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Elucidating the Relationship Between Indirect Effects and Ecosystem Stability

Jessica Robbins
University of California Berkeley & The University of Georgia

How Do We Model Ecosystems? TREES BIRDS FISH INSECTS DETRITUS PARASITES MICROBES

Terms & Concepts

S: Species richness (number of species in the system) C: Connectance (number of connections present in the system / number of possible connections) Basal Species: Species which has no prey Stability: Staying essentially unchanged in the presence of perturbations; returning to a referential state or dynamics after a disruption

Indirect Effects
.5 .5

.25

Indirect Effects Ratio


+ + ] [ = = = =

G = Flow intensity matrix Gij = the fraction of species js flow material that goes directly to species i T = Throughflow vector Ti = all the energy species i obtains by consuming other species + all the energy species i obtains from the environment fij = Rate of direct flow from species j to species i

Objective

Investigate the connection between the indirect effects ratio of an ecosystem and the degree of its response to environmental perturbation

The Niche Model

Image Source: Williams, R.J., Martinez, N.D., 2000. Simple rules yield complex food webs. Nature 404, 180-183.

1. Each species is assigned a niche value (ni) between 0 and 1 according to a Beta distribution with =1.5 and =5 2. Each species is assigned a feeding range size (ri), determined by multiplying the result of a Beta function with mean 2C by ni 3. The location of the center of each range (ci) is drawn uniformly between the values ri/2 and ni 4. Species prey on those that fall within their feeding range, with a higher probability of feeding on species near the center of their range

Additional Conditions

Networks to be used for analysis must possess the following characteristics: 1. No cannibalism 2. All energy must ultimately derive from basal species 3. No isolated species or sub-networks

Possible Confounding Variables


Number of Species Food webs that can exhibit a wide range of functional responses to environmental change are often more stable Connectance Networks with higher connectance tend to be more stable Number of basal species There is thought to be a connection between abundance of basal species and system stability Body Mass Ratio Predator-prey body-mass ratios between 10 and 100 tend to promote stability Predator-Prey functional response System stability is promoted when predator response to prey abundance is not completely linear

Methodology
1. Networks were simulated using combinations of S (network size) equal to 5, 10, and 15 and expected connectance equal to .10 , .15, .20, .25, and .30. 10 networks with each parameter combination were created (1500 in total) 2. Once network dynamics had reached steady state, one species was chosen at random and its biomass was multiplied by a factor uniformly distributed between 0 and 5. The perturbed network was then allowed to go to steady state, and the norm of the biomass difference between the original and altered networks was calculated. This process was repeated 50 times for each network 3. Interference from confounding variables was minimized by grouping networks by connectance value, number of species, and abundance of basal species prior to analysis and maintaining a similar distribution of functional response and body mass ratio

Change In Biomass
For basal species: =
=

For consumer species: = +


=

() ()

Bi = biomass of species i xi= mass-specific metabolic rate yij= maximum ingestion rate of prey species j by predator species i eij= assimilation efficiencythe fraction of biomass of species j lost due to consumption by species i that is actually metabolized Gi(B) = normalized growth rate Fij(B) = functional response of consumer i and resource j

Results

Conclusions
1. There is a strong correlation between indirect effects ratio and connectance. This means that is difficult to delineate their impacts on ecosystem dynamics, and that stability that has been previously been attributed to connectance may also result from indirect effects 2. Both connectance and indirect effects promote ecosystem stability. However, this is not a guarantee that the ecosystems with the highest levels of connectance or highest indirect effects ratio will be the most stable, or vice versa 3. The indirect effects ratio may be a more informative metric than connectance 4. Increasing the abundance of basal species tends to decrease system stability 5. Increasing the network size both promotes instability and increases the range of ecosystem response 6. However, the influence of network size and basal species abundance on network stability may be poorly represented by computational modeling techniques

Possible Sources of Error


1. Each network simulation was run for 500 time units and networks that did not reach steady state within this time frame were not analyzed. This may bias the results in favor of more stable networks 2. Networks with significantly more than 15 species may exhibit different dynamics 3. Other ways of quantifying stability may yield different insights

Avenues For Future Investigation


1. Adding parasite species to food web 2. Adding detrital compartment to food web 3. Modeling non-feeding interactions 4. Simulating larger webs

Acknowledgments
This work would not have been possible without the support, advice, and assistance of: Dr. Caner Kazanci Qianqian Ma Dr. Alice Boit Dr. Rosalyn Rael Dr. Kevin Lafferty The University of Georgia The Ohio State University

Resources
1. 2. Allesina, S., Tang, S., 2012. Stability criteria for complex ecosystems. Nature 483, 208-208. Bellingeri, M., Cassi, D., Vincenzi, S., 2013. Increasing the extinction risk of highly connected species causes a sharp robust-to-fragile transition in empirical food webs. Ecological Modeling 251, 1-8. Brose, U., 2008. Complex food webs prevent competitive exclusion among producer species. Proceedings of the Royal Society B: Biological Sciences 275, 2507-2514. Brose, U., Williams, R.J., Martinez, N.D., 2006. Allometric scaling enhances stability in complex food webs. Ecology Letters 9, 1228-1236. Dunne, J.A., Williams, R.J., Martinez, N.D, 2002. Network structure and biodiversity loss in food webs: robustness increases with connectance. Ecology Letters 5, 558-567. Grimm, V., Schmidt, E., Wissel, C., 1992. On the application of stability concepts in ecology. Ecological Modeling 63, 143-161. Kalinkat, G., Scheider, F.D., Digel, C., Guill, C., Rall, B.C., Brose, U., 2013. Body masses, functional responses and predator-prey stability. Ecology Letters 1-9. Lin, Y., Sutherland, W.J., 2013. Color and degree of interspecific synchrony of environmental noise affect the variability of complex ecological networks. Ecological Modeling 263, 162-173. 1. Ma, Q., Kazanci, C., 2012. Analysis of indirect effects within ecosystem models using pathwaybased methodology. Ecological Modeling 252, 238245. Martinez, N.D., Williams, R.J., Dunne, J.A., 2005. Diversity, complexity, and persistence in large model ecosystems. Santa Fe Institute. McCann, K.S., 2000. The diversity-stability debate. Nature, 405 228-233. Rall, B.C., Guill, C., Brose, U., 2008. Food-web connectance and predator interference dampen the paradox of enrichment. Oikos 117, 202-213. Rip, J.M.K., McCann, K.S., 2011. Cross-ecosystem differences in stability and the principle of energy flux. Ecology Letters, 14 733-740. Williams, R.J., 2008. Effects of network and dynamical model structure on species persistence in large model food webs. Theoretical Ecology 3, 285-294. Williams, R.J., Anandanedesan, A., Purves, D., 2010. The probabilistic niche model reveals the niche structure and role of body size in complex food webs. PLoS ONE 5, e12092. Williams, R.J., Martinez, N.D., 2000. Simple rules yield complex food webs. Nature 404, 180-183. Williams, R.J., Martinez, N.D., 2004. Stabilization of chaotic and non-permanent food-webs dynamics. European Physical Journal B 38. 297303.

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Functional Response
=
+ + 0 0

i = consumer species j = resource species

B0= half-saturation density Bi= biomass of species I ij= relative prey preferences di= predator interference term h= hill coefficient, controls the strength of functional response

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