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Spring 2014 Community Ecology Symposium
Sharon P. LawlerUC Davis Department
of Entomology and Nematology
Professor of Aquatic Entomology and Community Ecology
Sarah BolingerCommunity EcologyApril 2014
Education
B.A., Lehigh University
M.S., Rutgers University
Ph.D., Rutgers University Worked with well-known community ecologist
Peter J. Morin on competition in aquatic systems between insects and vertebrates
Continued to work with him on studies of population dynamics in laboratory protist microcosms
Research Interests
Insect-vertebrate competition in aquatic systems
Food web architecture and population dynamics Diversity effects on ecosystem function Ecotoxicology – effects of toxics on stream
ecology Mosquito control
MODEL SYSTEMS
Laboratory protist microcosms to research population and metapopulation dynamics; food chain architecture
Ecotron research “Ecology in a Bottle”
Protist Microcosm Studies
Microcosms are small, bounded habitats containing the desired number of organisms
Used to study ecological interactions on a scale that is highly replicable and easily controlled
Some population dynamics scale well; others don't
Protist Microcosm Studies
KINGDOM PROTISTA
(may actually be 8 separate kingdoms)
Eukaryotes that don't fit into other kingdoms
Euglena Paramecium
Publications1993 Lawler, Sharon P,. Morin, Peter J. Food web architecture and population dynamics in laboratory microcosms of protists. The American Naturalist, 141(5): 675-686.
1993 Lawler, Sharon P. Species richness, species composition and population dynamics of protists in experimental microcosms. Journal of Animal Ecology, 62: 711-719.
1993 Lawler, Sharon P. Direct and indirect effects in microcosm communities of protists. Oecologia, 93: 184-190.
1995 Balciunas, Dalius and Sharon P. Lawler. Effects of Basal resources, predation, and alternative prey in microcosm food chains. Ecology, 76(4): 1327-1336.
1995 Morin, Peter J. and Sharon P. Lawler. Food web architecture and population dynamics: Theory and empirical evidence. Annual Review of Ecology and Systematics, 26: 505-529.
1996 Morin, Peter J. and Sharon P. Lawler. Effects of food chain length and omnivory on population dynamics in experimental food webs. Food Webs - Integration of Patterns & Dynamics, 218-230.
1996 Holyoak, Marcel and Sharon P. Lawler. The role of dispersal in predator- prey metapopulation dynamics. Journal of Animal Ecology, 65: 640-652.
2000 Holyoak, M., S.P. Lawler and P.H. Crowley. Predicting extinction: Progress with an individual-based model of protozoan predators and prey. Ecology, 81(12): 3312-3329.
2004 Orland, M.C. and S.P. Lawler. Resonance inflates carrying capacity in protist populations with periodic resource pulses. Ecology, 85(1): 150-157.
2005 Holyoak, M. H. and S. P. Lawler. The contribution of laboratory experiments on protists to understanding population and metapopulation dynamics. Advances in Ecological Research, 37: 245-271.
Food Web Architecture and Population Dynamics
In theory, food chain length and presence of omnivory are important to population dynamics
Food web theory was controversial because experimental evidence of the effects of food chain length and omnivory (and other food web characteristics) on population dynamics were relatively few at the time
Lawler and Morin 1993
Food web architecture and population dynamics in laboratory microcosms of protists.
Look at population dynamics in protist microcosms Do protist communities in longer food chains
experience more instability? Does the presence of omnivory by top predators
destabilize population dynamics? Complications: stability as evaluated in model
systems is harder to measure empirically
Look for parallels between model behavior and measurable dynamics in experimental populations (used as proxy for stability)
Dynamics used: persistence time temporal variability of population size return time
Lawler and Morin 1993
Detritus-based food webs
Bacterivorous ciliates
Tetrahymena pyriformis Colpidium striatum
Facultatively omnivorous ciliate
Blepharisma americanum
Experimental Setup
Experimental Setup, cont.
Testing effect of position in food web on stability of population of a species
Compare mean abundance and temporal variation in abundance in populations of bacterivores (T. pyriformis and C. striatum) when each is top predator (short food chain length) or penultimate predator (long food chain length)
Long food chains also differ in whether top predator is omnivore or nonomnivore
Experimental Setup, cont.
Testing effects of omnivory Two conditions for facultative omnivore B.
americanum: 1. feeds only as bacterivore 2. feeds as omnivore
Compare population dynamics between the two Also compare to population dynamics of a
nonomnivore top predator Lastly, compare effects on prey population stability by
looking at population dynamics in bacterivores preyed on by omnivores vs. nonomnivores
Do longer food chains and food chains containing omnivory show signs of unstable population dynamics?
Results and Conclusions
Results and Conclusions
Addition of top predator reduces abundance of bacterivores Blepharisma increases more rapidly and has higher mean
abundance when feeding as omnivore; max population was the same
Population dynamics of bacterivores vary more in longer food chains except in one case
Omnivore abundance varies less than that of nonomnivores at third trophic level
Blepharisma shows greater variation when restricted to bacteria, because of slower growth of these populations
Results and Conclusions
Tentatively support population fluctuation and extinction increase with increased chain length
Omnivore study seems to indicate that species feeding at multiple trophic levels better endure fluctuations in prey abundance
Generalization requires more research, but it's important that real communities of organisms display theoretical food web phenomena
Lawler and Morin 1995
How to find experimental evidence of issues within food web theory:
Factors limiting food chain length How length and complexity affect trophic
cascades How length and complexity affect population
dynamics
Experimental Evidence of Food Web Theory
Lots of theoretical work existed, but much less experimental work – why?
Hard to get from long-lived organisms in natural systems
Skepticism exists over whether food web models are accurate and even applicable to natural systems at all
Lawler and Morin: Theories are testable, especially in artificially constructed environments (ie microcosms)
Food Web Theory: Background
Elton: Food chains are short Hypotheses: energetic transfer efficiency;
dynamic instability Omnivory
Lotka-Volterra models destabilize with omnivory
Thought to be uncommon until recently
Food Web Complexity and Population Dynamics
Relationship between complexity and stability – positive or negative?
Empirical studies hard to interpret No studies vary connectance while holding
species richness constant
Testing Food Web Theory – Future Work
Show quantitatively that dynamics are similar between model and real system
Study food chains longer than 2 or 3 levels Increase species richness in studies More work needed in studies of:
Omnivory effect on population dynamics Effects of nutrient enrichment Effects of more complex food chains on
trophic cascades
Contributions of Laboratory Microcosm Studies
Holyoak, M., and S. P. Lawler 2005. The contribution of laboratory experiments on protists to understanding population and metapopulation dynamics. Advances in Ecological Research, Vol. 37: Population Dynamics and Laboratory Ecology 37:245-271.
Huge variety of protists useful in constructing communities
Many protists make good analogs of larger species with similar ecological strategies
Studying protists is convenient – short generation time, high replicability
Protist study has been historically important in verifying models, like Gause's studies of logistic growth and competitive exclusion
How useful are natural microcosms for study?
Whole ecosystem vs. laboratory microcosm studies: tractability vs realism in ecological study
Can natural microcosms help circumvent the conflict?
Potential for replication; natural boundaries; small size; short generation time of most organisms
Some questions well-suited to these systems:
How does diversity affect ecosystem function? How does the metacommunity affect species
richness? So why use natural? How good is the external validity,
actually?
Srivastava, D.S, Kolasa, J., Bengtsson, A. Gonzalez, Lawler, S.P., Miller, T.E., Munguia, T, Romanuk, Schneider, D.C., Trzcinski, M.K. 2004. Are natural microcosms useful model systems for ecology? Trends in Ecology & Evolution, 19(7): 379-384.
Ecotron
Climate-controlled facilities for ecological experiments
16 chambers
Uses of Ecotron
Create simplified communities to study in a lab
Real-life simplified model of nature
Because of climate control, experiments can be replicated across the chambers and statistical analysis is more robust
Species Diversity and Ecosystem Performance
Naeem, S. et al. 1995. Empirical evidence that declining species diversity may alter the performance of ecosystems. Philosophical Transactions of the Royal Society of London Series B, 347: 249-262.
Hector, A., J. Joshi, S.P. Lawler, E.M. Spehn, and A. Wilby. 2001. Conservation implications of the link between biodiversity and ecosystem functioning. Oecologia, 129: 624-628.
Ecotron Experiment
Direct manipulation of diversity Replication: 14 chambers; all
conditions held constant except diversity – high, med, low
Four trophic levels Keep at least one member of each
trophic group and functional group Look at effects on identified
ecosystem processes Community respiration, productivity,
decomposition, nutrient retention, water retention
Using Mesocosms to Study Effects of Diversity
Loss of a whole trophic level or functional group has clear impact, but what about part? As biodiversity declines, will ecosystem function change?
Know about causes of diversity, and about biogeochemical cycles and energy in ecosystems, but what about how diversity affects cycling and energy flow?
Four hypotheses at the time
Results and Conclusions
“Higher diversity systems had more dense, more complex canopies, higher numbers of earthworms and insect herbivores, greater rates of CO2 flux, greater productivity and greater accumulation of phosphorus and potassium.”
Doesn't appear to be an artifact of particular plant community used
Limitations
Some caveats when attempting to extrapolate results to natural systems, but it appears that affecting diversity can cause ecosystem function to change even if trophic structure is unmanipulated, but changes vary across functions. Also it appears that if loss of diversity affects canopy structure, CO2 and productivity are affected.
Other Research
Cascade frogs What are indirect effects of introduced trout on
Rana cascadae? Competition for prey appears to be limiting
populations of R. cascadae Mosquito control
Literature Cited
Lawler, Sharon P,. Morin, Peter J. 1993. Food web architecture and population dynamics in laboratory microcosms of protists. The American Naturalist, 141(5): 675-686.
Naeem, S. et al. 1995. Empirical evidence that declining species diversity may alter the performance of ecosystems. Philosophical Transactions of the Royal Society of London Series B, 347: 249-262.
Morin, Peter J. and Sharon P. Lawler. 1995. Food web architecture and population dynamics: Theory and empirical evidence. Annual Review of Ecology and Systematics, 26: 505-529.
Srivastava, D.S, Kolasa, J., Bengtsson, A. Gonzalez, Lawler, S.P., Miller, T.E., Munguia, T, Romanuk, Schneider, D.C., Trzcinski, M.K. 2004. Are natural microcosms useful model systems for ecology? Trends in Ecology & Evolution, 19(7): 379-384.
Holyoak, M., and S. P. Lawler 2005. The contribution of laboratory experiments on protists to understanding population and metapopulation dynamics. Advances in Ecological Research, Vol. 37: Population Dynamics and Laboratory Ecology 37:245-271.
Joseph, M., J. Piovia-Scott, S. Lawler and K. Pope. 2011. Indirect effects of introduced trout on Cascades frogs (Rana cascadae) via shared aquatic prey. Freshwater Biology, 56: 828-838.