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Frontiers of Ecology

JOHN N. THOMPSON, 0. J. REICHMAN, PETER J. MORIN, GARY A. POLIS, MARY E. POWER, ROBERT W.
STERNER, CAROL A. COUCH, LAURA GOUGH, ROBERT HOLT, DAVID U. HOOPER, FELlClA KEESING, CHARLES R
LOVELL, BRUCE T. MILNE, MANUEL C. MOLLES, DAVID W. ROBERTS, AND SHARON Y. STRAUSS

E cological research is entering a new era of


integration and collaboration as we meet the challenge
of underitanding the great complexityof biological systems.
AS ECOLOGICAL RESEARCH ENTERS A NEW
ERA OF COLLABORATION, INTEGRATION,
Ecological subdisciplinesare rapidly combining and incor-
porating other biological, physical, mathematical, and soci- AND TECHNOLOGICAL SOPHISTICATION,
ologicafdisciplines.The b;rgeoning base of theoretical and
empiricalwork,made possible by new methods,technologies, FOUR FRONTIERS SEEM PARAMOUNT FOR
and funding opportunities, is providing the opportunity to UNDERSTANDING HOW BIOLOGICAL AND
reach robust answers to major ecological questions.
In December 1999 the National Science Foundation con- PHYSICAL PROCESSES INTERACT OVER
vened a white paper committee to evaluate what we know
and do not know about important ecological processes, MULTIPLE SPATIAL AND TEMPORAL SCALES
what hurdles currently hamper our progress, and what in-
tellectual and conceptual interfaces need to be encouraged.
The committee distilled the discussion into four frontiers in
research on the ecological structure of the earths biological Each of the four research frontiers takes a different ap-
diversity and the ways in which ecological processes con- proach to the overall ecological dynamics of biocomplexity,
tinuously shape that structure (i.e., ecological dynamics).This and all require integration and collaboration among those
article summarizesthe discussions of those frontiers and ex- approaches. These overlapping frontiers themselves are not
plains why they are crucial to our understanding of how eco- necessarily new. Within each frontier, however, are emerg-
logical processes shape patterns and dynamics of global ing questions and approaches that will help us understand
biocomplexity. The frontiers are how ecological processes are interconnected over multiple
1. Dynamics of coalescence in complex communities spatial and temporal scales, from local community structure
2. Evolutionary and historical determinants of ecological to global patterns.
processes: The role of ecological memory
3. Emergent properties of complex systems: Biophysical Research frontier 1: Dynamics of
constraints and evolutionary attractors coalescence in complex communities
4. Ecological topology: Defining the spatiotemporal do- We use the term community coalescence to refer to the
mains of causalityfor ecologicalstructure and processes development of complex ecological communities from a

John N. Thompson (e-mail: thompson@biology.ucsc.edu)is a professor in the Department of Ecology and Evolutionary Biology, University of California,
Santa Cruz, CA 95064. 0. J. Reichman is director of the National Center for EcologicalAnalysis Synthesis, 735 State Street, Santa Barbara, CA 93101.
Peter J. Morin is a professor in the Department of Ecology, Evolution, and Natural Resources, Cook College, Rutgers University, New Brunswick, NJ
08901-8551. Gary A. Polis (deceased) was professor and chair of the Department of Environmental Sciences and Policy, University of California, Davis,
CA 95616. Mary E. Power is a professor in the Department of Integrative Biology, University of California, Berkeley, CA 94720. Robert W. Sterner is
professor and head of the Department of Ecology Evolution and Behavior, University of Minnesota, St. Paul, MN 55108. Carol A. Couch is a senior
research scientist in the Water Resources Division of the US Geological Survey, Norcross, GA 3036@2824. Laura Gough is an assistant professor in
the Department of Biological Sciences, University of Alabama, Tuscaloosa, AL 35487-0206. Robert Holt is a professor in the Department of Ecology
and Evolutionary Biology, Natural History Museum, University of Kansas, Lawrence, KS 66045. David U. Hooper is an assistant professor in the
Department of Biology, Western Washington University, Bellingham, WA 982259160. Felicia Keesing is an assistant professor in the Department of
Biology, Bard College, Annandale, NY 12504. Charles R. Lovell is a professor in the Department of Biological Sciences, University of South Carolina,
Columbia, SC 29208. Bruce T. Milne and Manuel C. Molles are professors in the Department of Biology, University of New Mexico, Albuquerque, NM
87131. David W. Roberts is an associate professor in the Forest Resources Department, Utah State University,Logan, UT 84322. Sharon Y Strauss
is an associate professor in the section of Evolution and Ecology, University of California, Davis, CA 95616.

January 2001 / Vol. 51 No. 1 BioScience 15


Articles

regional species pool. This coalescence depends on inter- possible assembly sequencesand endpoints. The diversityof
actions among species availability,physical environment,evo- initial sequences is less problematic if many of the alternative
lutionary history, and temporal sequence of assembly. pathways tend to converge on a limited subset of endpoints.
Ecologists have made important strides in understandingthe To that end, we need to find out whether developing com-
process of community coalescence, but it will take even munities move toward single or multiplepoints or states (at-
greater integration of approaches for us to be able to con- tractors). Past some stage of assembly, community
fidentlypredct the pathways or endpoints of community as- composition may become canalized, but it may be highly
sembly (Belyeaand Lancaster 1999,Gotelli 1999,Weiher and susceptible to perturbations in the early stages of coales-
Keddy 1999). So far, we cannot predict which species are cence. Anthropogenic change may limit pathways of com-
likely to invade or to be lost from particular natural com- munity development by changing the physicochemical
munities, although some patterns are beginning to emerge environment, altering biogeochemical cycles, or changing
(Petchey et al. 1999). We also have much to learn about the genetic structure of populations. Global homogeniza-
community responses to perturbations at different stages in tion of the species pool may itself alter the pathways or end-
development. points of communityassembly,reshapingboth short-termand
Much of what we do know comes from a handful of eas- long-term successional patterns across landscapes.
ily studied systems, whereas the functioning of communi-
ties undoubtedly also depends heavily on little-studied Functional groups. Ecologists recognize that the con-
hidden players, such as microbes, fungi,and soil invertebrates cept of functional groups is a valuable tool for simplifying
(de Ruiter et al. 1995).Indeed, many attempts to create or communitycomplexityto manageablelevels.Currently,most
restore communities (e.g., freshwater wetlands or salt groups are defined in a system-specificway (Wilson 1999)
marshes) fail for reasons that remain poorly understood. based on biochemical, morphological, or trophic criteria
Until we can fill these fundamental gaps in our knowledge, (Vitousek and Hooper 1993, Hooper and Vitousek 1997,
human impacts on community patterns will remain hard to Naeem and Li 1997,Tilman et al. 1997),although some at-
predict. tempts have been made to develop general classifications
Research conducted over the last decade suggests what (Grimeet al. 1997).Each of these attempts has helped us iden-
kinds of studies are needed to fill the gaps. Progress will al- tify the advantagesand disadvantagesof different approaches.
most certainly depend on developing new ways to simplify Together, they are moving us toward the development of
the study of complexcommunities. Focusing on functional robust frameworks that work across multiple taxa and
groups or other as yet undevelopedconstructs may help. We ecological communities. Those advances should ultimately
are also aware that we need to learn far more about the sys- help us to understand how functionalgroup structure shapes
tematics of many smaller or cryptic organisms,which may assembly dynamics, as well as to evaluatehow different con-
form links that are crucial to our understanding of com- figurations of functional groups affect the development of
munity coalescence. The difficulty of linking important communities or the dynamics of ecosystems (e.g.,rates and
ecosystem functionsto key taxa and interspecific interactions trajectories of community development with and without
is a particular challenge. nitrogen-fixingspecies).
Research in five key areas, described below, could provide
important insights into the community assembly process. Hi& players. Community assemblyalmost certainlyde-
pends upon cryptic invertebrate,microbiological, and my-
Functional traits and community composition. As cological groups, which include a vast array of free-living
we study communities that are increasingly a mix of native species,parasites, and mutualistic syrnbionts.These groups and
and introduced taxa, we need to find out whether coevolved interactions have historically been ignored or unrecognized
species or local populations differ from coevolutionarilynaive because of technological limitations and biases in the train-
populations in the strengthor nature of their interactionsdur- ing of ecologists. Many microbial species cannot yet even be
ing or after communityassembly.We need to improveour abil- cultured.Nevertheless,they may play a keystone role in com-
ity to pinpoint the traits of species that affect the probability munity developmentand function,and their absence may be
of invasion or extinction within developing communities responsible for some failed attempts at community restora-
(Pimm 1989, Rejminek and Richardson 1996, Belyea and tion (Wall Freckrnan et al. 1997,Brussaard 1998).The im-
Lancaster 1999). Moreover, we need to further our under- portance of these hidden players may become apparent only
standingof how the importanceof those traits depends on ex- when they become problems,as sometimeshappens when we
isting species composition, critical thresholds (e.g., species alter community structure. Lyme disease, for example, may
richness, functional group composition), and history of a be the result of the emergence of a hidden player in response
community (Rejmanek 1989,Burke and Grime 1996,Levine to changes in communitycompositionand landscapepatterns
and D'Antonio 1999). (Joneset al. 1998).
Part of our challenge will be to determine how the struc-
Pathways toward community coalescence. For any ture of microbial diversity should be incorporated into dif-
pool of potential community members there are many ferent kinds of analyses of the local, regional, and global

16 BioScience January 2001 / Vol. 51 No. 1


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Fzgure I . 1 ne Salmon ~ t v ewithtn


r the large wilderness core of Idaho. Thefav remaining relatively pristine
landscapes worldwide will become increasingly important for testing hypotheses at the frontiers of ecology. These
unfragmented landscapes are our only way of testing hypotheses on the dynamics and trajectories of ecosystem
and evolutionary processes acting on extant species over long time scales and across broad geographic areas.
Photo: John Thompson.

dynamics of ecological processes. Traditional definitions of comprehensivetheory of communitycoalescenceif we are to


species and traditional metrics of distribution and abundance succeed in efforts to mitigate the effects of invasive species on
often make a poor fit with the structure of microbial pop- local communitiesand to make those communitiesless prone
ulations and communities. to disruption. The deliberate introduction of species, which
Our research choices have been biased not only toward takes place now in biological control efforts,will surely be less
particular taxa but also toward particular communities. We risky once we understand how to enhance the control of
have detailed community assembly information on only a species through the assembly of persistent but self-limiting
small number of well-studied kinds of communities (e.g., subwebs. Similarly, bioremediation efforts will benefit from
successional fields, temperate lakes, rocky intertidal shores). understanding how to develop persistent, self-limiting as-
Studies of underrepresented communities, taxa, and inter- semblages of species that accumulate or degrade toxic wastes.
actions will together provide important insights into gen-
eral patterns in the assembly process (e.g., roles of particular
kinds of microbial interaction or mutualistic interaction). f
Research ontier 2: Evolutionary and
historica determinants of ecological
processes
Restoration. The best measure of our understandingof any The genetic and evolutionary structure of organisms, in
complex system is whether we can reconstruct it from its combination with the recent history of environmental
parts (Jordan et al. 1987).Successful restoration of sustain- events, continually shape the dynamic patterns of commu-
able communities, and de novo creation of persistent com- nity coalescence outlined in frontier 1. Biology is a science
plex systems that provide essential ecosystem servicesin novel strongly influenced by historical events. Evolution and en-
environments, are therefore the true tests of our under- vironmental history impose "ecological memory" on com-
standingof community dynamics. Those tasks, in short, con- munities, introduce time lags in ecological processes, and
stitutethe shared frontier of community ecology and ecological constrain the trajectories of community coalescence in ways
restoration. We need to know which aspects of community that are poorly understood. Ecological memory-the result
structure are restorable once disassembled and which are of past environmental conditions and subsequent selection
not. Although our understanding of what is restorable is lirn- on populations-is encoded in the current structure of
ited, our societies are nonetheless moving apace to manipu- biological communities and reflected in the genetic struc-
late community assemblages worldwide (Simberloff et al. ture of species. It affects how communities assemble, and it
1997).We must therefore continue to work toward a more may also affect the likelihood that they can be restored once

January 2001 / Vol. 51 No. 1 BioScience 17


Articles

disassembled. We therefore need a much better under- such as coalescence theory may allow us to estimatethe length
standing of which aspects of phylogeny,ongoing evolution, of time that locally and regionally interacting species have
and recent history are most important in shaping current cooccurred (Pellmyr et al. 1998).These techniques could en-
ecological processes and patterns across landscapes. able us to determine whether populations, species, and com-
munities that share a history of cooccurrenceexhibit properties
The combined roles of phylogeny and ongoing different from those created largelyfrom recent invasions or
evolution. Although biologists commonly divide the tem- restoration efforts. They will help us determinewhether long-
poral continuum into ecologicaltime and evolutionarytime, term coevolution of taxa affects community properties such
evolution shapesecologicalprocesses across aU time intervals. as invasibility,nutrient cycling, and productivity.
The phylogenetichistoryof species creates large-scalepatterns
in the ecological relationships of taxa and combines with Scale of evolutionarydynamics. How do landscape structure
rapid evolutionover the scale of decadesto generate ongoing and genetic structure act together in shaping ecological
ecological dynamics.We need to answer six crucial questions processes? Ecologists are beginning to look hard at how the
if we areto develop a theory of ecology that takes into account spatial scale of ongoing evolution and coevolution shapes
the genetics and evolution of organisms. ecological dynamics. Species are groups of geneticallydiffer-
entiated populations connected by various levels of gene
Phylogeneticstructureof ecologicalprocesses. How does the flow. Adaptations developed within local communities can
shared phylogenetic history of species shape ecological spread to populations in other communities,therebyaltering
processes? Because of shared phylogenetic history, species local ecological dynamics. Natural and anthropogenic frag-
cannotbe treated as independentunits in their ecologicalroles. mentation of populations restricts the exchange of individ-
We know, for example,that ecological specialization is phy- uals among communities both historically and currently.We
logenetically constrained (Futuyma and Mitter 1996,Webb arejust beginningto understand how the landscape structure
2000).A few studies have analyzed how shared species' traits of fragmentation interacts with the genetic configuration of
and historicalbiogeography combine to constrain and shape species and ongoing evolution to shape ecologicalprocesses
community and ecosystem structures. For example,analysis at local, regional, and continental scales.We need a refined the-
of the phylogeny of Caribbean anole lizards casts doubt on the ory of spatiallystructured evolutionarydynamicsto assess the
overall applicabilityof a widely cited ecological theory, the overall effects of increased anthropogenic fragmentation on
taxon cycle (Butler and Losos 1997).Recent analyses of rain communities.
forest in Borneo indicatethat more closely related specieswere
found in the same microhabitats more often than couldbe ex- Genetic diversityandecologicaldynamics. How does genetic
plained by chance alone (Webb2000).We need similar analy- diversitywithin species shape the temporal dynamics of pop-
ses across taxa and ecosystemsto furtherintegrate phylogenetic ulations,trophic interactions,and energy flow in ecosystems?
nonindependence into community and ecosystem ecology. By incorporatingthe effects of genetic diversityper se into our
models of the temporal dynamics of ecosystem structure
Rapid evolutionand ecologicaldynamics. To what extent are and function,we may better understand and predict ecolog-
succession, population dynamics, and ecosystem dynam- ical systems.The study of plant hybrid zones has given us a
ics-historically consideredto be solelyecological processes- glimpse of the potentially far-reaching effects of the structure
governed by rapid evolutionary change in species and their of geneticdiversityon community dynamics (Whitham et al.
interactions?There are dozens of examples in which ecolog- 1999).Particular genotypes or hybrids of some species are
ically important traits of species are known to have evolved much more vulnerable to herbivores or are more productive
during the past century (Thompson 1998,1999).Rapid evo- than other genotypes or either of the parental species.
lution of bill morphology in Galapagos finches,wing color in Moreover, some hybrids and genotypes support greater di-
peppered moths, and pesticide resistance in many taxa are well- versity and biomass of insect herbivores, birds, and other
known examplesthat highlightthe speed with which natural species, thereby becoming local foci of diversity and sup-
populations can evolve when subjected to environmental porting distinct food web interactions.
change. These evolutionary changes have the potential to
ripple throughout communities.For example,the recently dis- Genomicsof ecologicaldynamics. How do genomic con-
covered rapid evolution of Daphnia in response to pollu- tent and patterns of gene expression shape species distrib-
tion in Lake Constance may change phytoplankton dynamics utions, species interactions, and responses to changing
in ways that are important to community function (Hairston environments?We are just starting to evaluate the ways in
et al. 1999). which chromosome number shapes large-scale patterns in
community structure and dynamics. From a small number
Coevolutionand ecologicaldynamics. How does the history of studies we know that chromosome number is tied to
of species cooccurrenceshape ecological processes,commu- ecologically important attributes of taxa, such as species
nity function, and community stabilityand invasibility?Use distributions (e.g.,the proportion of polyploid plant species
of molecular markers coupled with new analytical techniques increases with latitude), invasiveness, and susceptibility of

18 BioScience January 2001 / Vol. 51 No. I


Figure 2. Thefiagmented wet tropics of northern Queensland, Australia, north of Cairns. Similar landscapes now
characterize most terrestrial environments worldwide, illustrating the need for understanding the dynamics of population,
communiry, ecosystem, and evolutionary processes in environments whose boundaries are sharp and constantly changing
across large spatial and temporal scales. Photo: John Thompson.

plants to herbivores (e.g., moths on saxifrages) (Rosenzweig working on particular taxa or communities. Nevertheless,
1995, Thompson et al. 1997). Thus, the evolutionary ge- we are only in the early stages of developing a general body
nomics of component species may have strong impacts on of theory on how past periodic or pulsed productivity affects
the ecological functioning of communities. the dynamics of populations, interactions between resources
and consumers, food webs, communities, and ecosystems.
The role of environmental history and temporal We need to continue to work toward a synthetic frame-
variability. Ecological research needs to build on past ef- work for explainirlg how temporally variable productivity
forts to integrate processes acting across different time scales. influences food web processes, community dynamics, and
All environmentsvary through time at scales from very short ecosystem function. For example, masting events and.peri-
(hours, days) to long (decades,centuries,millennia).Because odic emergence of insects (e.g., cicadas) can greatly alter
of this temporal variability, we often cannot understand com- species interactions and community dynamics for years
munities without understanding past events or processes. (Joneset al. 1998).Long periods between pulses of high pro-
We know that three components of environmentalhistory and ductivity reduce consumer abundance, consequently im-
temporal variability appear to be particularly important in peding resource suppression at the next productive period.
shaping community dynamics: variability in productivity This process appears to underlie the evolution of masting in
and the input of resources; variability in species composition, bamboos and trees and synchronous reproduction in insects
abundance, and interaction strength; and,variabilityin key abi- and ungulates (Janzen 1976).
otic events. The impact of these factors can change over dif- In general, periodic pulses of productivity (e.g., good
ferent time scales, shaping species composition,abundance, versus bad years or seasons) may produce resource reserves
and food web interactions in the short term (e.g., via recent that last for varying periods before they are depleted, alter-
invasion), medium term (range expansion and contraction), ing food web dynamics long after the productivity pulse
and long term (historical biogeography). (Noy-Meir 1974,Sears et al. in press). For example, El Niiio
rains (or La Niiia drought) may stimulate or depress
Historical variability in productivity or resource input. terrestrial productivity, thereby allowing subsequently high
Resources vary in availability over time in all communities or low abundances of consumers and altering interactions
through abiotic (e.g., wet versus dry seasons; El Nifio versus between resources and consumers (Polis et al. 1997). On
La Niiia years) and biotic influences (mast years in long- smaller spatial scales, the pattern of local competitive
lived plants, pig wastes flowing into watersheds) (Polis et al. outcomes in forests may depend on the pattern of tree falls,
1996). Such variation has been characterized by ecologists old allelopathic footprints, and past local patterns of

January 2001 / Vol.51 No. I BioScience 19


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nutrient depletion or regeneration, each of which may pro- evolution by natural selection can successfully predict the
duce long-lasting changes in local soil chemistry. composition, structure, and functioning of ecosystems.If so,
evolutionary first principles could enhance ecosystem study.
Historicalvariabiityin speciescomposition,abundance, and Reciprocally,advances in studies of matter and energy flow
interactionstrength.We are beginning to disentanglethe ways can provide the information necessary to realistically assess
in which past community configurations shape current biophysical constraints on evolutionary change.
processes. We know, for example, that the effects of intro- If we imagine a community as a set of species plotted in
ductionsof new species into communities can depend on past multidimensional phenotypic space, then enclosing this
presence of particular taxa. Introduction of cows into inter- space is a set of boundaries and constraints imposed by the
mountain western North America led to effects on grassland principles of energy and thermodynamics. Matter devoted
and steppe communitiesthat were far differentfrom those re- to one use is unavailable for other uses, and these con-
sulting from similarintroductions east of the Rockies. These straints generate evolutionary tradeoffs. Selection maxi-
differing impacts came about in part from differencesin the mizes fitness within those constraints. In the terminology of
history of associationof these grasslands with bison and the dynamic systems theory, the result is an attractor, a condi-
selection these large mammals imposed on plant taxa. In ad- tion (such as a set of phenotypes) toward which a system
dition, short-term irruptions of species can produce long- (such as a community) is drawn over (evolutionary) time.
lasting effects. The irruption of rinderpest in some African As each organism responds to its environment in a man-
grasslands resulted in a singleflush of recruitment of acacias ner prescribed by its genome and the specifics of its bio-
duringthis century.The acaciasare still extant,yet current eco- physical and biotic environment, it simultaneouslymodifies
logical conditions or their recent life history statistics (e.g., that environment. The result is a coupled, complex, dy-
measurement of intrinsic rate of natural increase) do not namic system of organism and environment, wherein nat-
account for their presence. ural selection optimizes the fitness of populations amid a
continuallychanging,bioticallydriven environment. If a sin-
Historical variability in key abiotic events. Erratic large- gle species of plant evolves to have larger leaf area, a host of
scale disturbanceregimes can leave profound ecologicallega- allometric tradeoffswithin that speciescome into play, such
cies that may propagate historical echoes. Volcanoes, as corresponding increases in sapwood conducting volume.
hurricanes, El Nifio-southern oscillation events, or inter- The cumulative effect of these changes may be an alter-
decadal oscillations all reshape biological communities in ation in how resources are used within a community.
ways that have long-lastingeffects.For example,interdecadal Material and energy fluxes for competing species and con-
changes in anchor ice play a major role in altering the abun- sumers of that plant may in turn also be affected.
dance, composition, and interaction dynamics of the entire An example of an attractor resulting from biophysical
bottom community in the Antarctic (Dayton 1989).We must properties is the 0.75 power law for metabolism in en-
striveto develop a fuller appreciation of when and how these dothermic animals, which states that whole-body meta-
major periodic environmental events are important in shap- bolic rate scaleswith body mass to the 0.75 power.Although
ing global and regional patterns in ecological processes. there has been turnover of species along this curve over
time, the physical basis of this power law ensures that the
Research frontier 3: Emergent properties curve itself has remained fixed. Such an evolutionary at-
of complex systems tractor works through a combination of physical principles
The evolutionary and historical determinants of ecological and evolutionary optimization. Phenotypes that deviate
processes outlined in research frontier 2 are in turn shaped from this attractor are at a significantdisadvantagecompared
in large part by the laws of physical science. Hence, the with those near the attractor.
third frontier is to understand how the joint evolutionary The frequency distributions of body mass in mammals,
forces acting on an entirespecies assemblageresult from con- which demonstrate a unimodal relationship, maybe another
straints imposed by biophysical processes. example. Brown (1995) suggested that modal mass corre-
Phenotypic traits are products of adaptive evolution sponds to an evolutionary optimum (attractor) for a given
operating within bounds set by physical and chemical body plan. Body mass may be governed by a tradeoff between
constraints. Important progress in determining how these resource harvest ability and bioenergetic patterns deter-
constraints affect the morphology and behavior of indi- mining the rate at which resources can be turned into
vidual species has been registered in the subdisciplines of reproduction. Deviations from the optimum may be driven
physiological and functional ecology. Today's research by niche partitioning. Ritchie and Olff (1999) used spatial
frontier is the expansion of this viewpoint to collections of scaling laws to describe how species of different sizes find
species and communities and ecosystems.We need to know food in patches of varying size and resource concentration.
how specific evolutionary responses at the species level They derived a mathematical rule for the species that share
produce the collective morphology (physiognomy) and resources and used the rule to develop a predictive theory
behavior (ecosystem function) of ecosystems.One central of the relationship between productivity and diversity.
question is whether first principles of physics, chemistry,and Spatial scaling laws, they argued, provide potentially

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unifying first principles that may explain many important less direct means of testing mechanistic hypotheses will be
patterns in species diversity. necessary.
Biochemical constraints may also result in an evolution-
ary attractor on life history evolution. Evidence for such con- New techniques. New approaches in the study of structural
straints is accumulating through studies of ecological and functional genomics can further this highly integrated
stoichiometry, which considers patterns in element con- view of evolution and ecosystems. A community is com-
centration among different species (Sterner 1995). Recent posed of suites of genes that are collectedinto epistatic groups,
studies have indicated that rapid growth in biomass re- these into genomes, and finally genomes into communities.
quires an element composition high in phosphorus (P), a Breakthroughs in gene sequencing and in measuring pat-
result of the element composition of anabolic biochemical terns of gene expression should make it increasingly practi-
machinery-in particular, ribosomes (Elser et al. 1996). cable for ecologists to study the relationships between
Thus, life history evolution is constrained in a phenotypic biophysical constraints and evolutionary attractors. For ex-
space that includes phosphorus as one of its axes: High ample, studies have described the patterns of expression of
growth rate depends upon high P content. Another stoi- 6347 genesin mice raised under caloric restriction (Lee et al.
chiometric example is the evolution of structural material, 1999).Caloric restriction retarded aging by causing a meta-
such as wood or bones, in which different biochemical so- bolic shift toward increased protein turnover and decreased
lutions to the same or similar structural problems have macromolecular damage. These experiments provide a
been achieved. Hence, we see alternativechemical signatures glimpse of a single genome's response to the changed avail-
in living organisms. The larger the organism, the greater the ability of energy. They suggest intriguing links between ge-
reliance on carbon (C) (for wood) or calcium (Ca) and nomics and energy and nitrogen fluxes.We need to explore
phosphorus (for bone), generating an attractor in a phe- how species' genomes respond to each other as material and
notypic space that includes elements such as C, Ca, or P as energy availability change. Such studies can be viewed as
well as organism body size. ecological,community, or even ecosystem genomics.

Specific research questions. We need to answer several Research frontier 4: Ecological topology
questions about the physical basis of evolutionaryattractors. Frontiers 1,2, and 3 require that we reevaluate the scale of
Among them are these: ecological processes and the analytical methods by which we
How are species arranged as collections in phenotypic study them.We need to develop a way of evaluating how our
space bounded by biophysical constraints? scales of information (scope and resolution in space and
How do adaptive solutions vary as species diversity time) affect our understanding of ecological processes-that
varies within those constraints? is, an ecological topology. We can think of this ecological
Can different,coherent sets of community members be- topology as a study of the domains of causality. For in-
come established as multiple stable states, or does stance,what are the factors that control rates of primary pro-
each situation engender a specificecological and evo- duction by native plants on ancient lava surfaces in Hawaii?
lutionary solution (based on ecologicalstoichiometry The most important controls may be local (e.g.,competition
or similar constraints)? among neighboring plants), regional (e.g.,grazing by wan-
dering goats introduced by Europeans centuries earlier and
What effect does fitness maximization of individuals
sustained by productivity of the larger surrounding land-
have on the collective behavior of all species and on
scape), or even global (set by the levels of phosphorus in
the fluxes of matter and energy within these collec- eolian dust from Central Asia, mobilized during the last
tions?
glacial period) (Jackson et al. 1971, Chadwick et al. 1999).
Can first principlesbe used to predict the statisticaldis- The appropriate domains of causality in many ecological
tribution of species traits, not just a single optimum? studies could extend far beyond previously assumed spatial
Can that distribution be extrapolated to predictions and temporal bounds.
or estimates of diversity? We therefore need a unified understanding of the spatial
We cannot begin to cross this frontier successfullyunless and temporal context of ecological interactions, such as the
we find the means to perform rigorous experimental tests magnitude and importance of cross-habitatfluxes of matter,
or find other ways to test hypotheses. Multispecies correla- energy, and information, sometimes acting over very large
tions of phenotypic traits will be relatively easy to find, but scales.Even our understanding of fundamental matters such
testing among competing hypotheses for these broad-scale as the structure of food webs generally lacks a spatiotemporal
patterns will require considerable ingenuity. Experimental context, strongly limiting our ability to explain their origin,
tests of some evolutionary hypotheses are impossible for all maintenance, or consequences.We suggest building an eco-
but the smallest (and most quickly reproducing) organ- logical topology that addresses these needs. Doing so re-
isms, but microbial taxa make excellent experimental tools quires characterizing patterns of movement of individuals
for answering some of these questions. For most other taxa, (Holt and Gaines 1992, Holt 1993) and flow of materials

January 2001 / Vol. 51 No. 1 BioScience 21


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Figure 3. Coalescence of a new, complex plant community of native and introduced species at Tejon Pass, California. The
community includes a mix of annual Mediterranean grass species and a diverse array of native annual wildfowers. Before
invasion by the introduced grasses, the community was probably dominated by perennial, tussock-forminggrasses, along
with annualgrasses andforbs. Photo: Courtesy of Frank Davis, University of California,Santa Barbara.
(Power and Rainey in press) across space and through time research in scientific fields outside ecology, and some of the
(Polis et al. 1997).This frontier is consistent with todafs great methods of those fields could inform us about the bounds of
need for scaling up from local processes to much larger ecological systems.
ones, including the planetary scale. Global change is one of
the most pressing issues in ecology today. Understanding the Changing the bounds. How will ecological communities
topology of causation of ecological patterns and processes change when their bounds change? If there are natural con-
could provide a formal structure for linking studies at the straints on the bounds of domains of ecological processes, will
local scale to larger ones. truncating or stretchingthem change system behavior,stability,
We already have some tools, and could soon develop oth- or sustainability?This is a crucial consideration,given human
ers, to investigate and characterize the bounds of such domination and rearrangement of ecosystems. For example,
spatiotemporaldomains. Genetic and isotopic tracers for fol- the California water system has been called the most massive
lowing the movement of matter and organisms within and rearrangement of nature ever attempted. Interbasin water
across ecosystem boundaries are becoming more accessible. transfers are now common throughout many arid regions of
We have new satellite and ground-based technologies that the world, depleting or artificially augmenting local ecosys-
expand our ability to capture and analyze spatial data. With tems and groundwater systems. Release of carboniferousfos-
these conceptual and technical tools, we can address a broad sil fuels back into the active global carbon cycle is another
range of important ecological problems, such as those that example of human domain stretching along a deep tempo-
follow. ral axis. Our global homogenization of species could also be
considered in this light. Humans, however, do not always
Choosing the right spatial and temporal context. stretch causality domains. Sometimes we greatly contract
How do different spatial and temporal domains of causality them, as when we replace wild populations of salmonids,
combine to produce local community patterns and processes? which harvest huge areas of ocean production, with cage-
Moreover, how far back in time or out into the spatial land- cultured fish that feed only in the mouths of estuaries. It is
scape must we probe to understand local phenomena or in- worth exploring whether causalitydomain theory can enhance
teractions?We need to h d out just how far we should expand our ability to predict the longer-term consequences of eco-
the domains of our search if we are to interpret ecologicalpat- logical distortions of various types and magnitudes.
terns and processes correctly. Focus on these ecological topologies will require a change
in how ecologists typically think of ecosystems. Conventionally,
Identifiing the rules. What rules govern the origin and the term ecosystem refers to a particular contiguous unit of the
maintenance of ecological topologies?Clearly, some general landscape. The system has inputs and outputs and internal dy-
rules govern the geometries of circulatory systems or the namics. Most management practices similarly focus on eco-
drainage networks of watersheds. This is an area of active logical processes and patterns defined by such habitat units.

22 BioScience January 2001 / Vol. 51 No.1


Nevertheless, many, if not most, ecological processes can be developing ecological theory that builds on the structure of
envisaged as caused by a set of rules, each of which operates empirical results in ecology.
over different spatial and temporal scales. Understanding
those rules is among our great current challenges. Access to research and monitoring databases.We also
need ecologists trained to manage large databases, who can
Meeting the challenge organize the storehouse of past ecological data, mine it for new
The challenges ahead will require new ways of working to- results, and make it accessible to others. Lack of ready access
gether, new tools, and greater access to an ever growing to the already available mass of ecological data is a major hur-
number of databases. dle in ecological analysis.The experience of dozens of work-
ing groups at the National Center for Ecological Analysis
Collaboration and integration. No single scientist has and Synthesis in Santa Barbara, California, who are attempting
depth enough across all the physical and biological sciences to summarize and compare data collected over decades for a
to address the questions at the frontiers. Therefore, we must wide variety of purposes, confirms this need. Hence, the need
foster research collaborations among scientists who are for stronger links between empirical ecologists and database
trained to understand the language of one another. Meeting managers will continue to grow as new ecological data are
that challenge will require renewed effort to make sure that gathered for ever-changing questions.
ecologists are well-grounded in earth sciences and mathe- The existing storehouse of ecological data includes not
matics, and physical scientists solidly grounded in the the- only the results of individual studies but also the many
ory of ecology. databases that come from environmental monitoring efforts.
In addition, stronger links between ecology and evolu- Maintenance of these monitoring efforts is important if we
tionary biology must be forged across all the frontiers. are to understand how past environmental events shape
Despite the formation of departments of ecology and evo- current ecological processes. Monitoring efforts, however, are
lutionary biology in many universities during the last 30 useful for ecological analyses only if their results are readi-
years, there remains a major gap between these disciplines. ly accessible to ecologists.
Relatively few ecologists have ever had a formal course in evo- Moreover, to work with diverse data sets, collection meth-
ods and data entry methods must be standardized wherever
lutionary processes, and very few have ever had advanced
possible. Techniques continue to change, and different meth-
training in evolutionary theory and methods. Similarly,few
ods are needed for different ecological situations.
evolutionary biologists have much advanced training in
Nevertheless,we need to take a fresh look at some of the com-
current ecological theory and methods. Incorporating an
mon methods used in studies within subdisciplines and at-
evolutionary framework into ecological research requires that
tempt standardization.
we train scientists to be well versed in both ecological and
evolutionary theory.
Analogously, there is a need to strengthen the links be-
Conclusion
Ecological research is undoubtedly entering a new era, build-
tween ecosystem and community ecology. Understanding ing on a firm base of advancements in our ecological knowl-
how biological communities shape, and are shaped by, phys- edge achieved in recent years. Each of the frontiers of ecology
ical environmental processes will transpire as the discipli- requires increased collaboration and technology if we are to
nary barriers between ecosystem ecology and community understand how the complexity of ecological processes con-
ecology break down. tinues to reshape the earth. Most of the unresolved challenges
The need for stronger links between systematics and ecol- in ecology arise from incomplete understanding of how bi-
ogy is evident for all taxa, but it is especially critical for mi- ological and physical processes interact over multiple spa-
crobial groups. We need to turn out a generation of biologists tial and temporal scales. These challenges permeate the four
who are well grounded both in microbial ecology and mi- interrelated research frontiers we have identified. Crossing
crobial systematics. These will be scientists capable of grap- the frontiers demands strong cross-disciplinary training
pling with the complex structure of microbial taxa in an and collaboration, coordination of observational and ex-
ecological context. We are only in the early stages of what will perimental approaches across large scales, incorporation
become a major field of study: how microbial diversity con- of new technologies, and greater access to the growing data-
tributes to the dynamics of biocomplexity. The same needs base of ecological results. Developing the means to cross the
hold for studies of the systematics and ecology of cryptic in- frontiers is essential if the field of ecology is to move forward
vertebrates and fungi. as a predictive science.

Analytical tools. The mathematical and statistical Acknowledgments


constructs used in ecological research come mostly from We are grateful to Scott Collins, William Michener, and
linear theory. In contrast, many ecological processes are gov- Margaret Palmer, program officers for the National Science
erned by nonlinear dynamics, threshold effects, and more Foundation, for initiating this effort to assess the frontiers
complex relational structures. We must continue to work on of ecology. We are also grateful to three anonymous

January 2001 / Vol. 51 No. 1 . BioScience 23


Articles iFd l

Petchey OL, McPhearson PT, Casey TM, Morin PJ. 1999. Environmental
reviewers for very helpful and thoughtful reviews. This
warming alters food web structure and ecosystem function. Nature 402:
work was supported by NSF grant 97-07781 to J. N. T. 69-72.
Pimm SL. 1989. Theories of predicting success and impact of introduced
References cited species. Pages 351-367 in Drake JA, DiCastri F, Groves RH, Kruger FJ,
Belyea LR, Lancaster J.1999.Assembly rules within a contingentecology. Oikos Mooney HA, Rejmanek M, Williamson MH, eds. Biological Invasions:
86: 402416. A Global Perspective. Chichester (UK): JohnWiley & Sons.
Brown JH. 1995.Macroecology. Chicago: University of Chicago Press. Polis GA, Holt RD, Menge BA,Winemiller K. 1996.Time, space and life his-
Brussaard L. 1998. Soil fauna, guilds, functional groups and ecosystem tory: Influences on food webs. Pages 435460 in Polis GA, Winemiller
processes. Applied Soil Ecology 9: 123-135. K, eds. Food Webs: Integration of Patterns and Dynamics. New York:
Burke MJW, Grime JP. 1996.An experimental study of plant community in- Chapman and Hall.
vasibility. Ecology 77: 776-790. Polis GA,Anderson W, Holt RD. 1997.Towards an integration of landscape
Butler MA, Losos JB. 1997.Testing for unequal amounts of evolution in a con- ecology and food web ecology: The dynamicsof spatiallysubsidized food
tinuous character on different branches of a phylogenetic tree using webs. Annual Review of Ecology and Systematics 28: 289-316.
linear and squared-change parsimony. Evolution 51: 1623-1635. Polis GA, Hurd SD, JacksonCT, Sanchez-Pifiero F. 1997. El Nifio effects on
Chadwick OA, Derry LA, Vitousek PM, Huebert BJ, Hedin LO. 1999. the dynamics and control of a terrestrial island ecosystem in the Gulf of
Changingsources of nutrients during four million years of ecosystemde- California. Ecology 78: 1884-1897.
velopment. Nature 397: 491-497. Power ME, RaineyWE. In press. Food webs and resource sheds:Towards spa-
Dayton P. 1989.Interdecadal variation in an Antarctic sponge and its preda- tially delimiting trophic interactions. In Hutchings MJ, John EA, eds.
tors from oceanographic climate shifts. Science 245: 1484-1486. Ecological Consequences of Habitat Heterogeneity.London: Blackwell
de Ruiter PC, Neutel A, Moore JC. 1995.Energetics,patterns of interaction Scientific.
strengths, and stability in real ecosystems. Science 269: 1257-1260. Rejmanek M. 1989. Invasibility of plant communities. Pages 369-388 in
Elser JJ,Dobberfuhl D, MacKay NA, Schampel JH. 1996.Organism size, life Drake JA,DiCastri F, Groves RH, Kruger F J,Mooney HA, Rejmanek M,
history, and N:P stoichiometry: Towards a unified view of cellular and Williamson MH, eds. Biological Invasions: A Global Perspective.
ecosystem processes. BioScience 46: 674484. Chichester (UK): JohnWiley & Sons.
Futuyma DJ, Mitter C. 1996. Insect-plant interactions: The evolution of Rejmanek M, Richardson DM. 1996. What attributes make some plant
component communities.Philosophical Transactions of the Royal Society species more invasive?Ecology 77: 1655-1661.
of London B 351: 1361-1366. Ritchie M, Olff H. 1999. Spatial scalinglaws yield a synthetic theory of bio-
Gotelli NJ. 1999. How do communities come together? Science 286: diversity. Nature 400: 557-560.
1684-1685. Rosenzweig ML. 1995.Speciesdiversityin space and time. Cambridge (UK):
Grime JP,et al. 1997.Integrated screeningvalidatesprimary axes of special- Cambridge University Press.
ization in plants. Oikos 79: 259-281. SearsA, Holt RD, Polis GA. In press.The effectsof temporal variabilityin pro-
Hairston NG Jr,Lampert W, Caceres CE, Holtmeier CL, Weider LJ,Gaedke ductivity on food webs and communities. In Polis GA, Power ME,
U, Fischer JM,Fox JA,Post DM. 1999.Rapid evolution revealed by dor- Huxelm G, eds. Food Webs at the Landscape Level. Chicago: University
mant eggs. Nature 401: 446. of Chicago Press.
Holt RD. 1993.Ecology at the mesoscale:The influenceof regional processes Simberloff D, Schmitz DC, Brown TC, eds. 1997. Strangers in Paradise:
on localcommunities. Pages 77-88 in Ricklefs R, SchluterD, eds. Species Impact of Managementof NonindigenousSpeciesin Florida.Washington
Diversity in Ecological Communities. Chicago: University of Chicago (DC):Island Press.
Press. Sterner RW. 1995.Elemental stoichiometry of species in ecosystems. Pages
Holt RD, Gaines MS. 1992. Analysis of adaptation in heterogeneous land- 240-252 in JonesCG, Lawton JH,eds. Linking Species and Ecosystems.
scapes:Implications for the evolution of fundamentalniches. Evolutionary New York: Chapman and Hall.
Ecology 6: 433-447 Thompson JN. 1998. Rapid evolution as an ecological process. Trends in
Hooper DU, Vitousek PM. 1997.The effects of plant composition and di- Ecology and Evolution 13: 329-332.
versity on ecosystemprocesses. Science 277: 1302-1305. . 1999.The evolution of species interactions.Science284: 2116-2118.
Jackson ML, Levelt TVM, Syers JK, Rex RW, Clayton RN, Sherman GD, Thompson JN, Cunningham BM, Segraves KA, Althoff DM, Wagner D.
Uehara G. 1971. Geomorphological relationships of tropospherically 1997.Plant polyploidy and insect/plant interactions.American Naturahst
derived quartz in soils of the Hawaiian Islands. Soil Science Society of 150: 730-743.
America Journal 35: 515-525. Tilman D, Knops J,Wedin D, Reich P, Ritchie M, Siemann E. 1997.The in-
Janzen DH. 1976.Why bamboos wait so long to flower.Annual Review of fluence of functional diversity and composition on ecosystem processes.
Ecology and Systematics 7: 347-91. Science 277: 1300-1302.
Jones CG, Ostfeld RS, Richard MP, Schauber EM,Wolff J0.1998. Chain re- Vitousek PM, Hooper DU. 1993. Biologicaldiversity and terrestrial ecosys-
actions linking acorns to gypsy moth outbreaks and Lyme disease risk. tem biogeochemistry. Pages 3-14 in Schulze E-D, Mooney HA, eds.
Science 279: 1023-1026. Biodiversity and Ecosystem Function. Berlin: Springer-Verlag.
Jordan WR 111,Gilpin ME, Aber JD,eds. 1987.RestorationEcology.Cambridge Wall Freckman D, Blackburn TH, Brussaard L, Hutchings P, Palmer MA,
(UK):Cambridge University Press. SnelgrovePVR. 1997.Linking biodiversityand ecosystem functioning of
Lee C-K, Klopp RG, Weindruch R, Prolla TA. 1999.Gene expression profile soils and sediments. Ambio 26: 556-562.
of agingand its retardation by caloricrestriction.Science285: 239g2393. Webb CO. 2000. Exploring the phylogenetic structure of ecological com-
Levine JM,D'Antonio CM. 1999.Elton revisited: A review of evidencelink- munities: An example from rain forest trees. American Naturalist 156:
ing diversityand invasibility. Oikos 87: 15-26. 145-156.
Naeem S, Li S. 1997.Biodiversity enhances ecosystem reliability.Nature 390: Weiher E, Keddy P, eds. 1999. Ecological Assembly Rules: Perspectives,
507-509. Advances, Retreats.New York: Cambridge University Press.
Noy-Meir I. 1974.Desert ecosystems: Higher trophic levels.Annual Review Whitham TG, Martinsen GD, Floate KD, Dungey HS, Potts BM, Keim P. 1999.
of Ecology and Systematics 5: 195-214. Plant hybrid zones affect biodiversity:Tools for a genetic-based under-
Pellmyr 0 , Leebens-Mack J,Thompson JN. 1998.Herbivores and molecu- standing of community structure. Ecology 80: 41W28.
lar clocks as tools in plant biogeography. Biological Journal of the Wilson JB. 1999. Guilds, functional types and ecological groups. Oikos 86:
Linnean Society63: 367-378. 507-522.

24 BioScience January 2001 /Val. 51 No. 1

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