Identifying key species in ecosystems with stochastic sensitivity analysis

The development of approaches to estimate the vulnerability of biological communities and ecosystems to extirpations and reductions of species is a central challenge of conservation biology. One key aim of this challenge is to develop quantitative approaches to estimate and rank interaction strengths and keystoneness of species and functional groups, i.e. to quantify the relative importance of species. Network analysis can be a powerful tool for this because certain structural aspects of ecological networks are good indicators of the mechanisms that maintain co-evolved, biotic interactions. A static view of ecological networks would lead us to focus research on highly-central species in food webs (topological key players in ecosystems). There are a variety of centrality indices, developed for several types of ecological networks (e.g. for weighted and un-weighted webs). However, truly understanding extinction and its community-wide effects requires the use of dynamic models. Deterministic dynamic models are feasible when population sizes are sufficiently large to minimize noise in the overall system. In models with small population sizes, stochasticity can be modelled explicitly. We present a stochastic simulation-based ecosystem model for identification of “dynamic key species” in situations where stochastic models are appropriate. To demonstrate this approach, we simulated ecosystem dynamics and performed sensitivity analysis using data from the Prince William Sound, Alaska ecosystem model. We then compare these results to those of purely topological analyses and deterministic dynamic (Ecosim) studies. We present the relationships between various topological and dynamic indices and discuss their biological relevance. The trophic group with the largest effect on others is nearshore demersals, the species mostly sensitive to others is halibut, and the group of both considerable effect on and sensitivity to others is juvenile herring. The most important trophic groups in our dynamical simulations appear to have intermediate trophic levels.

[1]  Ferenc Jordán,et al.  CoSBiLab Graph: The network analysis module of CoSBiLab , 2010, Environ. Model. Softw..

[2]  Ernesto Estrada,et al.  Characterization of topological keystone species: Local, global and “meso-scale” centralities in food webs , 2007 .

[3]  J. Elser,et al.  Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere , 2002 .

[4]  Peter A. Abrams,et al.  IS PREDATOR‐MEDIATED COEXISTENCE POSSIBLE INUNSTABLE SYSTEMS? , 1999 .

[5]  E. Brown,et al.  Aggregations of the jellyfish Aurelia labiata:abundance, distribution, association with age-0 walleye pollock, and behaviors promoting aggregation in Prince William Sound, Alaska, USA , 2000 .

[6]  Toshinori Okuyama,et al.  Local interactions between predators and prey call into question commonly used functional responses , 2009 .

[7]  Ferenc Jordán,et al.  Identifying important species: Linking structure and function in ecological networks , 2008 .

[8]  Ricard V Solé,et al.  Press perturbations and indirect effects in real food webs. , 2009, Ecology.

[9]  Neo D. Martinez,et al.  Network structure and biodiversity loss in food webs: robustness increases with connectance , 2002, Ecology Letters.

[10]  Carl J. Walters,et al.  Ecopath with Ecosim: methods, capabilities and limitations , 2004 .

[11]  Ferenc Jordán,et al.  Network ecology: Topological constraints on ecosystem dynamics , 2004 .

[12]  D. Doak,et al.  7. Predicting the Effects of Species Loss on Community Stability , 2003 .

[13]  Stuart H. Hurlbert,et al.  Functional importance vs keystoneness: Reformulating some questions in theoretical biocenology , 1997 .

[14]  Thomas A. Okey,et al.  Shifted community states in four marine ecosystems : some potential mechanisms , 2004 .

[15]  D. Doak,et al.  The Keystone-Species Concept in Ecology and ConservationManagement and policy must explicitly consider the complexity of interactions in natural systems , 1993 .

[16]  John Scott What is social network analysis , 2010 .

[17]  M. Coll,et al.  Decadal changes in a NW Mediterranean Sea food web in relation to fishing exploitation , 2009 .

[18]  Neo D. Martinez Artifacts or Attributes? Effects of Resolution on the Little Rock Lake Food Web , 1991 .

[19]  Ernest W. Tollner,et al.  Cycling in ecosystems: An individual based approach , 2009 .

[20]  S. Jørgensen,et al.  Movement rules for individual-based models of stream fish , 1999 .

[21]  John D. Stevens,et al.  The effects of fishing on sharks, rays, and chimaeras (chondrichthyans), and the implications for marine ecosystems , 2000 .

[22]  Corrado Priami,et al.  BetaWB: modelling and simulating biological processes , 2007, SCSC.

[23]  J. Jeffers,et al.  Theoretical Studies of Ecosystems: The Network Perspective , 2009 .

[24]  J. Finn,et al.  Measures of ecosystem structure and function derived from analysis of flows. , 1976, Journal of theoretical biology.

[25]  C. Walters,et al.  Structuring dynamic models of exploited ecosystems from trophic mass-balance assessments , 1997, Reviews in Fish Biology and Fisheries.

[26]  P. Yodzis,et al.  DIFFUSE EFFECTS IN FOOD WEBS , 2000 .

[27]  D. Simberloff A succession of paradigms in ecology: Essentialism to materialism and probabilism , 2004, Synthese.

[28]  Carron Shankland,et al.  Developing the Use of Process Algebra in the Derivation and Analysis of Mathematical Models of Infectious Disease , 2003, EUROCAST.

[29]  I. Molnar,et al.  Persistence and flow reliability in simple food webs , 2003 .

[30]  T. F. Hansen,et al.  Copepods act as a switch between alternative trophic cascades in marine pelagic food webs , 2004 .

[31]  Stuart Banks,et al.  A trophic model of a Galápagos subtidal rocky reef for evaluating fisheries and conservation strategies , 2004 .

[32]  J. Castilla,et al.  Challenges in the Quest for Keystones , 1996 .

[33]  Tom A. B. Snijders,et al.  Social Network Analysis , 2011, International Encyclopedia of Statistical Science.

[34]  D. Gillespie Exact Stochastic Simulation of Coupled Chemical Reactions , 1977 .

[35]  Alan Feest,et al.  Biodiversity Quality: a paradigm for biodiversity , 2009 .

[36]  D. Pauly,et al.  A method for identifying keystone species in food web models , 2006 .

[37]  B. C. Patten Environs: The Superniches of Ecosystems , 1981 .

[38]  D. Simberloff 11. Community and Ecosystem Impacts of Single-Species Extinctions , 2003 .

[39]  C. Harley 3. Species Importance and Context: Spatial and Temporal Variation in Species Interactions , 2003 .

[40]  Corrado Priami,et al.  The Beta Workbench: a computational tool to study the dynamics of biological systems , 2008, Briefings Bioinform..

[41]  Ferenc Jordán,et al.  Species positions and extinction dynamics in simple food webs. , 2002, Journal of theoretical biology.

[42]  Chris Tofts Algorithms for task allocation in ants. (A study of temporal polyethism: Theory) , 1993 .

[43]  F. Jordán,et al.  Quantifying positional importance in food webs: A comparison of centrality indices , 2007 .

[44]  Daniel Pauly,et al.  Trophic Mass-Balance Model of Alaska's Prince William Sound Ecosystem, for the Post-Spill Period 1994-1996 , 1999 .

[45]  Astrid Jarre,et al.  Small pelagics in upwelling systems: patterns of interaction and structural changes in "wasp-waist" ecosystems , 2000 .

[46]  Craig R Powell,et al.  The effects of stochastic population dynamics on food web structure. , 2008, Journal of theoretical biology.