Tony Ives

Research Interests

My general areas of research are community ecology and population biology, although my work also extends into evolution and behavior. I do both theoretical and empirical work, and often try hard to combine them directly.


My main projects include
        Multispecies interactions and environmental fluctuations
        Population dynamics of insect predator-prey systems
        Phylogenetic patterns in comparisons among species and communities
        Theory, statistics, and complex population dynamics

Multispecies interactions and environmental fluctuations

        How do interactions among species within an ecological community influence the response of the community to environmental perturbations? This is a central question in understanding how community structure affects the resistance and resilience of ecosystems. I am particularly interested in the application of stochastic models to multispecies data sets, using patterns of population fluctuations to explore the consequences of species interactions for the general dynamical properties of the system.


 Selected publications

  • Ives, A. R., B. Dennis, K. L. Cottingham, and S. R. Carpenter. 2003.  Estimating community stability and ecological interactions from time-series data.  Ecological Monographs 73: 301-330.
  •  Kilpatrick, A. M.G, and A. R. Ives. 2003.  Species interactions can explain Taylor’s power law for time series.  Nature 422: 65-68.
  • Beisner, B. E.P, A. R. Ives, and S. R. Carpenter. 2003.  The effects of an exotic fish invasion on the prey communities of two lakes.  Journal of Animal Ecology 72: 331-342.
  •  Ripa, J.P, and A. R. Ives. 2003.  Food web dynamics in correlated and autocorrelated environments.  Theoretical Population Ecology 64: 369-384.
  • Ives, A. R., and B. J. CardinaleP. 2004.  Food-web interactions govern the resistance of communities following non-random extinctions.  Nature 429: 174-177.
  •  Ives, A. R., B. J. CardinaleP, and W. E. Snyder. 2005.  A synthesis of subdisciplines: predator-prey interactions, and biodiversity and ecosystem functioning.  Ecology Letters 8: 102-116.
  • Lehmann-Ziebarth, N.U, and A. R. Ives. 2006. The structure and stability of model ecosystems assembled in a variable environment. Oikos 114:451-464.
  • Weis, J. J.U, B. J. CardinaleP, K. J. Forshay, and A. R. Ives. 2007. Effects of species diversity on the production of community biomass change in form and magnitude over the course of succession. Ecology 88:929-939.
  • Ives, A. R., and S. R. Carpenter. 2007. Stability and diversity of ecosystems. Science 317:58-62.
  • Duffy, M. A.P, S. R. Hall, C. E. Caceres, and A. R. Ives. 2009. Rapid evolution, seasonality, and the termination of parasite epidemics. Ecology 90:1441-1448.
  • Abbott, K. C.P, J. Ripa, and A. R. Ives. 2009. Environmental variation in ecological communities and inferences from single-species data Ecology 90:1268-1278.
  • Duffy, M. A., C. E. Caceres, S. R. Hall, A. J. Tessier, and A. R. Ives. 2010. Temporal, spatial, and between-host comparisons of patterns of parasitism in lake zooplankton. Ecology 91:3322-3331.
  • Northfield, T. D., G. B. Snyder, A. R. Ives, and W. E. Snyder. 2010. Niche saturation reveals resource partitioning among consumers. Ecology Letters 13:338-348.
  • NorthfieldP, T. D. and A. R. Ives. 2013. Coevolution and the effects of climate change on interacting species. PLoS Biology 11.

Lab members:
            P = UW Postdoctoral researcher
            G = UW Graduate student
            U = UW Undergraduate
            H = High School student  

Population dynamics of insect predator-prey systems

   Pea aphids in Wisconsin alfalfa fields rarely reach densities sufficient to cause crop damage due to high levels of predation and parasitism. How do predators and parasitoids act to suppress aphid populations in a persistent, stable fashion? And how does evolution counteract the effects of environmental fluctuations and predators on aphids? To address these problems, I employ experiments both in the lab and in the field, and integrate the results of the experiments using a variety of theoretical tools. As a method to explore aphid evolution, I am taking advantage of the numerous bacterial symbionts of aphids (part of the aphid extended genome) that confer heat tolerance and resistance to parasitoids and parasites. This is part of a project with Jason Harmon (NDS), Kerry Oliver (UG-Athens), and Volker Radeloff (UW-Madison).


  Selected publications

  • Snyder, W. E.P, and A. R. Ives. 2003.  Interactions between specialist and generalist natural enemies: parasitoids, predators, and pea aphid biological control.  Ecology 84: 91-107.
  • Östman, Ö., and A. R. Ives. 2003.  Scale-dependent indirect interactions between two prey species through a shared predator.  Oikos 102: 505-514.
  • Cardinale, B. J.P, C. T. HarveyG, K. GrossG, and A. R. Ives. 2003.  Biodiversity and biocontrol: Emergent impacts of a multi-enemy assemblage on pest suppression and crop yield in an agroecosystem.  Ecology Letters 6: 857-865.
  • Lin, L. A.H, and A. R. Ives. 2003.  The effect of parasitoid host-size preference on host population growth rates: an example of Aphidius colemani and Aphis glycines.  Ecological Entomology 28: 542-550.
  • Aquilino, K. M.U, B. J. CardinaleP, and A. R. Ives. 2005.  Reciprocal effects of host-plant and natural enemy diversity on herbivore suppression: An empirical study of a model tri-trophic system.  Oikos 108: 275-282.
  • Gross, K.G, A. R. Ives, and E. V. Nordheim. 2005. Estimating time-varying vital rates from observation time series: a case study in aphid biological control. Ecology 86:740-752.
  • Langley, S. A.U, K. J. TilmonP, B. J. CardinaleP, and A. R. Ives. 2006. Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae) alters individual fixed preferences for pea aphid color morphs. Oecologia 150:172-179.
  • Meisner, M.U, J. P. HarmonP, and A. R. Ives. 2007. Presence of an unsuitable host diminishes the competitive superiority of an insect parasitoid: a distraction effect. Population Ecology 49.
  • Harmon, J. P.P, N. A. Moran, and A. R. Ives. 2009. Species response to environmental change: impacts of food web interactions and evolution. Science 323:1347-1350.
  • Meisner, M. H.U, J. P. HarmonP, C. T. HarveyG, and A. R. Ives. 2011. Intraguild predation on the parasitoid Aphidius ervi by the generalist predator Harmonia axyridis: the threat and its avoidance. Entomologia Experimentalis Et Applicata 138:193-201.
  • Meisner, M. H.U, J. P. HarmonP, and A. R. Ives. 2011. Response of coccinellid larvae to conspecific and heterospecific larval tracks: A mechanism that reduces cannibalism and intraguild predation. Environmental Entomology 40:103-110.
  • Straub, C. S., A. R. Ives, and C. Gratton. 2011. Evidence for a trade-off between host-range breadth and host-use efficiency in aphid parasitoids. American Naturalist 177:389-395.
  • Schooler, S. S., P. De Barro, and A. R. Ives. 2011. The potential for hyperparasitism to compromise biological control: Why don't hyperparasitoids drive their primary parasitoid hosts extinct? Biological Control 58:167-173.
  • Meisner, M. H.U and A. R. Ives. 2013. Foraging efficiency and the fitness consequences of spatial marking by ladybeetle larvae. Oikos 122:1238-1246.
  • Bianchi, F., A. R. Ives, and N. A. Schellhorn. 2013. Interactions between conventional and organic farming for biocontrol services across the landscape. Ecological Applications 23:1531-1543.
  • Barton, B. T.P and A. R. Ives. 2014. Species interactions and a chain of indirect effects driven by reduced precipitation. Ecology 95:486-494.
  • Barton, B. T.P, and A. R. Ives. 2014. Direct and indirect effects of warming on aphids, their predators, and ant mutualists. Ecology 95:1479-1484.

Lab members:
            P = UW Postdoctoral researcher
            G = UW Graduate student
            U = UW Undergraduate
            H = High School student   

Phylogenetic patterns in comparisons among species and communities

        Evolutionarily related species are likely to share many of the same traits due to their phylogenetic descent from a common ancestor. With Ted Garland (UC-Riverside) as a frequent collaborator, I develop statistical methods to investigate phylogenetic correlations of traits among species. Because the occurrence and abundance of species in ecological communities depend on their traits, community composition is also likely to show phylogenetic patterns with, for example, phylogenetically related species more likely to occur in the same communities. Thus, I am extending methods derived for comparisons of traits among species to compare the compositions of different ecological communities.

  Selected publications

  • Garland, T., and A. R. Ives. 2000.  Using the past to predict the present: confidence intervals for regression equations in phylogenetic comparative methods.  American Naturalist 155: 346-364.
  • Blomberg, S. P., T. Garland, and A. R. Ives. 2003.  Testing for phylogenetic signal in comparative data: behavioral traits are more labile.  Evolution 57: 171-745.
  • Ives, A. R., and H. C. J. Godfray. 2006. Phylogenetic analysis of trophic associations. American Naturalist 168:E1-E14.
  • Huey, R. B., B. Moreteau, J. C. Moreteau, P. Gibert, G. W. Gilchrist, A. R. Ives, T. Garland Jr., and J. R. David. 2006. Evolution of sexual size dimorphism in a Drosophila clade, the D. obscura group. Zoology 109:497-505.
  • Helmus, M. R.G, T. J. BlandU, C. K. WilliamsP, and A. R. Ives. 2007. Phylogenetic measures of biodiversity. American Naturalist 169:E68-E83.
  • Helmus, M. R.G, K. SavageU, M. W. Diebel, J. T. Maxted, and A. R. Ives. 2007. Separating the determinants of phylogenetic community structure. Ecology Letters 10:917-925
  • Ives, A. R., P. E. Midford, and T. Garland Jr. 2007. Within-species variation and measurement error in phylogenetic comparative methods. Systematic Biology 56:252-270.
  • Lavin, S. R., W. H. Karasov, A. R. Ives, K. M. Middleton, and T. Garland, Jr. 2008. Morphometrics of the avian small intestine, compared with non-flying mammals: a phylogenetic approach. Physiological and Biochemical Zoology 81:526-550.
  • Zheng, L., A. R. Ives, T. Garland, B. R. Larget, Y. Yu, and K. F. Cao. 2009. New multivariate tests for phylogenetic signal and trait correlations applied to ecophysiological phenotypes of nine Manglietia species. Functional Ecology 23:1059-1069.
  • Ives, A. R. and T. Garland. 2010. Phylogenetic logistic regression for binary dependent variables. Systematic Biology 59:9-26.
  • Ives, A. R. and M. R. HelmusG. 2010. Phylogenetic metrics of community similarity. American Naturalist 176:E128-E142.
  • Ives, A. R. and M. R. HelmusG. 2011. Generalized linear mixed models for phylogenetic analyses of community structure. Ecological Monographs 81:511-525.
  • Helmus, M. R. and A. R. Ives. 2012. Phylogenetic diversity-area curves. Ecology 93:S31-S43.
  • Rafferty, N. E.G and A. R. Ives. 2013. Phylogenetic trait-based analyses of ecological networks. Ecology 94:2321-2333.
  • Johnson, M. T. J., A. R. Ives, J. Ahern, and J. P. Salminen. 2014. Macroevolution of plant defenses against herbivores in the evening primroses. New Phytologist 203:267-279.

        Lab members:
            P = UW Postdoctoral researcher
            G = UW Graduate student
            U = UW Undergraduate
            H = High School student

    Theory, statistics, and complex population dynamics

        I am involved in several projects, most of them collaborative, investigating the complex dynamics that some species exhibit. For example, Árni Einarsson, Vincent Jansen, Arnthor Gardarsson and I demonstrated that the extremely large but irregular fluctuations in the abundance of midges in Lake Myvatn, Iceland, could be explained by a model showing alternative dynamical states. The observed dynamics appear to shift stochastically between the two states, a high stable point and a 5-orders-of-magnitude cycle, producing dramatic and unpredictable population patterns.

  Selected publications

  • Williams, C. K.P, A. R. Ives, and R. D. Applegate. 2003.  Population dynamics across geographical ranges: time-series analyses of three small game species.  Ecology 84: 2654-2667.
  • Ives, A. R., S. T. Woody, E. V. Nordheim, C. Nelson, and J. H. Andrews. 2004.  The synergistic effects of stochasticity and dispersal on population densities.  American Naturalist 163: 375-387.
  • Williams, C. K.P, A. R. Ives, R. D. Applegate, and J. RipaP. 2004.  The collapse of cycles in the dynamics of North American grouse populations.  Ecology Letters 7: 1135-1142.
  • Woody, S. T., A. R. Ives, E. V. Nordheim, and J. H. Andrews. 2007. Dispersal, density dependence, and population dynamics of a fungal microbe on leaf surfaces. Ecology 88:1513-1524.
  • Solbreck, C., and A. R. Ives. 2007. Density dependence vs. independence, and irregular population dynamics of a swallow-wort fruit fly. Ecology 88:1466-1475.
  • Forester, J. D., A. R. Ives, M. G. Turner, D. P. Anderson, D. Fortin, H. L. Beyer, D. W. Smith, and M. S. Boyce. 2007. State-space models link elk movement patterns to landscape characteristics in Yellowstone National Park. Ecological Monographs 77:285-299.
  • Ives, A. R., A. Einarsson, V. A. A. Jansen, and A. Gardarsson. 2008. High-amplitude fluctuations and alternative dynamical states of midges in Lake Myvatn. Nature 452:84-87.
  • Ives, A. R., K. C. AbbottP, and N. L. ZiebarthU. 2010. Analysis of ecological time series with ARMA(p,q) models. Ecology 91:858-871.
  • Ziebarth, N. L.U, K. C. AbbottP, and A. R. Ives. 2010. Weak population regulation in ecological time series. Ecology Letters 13:21-31.
  • Foufopoulos, J., A. M. Kilpatrick, and A. R. Ives. 2011. Climate change and elevated extinction rates of reptiles from Mediterranean islands. American Naturalist 177:119-129.
  • Rafferty, N. E.G and A. R. Ives. 2011. Effects of experimental shifts in flowering phenology on plant-pollinator interactions. Ecology Letters 14:69-74.
  • Ives, A. R., P. R. GlaumU, N. L. ZiebarthU, and D. A. Andow. 2011. The evolution of resistance to two-toxin pyramid transgenic crops. Ecological Applications 21:503-515.
  • Schooler, S. S., B. Salau, M. H. Julien, and A. R. Ives. 2011. Alternative stable states explain unpredictable biological control of salvinia in Kakadu. Nature 470:86-89.
  • Foufopoulos, J., A. M. Kilpatrick, and A. R. Ives. 2011. Climate change and elevated extinction rates of reptiles from Mediterranean islands. American Naturalist 177:119-129.
  • Ives, A. R., P. R. GlaumU, N. L. ZiebarthU, and D. A. Andow. 2011. The evolution of resistance to two-toxin pyramid transgenic crops. Ecological Applications 21:503-515.
  • Rafferty, N. E.G and A. R. Ives. 2012. Pollinator effectiveness varies with experimental shifts in flowering time. Ecology 93:803-814.
  • Strier, K. B. and A. R. Ives. 2012. Unexpected demography in the recovery of an endangered primate population. PLoS One 7.
  • Usinowicz, J.U, S. J. Wright, and A. R. Ives. 2012. Coexistence in tropical forests through asynchronous variation in annual seed production. Ecology 93:2073-2084.
  • Ives, A. R. and V. Dakos. 2012. Detecting dynamical changes in nonlinear time series using locally linear state-space models. Ecosphere 3:art58.
  • Jackson, M. M., M. G. Turner, S. M. Pearson, and A. R. Ives. 2012. Seeing the forest and the trees: multilevel models reveal both species and community patterns. Ecosphere 3:art79.
  • Behm, J. E.G, D. A. Edmonds, J. P. HarmonP, and A. R. Ives. 2013. Multilevel statistical models and the analysis of experimental data. Ecology 94:1479-1486.
  • Hansen, G. J. A., A. R. Ives, M. J. Vander Zanden, and S. R. Carpenter. 2013. Are rapid transitions between invasive and native species caused by alternative stable states, and does it matter? Ecology 94:2207-2219.

        Lab members:
            P = UW Postdoctoral researcher
            G = UW Graduate student
            U = UW Undergraduate
            H = High School student

Citizen Science

    With Kerry Oliver, I have started a citizen science project to collect aphids and assay them for symbiotic bacteria. These symbionts perform all kinds of help to the aphids, such as protecting them from high temperatures and parasites. The citizen science project, done with the help of John Losey and the Lost Ladybug Project (U Cornell), will get aphids from all over the country so that we can see how the communities of bacterial symbionts within aphids change geographically. A summary of this project is available as a pdf here.



        Zoology 260 - Introductory Ecology
        Zoology 540 - Theoretical Ecology
        Zoology 956 - Seminar - Ecology

    Note to prospective graduate students:

        I am particularly interested in attracting graduate students who want to combine empirical and theoretical approaches to ecological studies. My lab is diverse, with students having worked on malaria in lizards, biological control of pea aphids, the dynamics of freshwater phytoplankton, fish community composition, plant-pollinator mutualisms, tropical frogs, and other topics. A broad background and interest in collaborating on a wide range of ecological questions is essential.

        When I was applying for graduate school, the most important thing I looked for was a place where I had the flexibility and resources to do my own research. Now as a faculty advisor, I encourage students to develop their own projects and interests. There are limits, however, as I don't want to take on students who are so far a field that I can't give adequate advice and support. But the mix of people in the lab provides a rich collective knowledge that can help with a variety of projects. I actively encourage collaborations within the lab, and between lab members and others on campus. To select prospective graduate students to fit in with the collaborative atmosphere of the lab, the entire lab has a strong say in accepting graduate students.

Graduate students currently supervised:

 Kyle Webert
 Kyle started work at Lake Myvatn as an undergraduate and started his PhD in 2010. He is studying both the aquatic and terrestrial communities of Myvatn, taking advantage of the huge fluctuations in midge abundances as a natural disturbance to the system.

  Fan Huan
  Huan started her PhD in the lab in 2011. She is studying functional and phylogenetic genomics, with particular interests in methods that do not require the assembly and alignment of genomes. These methods are therefore suited for non-model organisms that do not have reference genomes. She is simultaneously completing a PhD at the Xishuangbanna Tropical Botanical Garden, Yunnan, China.

  Joe Phillips
  Joe entered the lab in 2014 and started fieldwork at Lake Myvatn, Iceland. His interests include the stability of complex ecosystems, and the energy and nutrient dynamics of trophic cascades, which he studies in both real and theoretical systems.

Former graduate students:

Jocelyn Behm, Ph.D. – 2012 (Postdoc at VU University, Amsterdam)
Responses of Xishuangbanna's frog species to their environment: habitat selection in adults and phenotypic plasticity in tadpoles.

Kate Forbes, Ph.D. – 2008 (Epic Systems)
Movement and population dynamics of two ladybeetle species in agricultural crops.

Johannes Foufopoulos, Ph.D. – 1998 (Associate Professor at University of Michigan)
Host-parasite interactions in the Mountain Spiny Lizard Sceloporus jarrovi.

Tucker Gilman, Ph.D.– 2010 (Lecturer at University of Manchester)
Evolution in changing environments: insights from theory.

Kevin Gross, Ph.D. – 2003 (Professor at North Carolina State)
The aphid, the wasp, and the matrix: aspects of modeling host-parasitoid and single-species dynamics.

Chad Harvey, Ph.D. – 2007 (Associate Professor at McMaster University)
How variation in the composition of insect food-webs affects the strength and outcome of both direct and indirect interactions among the constituent species.

Matt Helmus, Ph.D. – 2008 (Postdoc at VU University, Amsterdam)
Phylogenetic signal in ecological community responses to environmental variation.

Eric Klopfer, Ph.D. – 1997 (Professor at MIT)
Ecological and evolutionary consequences of explicit spatial structure in exploiter-victim systems.

Jennifer Lynn Klug, Ph.D. – 2000 (Professor at Fairfield University)
Complex effects of colored dissolved organic matter on algal growth and community composition.

Todd Palmer, M.A. – 1994 (Assistant Professor at University of Florida)
The influence of spatial heterogeneity on the behavior and growth of two herbivorous stream insects.

Nicole Rafferty, Ph.D. – 2011 (Postdoc at University of Arizona)
Climate change and shifts in flowering time: effects on plant-pollinator interactions.

Former postdocs

Karen Abbott 2006-09 (Associate Professor at Case Western University)

Brad Cardinale 2002-05 (Professor at University of Michigan)

Meg Duffy 2006-07 (Associate Professor at University of Michigan)

Jason Harmon 2005-09 (Assistant Professor at University of North Dakota)

Ralph Haygood 2002-05 (in private business)

Derek Johnson 2005 (Associate Professor at Virginia Commonwealth University)

John Losey 1996-97 (Associate Professor at Cornell University)

Tobin Northfield 2011-13 (Lecturer at James Cook University, Australia)

Jörgen Ripa 2000-01 (Researcher at Lund University)

Nancy Schellhorn 1998-99 (Research Leader for CSIRO, Australia)

Bill Snyder 1999-2000 (Professor at Washington State University)

Kelley Tilmon 2002-05 (Associate Professor at University of South Dakota)

Chris Williams 2001-04 (Associate Professor at University of Delaware)