, cited under Environment and Geography), but many of thehypotheses and logical frameworks are still relevant. Empirically, they report that sisterspecies pairs exhibit common axes of ecologicaldivergence, 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 temperatespecies 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 ofniche 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 withspatiotemporally 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 andlandscapegenetic parameters to model the populationlevel processes involved in speciation. This has the benefit of mergingpopulationgenetic and phylogenetic processes in a single framework with ecological niches. Future improvements could include amodel of adaptive nicheevolution, which is absent here.

Kozak, K. H., and J. J. Wiens. 2010. Accelerated rates of climaticniche evolution underlie rapid species diversification. EcologyLetters 13:1378–1389.

Kozak and Wiens uses models of character evolution to quantify rates of adaptive nicheevolution in a phylogenetic context. They findthat climaticniche rates are related to netdiversification rates, that groups with faster niche evolution have faster diversification. Theysuggest 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 relatedboth 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 allowresearchers to compare native realized niches to the potentially novel climatic spaces occupied by invaders and measure plasticity andlability in physiological tolerances. Broenniman, et al. 2007 and Gallagher, et al. 2010 show evidence for adaptive niche shifts ininvasive plants, beyond their native ranges. Liu, et al. 2014 shows that anthropogenic effects can strongly influence the spread ofinvasive 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 manymodeling techniques may not accurately capture niche dynamics in invasions. Rodder and Lotters 2009 and Schulte, et al. 2012 reporton terrestrial vertebrate invaders, finding similar evidence that apparent nicheshifts may actually represent movements within thefundamental niche, while the realized niches are constrained by different forces in the native and invasive ranges. Svensson 2012 and











Sexton, et al. 2014 review mechanisms that link geographic and ecological dynamics to niche evolution and speciation. This hasrelevance both for “natural” and invasive range expansions and divergence.

Broenniman, O., U. A. Treier, H. MullerScharer, W. Thuller, A. T. Peterson, and A. Guisan. 2007. Evidence of climatic niche shiftduring 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 speciescan occupy and are often used to predict invasion dynamics. The authors show evidence for significant niche shifts in invasive plants thatare not predicted by ordinary modeling techniques. This illustrates the potential outcomes of adaptive niche evolution and realizedniches 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 nativeand 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 overlapin most cases, and that alien species commonly inhabit climatic niches far outside their native habitats. Climatic differences may notpresent a strong barrier to colonization, particularly if mechanisms such as ecological release, phenotypic plasticity, or rapid adaptivenicheevolution come into play.

Liu, X., X. Li, Z. Liu, et al. 2014. Congener diversity, topographic heterogeneity and humanassisted dispersal predict spread ratesof 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 bytopographic 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 amongterrestrial 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 terrestrialplants. 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 invasiveranges 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 andconservatism in introduced lineages. They find variable evidence for shifts and conservatism based on input variables. This suggeststhat habitat selection, background effects, and predictor selection may influence conclusions. The use of null models is strongly indicatedto 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 mayconfound distribution modeling for alien species. Invasive lineages often had invasive niches outside their native range, suggesting thatrealized niches might be constrained by stochastic invasion dynamics, but that fundamental niches might be much broader. Thus, nichesmay be cryptically conserved, confounding modeling attempts.




Sexton, J. P., S. B. Hangartner, and A. A. Hoffman. 2014. Genetic isolation by environment or distance: Which pattern of gene flowis most common? Evolution 68:1–15.

Adaptive nicheevolution may be constrained or promoted by gene flow from surrounding populations. The authors review the literaturefor the prevalence of isolation by distance and environment, finding strong support for isolation by environment in a majority (74.3percent) 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? OrganismsDiversity & Evolution 12:229–240.

Svensson points out that reproductive isolation is often observed in concert with niche conservatism, potentially arguing against awidespread role for ecological speciation. He reviews ways in which reproductive isolation might be decoupled from adaptive nicheevolution.

Community Perspectives

While niche evolution is commonly studied within lineages or single species, the interaction of species in a community likely represents astrong driver of evolutionary dynamics in many cases. Thus, a community perspective on adaptive nicheevolution is desirable both tounderstand how species interact and how these interactions can affect niche shifts. Ackerly 2003 tackles many of these issues in abroadscale review, outlining scenarios that may promote adaptive niche evolution. The influential review Chesson and Huntly 1997provides models for how fluctuating conditions of environmental disturbance can promote species coexistence and select for certaintraits at the population level, leading to niche shifts. Araya, et al. 2012 suggests that traits related to harsh conditions, such as droughttolerance, might be highly labile and heavily convergent in different areas where they are observed. Pillar and Duarte 2010 introduces aphylogenetic framework for assessing niche conservatism within phylogenies. Ackerly, et al. 2006 presents data suggesting that withincommunity 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 coexistlocally. Finally, Sinervo, et al. 2010 presents sobering evidence that minor changes in climate at the upper end of lizards’ thermaltolerances 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. InternationalJournal of Plant Sciences 164:S165–S184.

Ackerly outlines the multiscale nature of organismenvironment interactions, comprising plasticity, ecological sorting, and adaptiveevolution of functional traits. Three broad scenarios are outlined that may promote adaptive nicheevolution: (i) colonization of novelhabitats, (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 varydrastically among lineages and ecological scenarios. Ackerly and colleagues report evidence that withincommunity niche differencesare most likely to evolve first during adaptive radiation, followed by climaticniche differences, though high rates of climaticnichedivergence 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 communitiesexhibit phylogenetic conservatism? A test case in an endemic clade. Journal of Ecology 100:1434–1439.











Combined ecological and phylogenetic analyses suggest that crucial traits related to hydrological niche responses (such as tolerance todrought) 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 andspecies coexistence. The mechanisms discussed have farreaching implications for mechanisms underlying adaptive niche evolution atthe population level.

Pillar, V. D., and L. D. S. Duarte. 2010. A framework for metacommunity analysis of phylogenetic structure. Ecology Letters13:587–596.

The authors introduce a phylogenetic framework for analyzing community assembly patterns. In particular, they address howincorporating species’ average trait values may provide evidence for conservatism or ecological filtering along resource 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 extinctions and future risks. Under even moderate climatechange scenarios, they predict that adaptive nicheevolution will not keep pace in a significant proportion of lizard diversity, which will beat 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 bioticvariables (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 Biogeography41:999–1006.

Zink reviews mechanisms of adaptive nicheevolution influenced by ecological selection and finds a strong role therein for promotingspeciation. However, niche conservatism also affects species’ traits, and morphological divergence may often be a mechanism tofacilitate 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, etal. 2005 presents a verbal model for how high rates of shortterm niche evolution may be homogenized by geographic variability toproduce the longterm stasis often seen in the fossil record. Ricklefs 2004 discusses the integration of evolution, including niches, overtemporal and spatial scales to understand diversity. Holland and Zaffos 2011 presents a dataset for benthic marine invertebrates thatshows strong conservatism in microhabitat and abundance, but high rates of niche evolution in environmental tolerances. Pearman, etal. 2008; Lavergne, et al. 2013; and Araujo, et al. 2013 use recentoccurrence datasets to document niche shifts over short timescalesand evaluate the performance of predictive modeling approaches. Roberts and Hamann 2012 and Stigall 2012 take a similar approachwith paleontological data. Prinzing, et al. 2001 suggests that paleoadaptations to climatic instability are reflected in presentday












conservatism of niches in higher plants. Together, these studies mirror neontological data suggesting that both conservatism anddivergence are common features of ecologicalniche evolution, while the exact drivers and patterns are often stochastic.

Araujo, M. B., F. FerriYanez, F. Bozinovic, P. A. Marquet, F. Valladeres, and S. L. Chown. 2013. Heat freezes niche evolution. EcologyLetters 16:1206–1219.

Using a broad comparative dataset, the authors show that heat tolerance is generally conserved, while cold tolerance shows large shiftsamong and between lineages. They suggest that physiological limits constrain heat tolerances, while cold tolerances should be morelabile. Thus, models of extinction risk under climate change may overestimate sensitivity of coldadapted lineages, while responses oflineages near their upper thermal tolerances may be more predictable. Nicheevolution estimates should take these differences intoaccount.

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 extantpopulations are so variable? They suggest a primarily verbal model based on ecological and demographic parameters that linksgeographical 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 benthicmarine 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 bypast 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 ofclimaticniche evolution. This is true at the species level, and for deeper lineages. This may allow for better prediction of future dynamicsbased 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 andadaptive nicheevolution. They suggest that combining ecological niche modeling with phylogenetic methods can help improve theaccuracy 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 nicheevolution has been constrained. They suggest that this represents the presentday persistence of adaptations to paleoenvironmentaltrends.

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



Ricklefs gives a broad overview of the range of temporal and spatial scales that need to be considered to understand diversity. Thisincludes niche evolution at local scales and over longer time periods. He lays out eight distinct research tracks that need to beconsidered 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: Apalaeoecological 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 validatingmodels using palynological and fossil data. They find moderate performance in most cases and suggest that noanalogue climates maynot 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 highmorphological stasis through time, but high ecological stasis (niche conservatism) only during periods of gradual climatic change. Incontrast, rates of adaptive nicheevolution keep the pace with the environment during rapid climatic changes.

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