| Peter Chesson Post-doctoral Associates Andrea Mathias Jessica Kuang Danielle Ignace Graduate Students Galen Holt Adam Miller Maureen Ryan Recent Graduates Megan Donahue Anna Sears Jessica Kuang Undergraduate Students Tara Woodcock Friends of the Lab Charlotte Lee Brett Melbourne Kendi Davies Scott Stark Graduation Photos ![]() |
Our study systems Some of us are fond of studying communities of annual plants. We find these in the deserts of Arizona and Eyre Pennisula (South Australia), in the vernal pools of the Central Valley of California, and in the coastal swales near Bodega Bay. We also study perennial herbs in the understorey of Eucalypt forests in Australia. Some of us are aquatic in orientation studying amphibia in ponds and streams or marine intertidal invertebrates. There are lots of systems that we think about
but do not collect data on ourselves, at least not right now. These
include coral
reef fish communities, tropical forests, and host-parasitoid systems. For studies of natural systems, we are developing new classes of quantitative tools to test coexistence mechanisms and hypotheses about dynamics of populations and communities subject to spatial and temporal variation. Then we go out into the field to count, measure, and manipulate. For theoretical studies, we use math and
computers. We are fond
of probability theory and statistical theory. We also get into
differential equations
and Fourier transforms.
|
Our questions Ecological
diversity
and its maintenance General
testable community theory Variation and
scale ![]() |

| Diversity
maintenance mechanisms Classification
of Mechanisms |
Dynamical
Concepts |
Classification of Ecological Variation |
| Community
dynamics in time and space Natural communities inhabitat a world of great complexity in time and space. The physical environment has many spatial complexities in its topography, hydrology and soils. And it changes over time: organisms experience varying weather and climatic changes on many scales interacting in a variety of ways with spatial environmental varation. Various processes, not just the physical environment, but also interactions between organisms, cause their densities to vary greatly in space and time too. It has often been hypothesized that these different kinds of variation have important roles in diversity maintenance. But what are those roles? Development of mathematical models with such complexities is not easy, and so it is perhaps not surprising that much mathematical theory, even if it addresses density variation in time or space, rarely includes variation in the physical environment. The theory coming from this lab is different. We have embraced environmental complexity, and gathered the mathematical tools necessary to study it. The result is a comprehensive multiscaled theory of diversity maintenance that includes the standard equilibrium theory as a special case. This work has led to the recognition of several new species coexistence mechanisms including the storage effect, nonlinear competitive variance, and fitness-density covariance. Multitrophic diversity maintenance The study of diversity maintenance mechanisms has traditionally focused on competition, with predation mostly seen as modifying what competition does. However, there is growing evidence that predation and competition can have very similar effects on the maintenance of species diversity. This similarity of action relies on the observation by Bob Holt (University of Florida) that both predation and competition can lead to mutually negative indirect interactions between species. Of most importance, such mutually negative indirect interactions apply not just between species but also to different individuals within the same species. Theoretical consequences of these facts mean that predation and competition each have the potential to limit diversity or promote diversity. For both predation and competition it all depends on the extent to which within species interactions outway between species interactions. When predation and competition have similar strengths for a given set of interacting species, the overall outcome for diversity |
maintenance may
represent a compromise between the separate tendencies of predation and
competition in the given circumstances. In other cases, new mechanisms
promoting diversity arise from the interaction between predation and
competiton. If one mechanism is much stronger than the other, the
tendency of the stronger mechanism prevails. How do species coexistence mechanisms work? And how do we know how strong they are? Our lab has developed measures of mechanism strength based on how much a mechanism increases the rate of recovery of a species from low density. In nature, species always fluctuate, and if they are to remain in a community, they must recover from their low excursions. Focusing on these recovery rates, we have found ways of quantifying mechanism strength in terms of the |
functional components that define the mechanism. This has led not only to a big increase in understanding of mechanism functioning, but also powerful new ways of testing mechanisms. Multiple
mechanisms, mulitple scales |
Peter Chesson
Community
Processes
in Variable Environments
The natural world is inherently variable. The
physical environment varies on all scales and so do the organisms
inhabiting it. Organisms
are adapted to this variability, not merely hedging against uncertainty
but
taking advantage of opportunities that variability brings. My main
interest is how the adaptation of organisms to variability promotes
species diversity and affects ecosystem functioning. I do this
primarily theoretically by developing
mathematical models using probability theory and the methods of
theoretical
statistics. Recent research topics have involved neighborhood
competition
models, community assembly, invasion resistance, and the various
manifestations
of temporal and spatial niches, also known as the storage effect and
relative
nonlinearity of competiton. In addition, I am involved with field
projects
on annual plants and herbaceous perennial plants. Other systems that
are
foci of theorizing are coral reef communities, tropical forests, and
communities
in mediterranean climates.

Some recent publications
Chesson,
P. 2008. Quantifying and testing species coexistence
mechanisms. Pp 119 – 164 in Valladares, F., Camacho, A., Elosegui, A.,
Estrada, M., Gracia, C., Senar, J.C., & Gili, J.M., ed. “Unity in
Diversity: Reflections on Ecology after the Legacy of Ramon Margalef.”
Andrea Mathias
Non-equilibrium dynamics and species coexistence
Non-equilibrium dynamics of natural ecosystems have a major role in generating and maintaining biodiversity. The main objective of my research is the mechanistic understanding of this role in both ecological and evolutionary context. In particular I am interested in life history evolution with frequency dependent selection in temporally fluctuating and spatially heterogeneous environments.
Danielle Ignace
Physiological Ecology of Plant Invasions
I am studying the causes of the dramatic
irruption of the Eurasian invasive plant, Erodium cicutarium, in the San
Simon Valley of southeastern Arizona. It many areas, it is now
95% or more of
the winter annual plant biomass, up from a miniscule
presence away from roads in the 1980s. At the same time, native
winter annual plants have become very rare and are showing changes in
species composition. This new monoculture potentially has
profound implications for biodiversity, habitat quality, ecosystem
functioning and forage quality of the rangeland. My research applies a
physiological, ecosystem, community and theoretical approach to
understanding the characteristics associated with the irruption and
distribution of E. cicutarium
and its impacts for biodiversity in the San Simon Valley.
Interactions
between Predation and Resource Competition
Galen Holt



Maureen Ryan
Environmental variation and species interactions in amphibian communitiesI'm interested in how environmental variation promotes diversity within species and communities, and how mechanisms such as the storage effect that function through environmental variation can affect endangered species, in particular. My research focuses on a grassland salamander community in the Central Valley of California that includes the threatened California tiger salamander (Ambystoma californiense), hybrid tiger salamanders (A. californiense crossed with A. tigrinum mavortium, an introduced tiger salamander from the central US), and the California newt (Taricha torosa).
I'm using a combination of artificial pond experiments and field experiments to look at whether and how environmental variation promotes coexistence between the two native species, Ambystoma californiense and Taricha torosa, and how coexistence mechanisms operating in the native community are influenced by the addition of hybrid salamanders. Additionally, I'm looking at how environmental variation influences hybridization dynamics between A. californiense and A. tigrinum mavortium and whether mechanisms associated with environmental variation can contribute to the maintenance of genetic polymorphism within hybridizing populations of tiger salamanders. Besides experiments, I'm using theoretical modeling and data from field surveys/sampling to investigate how dispersal and landscape structure interact with local community dynamics to influence species diversity and hybrid spread on a regional scale.
From a conservation perspective, my work is motivated by an interest in how landscape alteration may influence amphibian diversity and species coexistence within extant communities, and in particular how common pond manipulations and the loss of vernal pools in the California Central Valley might influence California tiger salamander populations.

Anna Sears
Environmental variation and species interactions
Ecology is challenged to find simple ways to understand the messiness of the natural world. I like to work from a metacommunity context, accepting that natural ecosystems contain a variety of different habitat types, and that this environmental variation affects the interactions of species within ecosystems. My PhD research specifically involves using natural annual plant systems to test how variation in the environment affects species coexistence. I am also interested in how environmental variation affects food-web interactions, and generally interacts with other forms of density dependence. My field sites are in the Chihuahuan desert and at the Bodega Marine Reserve.
Megan Donahue
Scale dependence in marine populations
For my dissertation research, I studied how
spatial
variation in local density-dependent processes influences regional
dynamics. In collaboration with Marcel Holyoak, I investigated the
effects of local resource distribution, dispersal rate, and the
strength of density dependence on regional dynamics in a highly
manipulable microcosm system, in which we could track population
dynamics over time. Applying the same principles to a field system, I
studied Petrolisthes cinctipes,
a filter-feeding porcelain crab that lives in intertidal cobblefields
along the Pacific coast. Based out of Bodega Marine Laboratory, I
parameterized models of local density-dependence in this organism using
field and laboratory experiments. For Petrolisthes
cinctipes, the primary sources of density dependence are
gregarious settlement, competition, and predation. With Leah Akins, I
quantified the spatial variation that influences these process at two
scales: variation in P. cinctipes
density, predator abundance, and larval supply among rocks within sites
and among sites. By integrating information about spatial variation
into empirically-based models of local dynamics, we can predict what
local processes are most important when "scaling up" to regional
dynamics. For links to publications and Megan's CV, see her website.
Brett Melbourne
Population dynamics and spatial heterogeneity
My main
research
interest is population dynamics,
especially how small-scale heterogeneity affects dynamics at larger
scales. Understanding how spatial and temporal heterogeneity affect the
dynamics of populations and communities is fundamental to ecology.
Indeed, answers to the most prominent questions in ecology involve the
pervasive influence of heterogeneity. Why do species coexist? Why are
there so many species? Why do some species cycle whereas others don't?
How does habitat fragmentation affect extinction?
Why are some species more invasive than others? Spatial and temporal
variation in population densities and in the physical environment are
essential considerations in these questions. My current research and
research to date focuses on the effects of spatial heterogeneity.
Whereas empirical studies have typically focused on heterogeneity
exclusively, theory demonstrates that population dynamics are altered
by spatial heterogeneity only when heterogeneity interacts with
nonlinear demographic processes - the pillar for research in the
Chesson lab. The aim of my research is to develop empirical tools to
measure the interaction
of nonlinearity and heterogeneity and to use these tools to probe real
populations.
Thus, I tackle the important issue of how to integrate theory and
models
with data from real systems, to understand how the interaction of
spatial
heterogeneity and nonlinearity alters population dynamics. I use these
insights
to address important basic and applied questions in ecology.
Kendi Davies
The persistence of species in patchy landscapes
My research focuses on testing and refining theory that makes predictions about the persistence of species in heterogeneous landscapes, including metapopulation and related spatio-temporal theory. So far, I have concentrated on the spatial and temporal dynamics of populations and communities in landscapes that have been modified by humans, through habitat fragmentation and grazing. I am interested in how the life history characteristics of species determine the response of species to landscape heterogeneity. I use quantitative methods to test theory and have approached this work using data from a large-scale, long-term field experiment for beetles and data for a continental scale study of arid zone birds in Australia.
I am currently broadening this focus to explore what this same theory predicts about patterns of biological diversity in naturally patchy landscapes. I am focusing on the grassland plant community in serpentine patches in a non-serpentine matrix (in the University of California’s McLaughlin Reserve). Right now, I am particularly interested in spatial scale and the relationship between diversity and invasibility (Shea and Chesson 2002).
Shea, K., Chesson, P. 2002. Community ecology theory as a framework for biological invasions. Trends in Ecology and Evolution 17, 170-176.
Some recent publications
Davies, K.F., Chesson, P., Harrison, S., Inouye, B.D.,
Melbourne,
B.A., Rice, K.J. 2005. Spatial heterogeneity explains the scale
dependence of the native-exotic diversity relationship. Ecology
86, 1602-1610.
Davies, K. F., C. R. Margules, and J. F. Lawrence. 2004. A synergistic effect puts rare, specialized species at greater risk of extinction. Ecology 85: 265-271.
Davies, K. F., B. A. Melbourne, C. R. Margules, and J. F. Lawrence (in press). Metacommunity structure influences the stability of local beetle communities. In M. Holyoak, M. A. Leibold and R. D. Holt eds. Metacommunities: Spatial Dynamics and Ecological Communities. Chicago, University of Chicago Press.
Davies, K. F., B. A. Melbourne, and C. R. Margules. 2001. Effects of within- and between-patch processes on community dynamics in a fragmentation experiment. Ecology 82: 1830-1846.
Davies, K. F., C. Gascon, and C. R. Margules. 2001. Habitat Fragmentation: consequences, management and future research priorities. Pages 81-97 in M. E. Soulé and G. H. Orians, editors. Conservation Biology: Research Priorities for the Next Decade. Island Press, Washington.
Davies, K. F., C. R. Margules, and J. F. Lawrence. 2000. Which traits of species predict population declines in experimental forest fragments? Ecology 81: 1450-1461.
Davies, K. F., and C. R. Margules. 1998.
Effects
of
habitat fragmentation on carabid beetles: experimental evidence.
Journal of Animal Ecology 67: 460-471.

Charlotte Lee
Theory of multispecies
interactions; human-environment interactions
I study the theory of multispecies interactions, with specific focus on the matrix of coefficients for all pairwise interactions in a community. Specific projects include: 1) Determining relationships between different patterns of direct and indirect competitive (or facilitative) interactions and the structure of model community matrices; 2) determining patterns of shared resource use that result in specific community matrices; and 3) developing techniques to do 1) and 2). I also study human-environment interactions using theory from ecosystem ecology and demography.
Scott Stark
Diversity maintenance in tropical plant communities