Linking trophic positions and flow structure constraints in ecological networks: Energy transfer efficiency or topology effect?

Abstract In the present work we investigate whether the distribution of energy flows in ecosystems responds to criteria of trophic organization. We analyzed weighted and unweighted food webs estimating, for each node, trophic position (TP), Shannon's index of inflow diversity ( H ) and individual contribution to the whole average mutual information (AMI). Finally, we performed the same analysis on simulated webs that were constructed using the following criteria: (a) preserving topology and varying link strength; (b) modifying position of links and their intensities. In real ecosystems, moving toward top species, higher pathway redundancy coupled to a strong specialistic trophic behavior was observed. This means that, beside the availability of multiple topological routes, top predators tend to establish “bipolar” predator–prey interactions feeding on preferred preys that, in turn, are mainly preyed by that specialized consumer. Although the analysis shows that, in qualitative food webs, links should be more numerous at the top of the trophic hierarchy, with magnitude more evenly distributed among the interactions, in weighted networks, the tendency of sharing the main flow between donor and receiving compartments (i.e. “bipolar” predator–prey interaction) increases from basal to top species. Patterns displayed by empirical data vanish when indices are calculated for simulated networks, pointing out how topology of energy delivery and flow intensities may be read as a function of the trophic hierarchy.

[1]  Guido Caldarelli,et al.  Universal scaling relations in food webs , 2003, Nature.

[2]  Louis-Félix Bersier,et al.  QUANTITATIVE DESCRIPTORS OF FOOD-WEB MATRICES , 2002 .

[3]  A twin-scale law about the minimum cost , 2003 .

[4]  Neo D. Martinez,et al.  Limits to Trophic Levels and Omnivory in Complex Food Webs: Theory and Data , 2004, The American Naturalist.

[5]  Claude E. Shannon,et al.  A Mathematical Theory of Communications , 1948 .

[6]  M. Emmerson,et al.  MEASUREMENT OF INTERACTION STRENGTH IN NATURE , 2005 .

[7]  Neo D. Martinez,et al.  Food-web structure and network theory: The role of connectance and size , 2002, Proceedings of the National Academy of Sciences of the United States of America.

[8]  Joel E. Cohen,et al.  Community Food Webs: Data and Theory , 1990 .

[9]  R. Paine,et al.  Intertidal community structure , 1974, Oecologia.

[10]  ROBERT M. MAY,et al.  Will a Large Complex System be Stable? , 1972, Nature.

[11]  Stefano Allesina,et al.  Food web networks: Scaling relation revisited , 2005 .

[12]  Anje-Margriet Neutel,et al.  Stability in Real Food Webs: Weak Links in Long Loops , 2002, Science.

[13]  U. M. Scharler,et al.  The consequences of the aggregation of detritus pools in ecological networks , 2005 .

[14]  Neo D. Martinez,et al.  Simple rules yield complex food webs , 2000, Nature.

[15]  Owen L. Petchey,et al.  Interaction strengths in food webs: issues and opportunities , 2004 .

[16]  Stefano Allesina,et al.  Who dominates whom in the ecosystem? Energy flow bottlenecks and cascading extinctions. , 2004, Journal of theoretical biology.

[17]  R. Paine,et al.  Food-web analysis through field measurement of per capita interaction strength , 1992, Nature.

[18]  Kevin McCann,et al.  Structural asymmetry and the stability of diverse food webs , 2006, Nature.

[19]  Robert E. Ulanowicz,et al.  Toward Canonical Trophic Aggregations , 1979, The American Naturalist.

[20]  S. Allesina,et al.  Secondary extinctions in ecological networks: Bottlenecks unveiled , 2006 .

[21]  Stefano Allesina,et al.  Effective trophic positions in ecological acyclic networks , 2006 .

[22]  Charles M. Newman,et al.  A stochastic theory of community food webs I. Models and aggregated data , 1985, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[23]  M. Pascual,et al.  Ecological networks : Linking structure to dynamics in food webs , 2006 .

[24]  Robert E. Ulanowicz,et al.  The effects of taxonomic aggregation on network analysis , 2002 .

[25]  R. Paine A Note on Trophic Complexity and Community Stability , 1969, The American Naturalist.

[26]  Raymond L. Lindeman The trophic-dynamic aspect of ecology , 1942 .

[27]  R. May,et al.  Stability and Complexity in Model Ecosystems , 1976, IEEE Transactions on Systems, Man, and Cybernetics.

[28]  Ferenc Jordán,et al.  Topological keystone species in ecological interaction networks : Considering link quality and non-trophic effects , 2006 .

[29]  G. Polis,et al.  Food Web Complexity and Community Dynamics , 1996, The American Naturalist.

[30]  C. E. SHANNON,et al.  A mathematical theory of communication , 1948, MOCO.

[31]  A. Hastings,et al.  Weak trophic interactions and the balance of nature , 1998, Nature.

[32]  Stefano Allesina,et al.  Steady state of ecosystem flow networks: a comparison between balancing procedures , 2003 .

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

[34]  J. Lawton,et al.  On feeding on more than one trophic level , 1978, Nature.

[35]  A. Neutel,et al.  Energetics, Patterns of Interaction Strengths, and Stability in Real Ecosystems , 1995, Science.

[36]  L. Amaral,et al.  The web of human sexual contacts , 2001, Nature.

[37]  P. Yodzis,et al.  The connectance of real ecosystems , 1980, Nature.

[38]  Robert E. Ulanowicz,et al.  Quantifying the complexity of flow networks: How many roles are there? , 2003, Complex..

[39]  Duncan J. Watts,et al.  Collective dynamics of ‘small-world’ networks , 1998, Nature.

[40]  Robert E. Ulanowicz,et al.  Quantitative methods for ecological network analysi , 2004, Comput. Biol. Chem..

[41]  Robert M May,et al.  Network structure and the biology of populations. , 2006, Trends in ecology & evolution.

[42]  J. Montoya,et al.  Food web complexity and higher‐level ecosystem services , 2003 .

[43]  J. Lawton,et al.  Number of trophic levels in ecological communities , 1977, Nature.

[44]  H. William Hunt,et al.  Influence of Productivity on the Stability of Real and Model Ecosystems , 1993, Science.

[45]  Johan van de Koppel,et al.  Reconciling complexity with stability in naturally assembling food webs , 2007, Nature.

[46]  R. Guimerà,et al.  QUANTITATIVE PATTERNS IN THE STRUCTURE OF MODEL AND EMPIRICAL FOOD WEBS , 2004, q-bio/0401023.

[47]  Jean-Pierre Gabriel,et al.  Phylogenetic constraints and adaptation explain food-web structure , 2004, Nature.

[48]  G. Polis,et al.  Food webs: integration of patterns and dynamics , 1997 .

[49]  Sven Erik Jørgensen,et al.  Complex ecology : the part-whole relation in ecosystems , 1995 .

[50]  Jennifer A. Dunne,et al.  Network structure and robustness of marine food webs , 2004 .

[51]  R. Paine Food webs : linkage, interaction strength and community infrastructure , 1980 .

[52]  M E J Newman,et al.  Community structure in social and biological networks , 2001, Proceedings of the National Academy of Sciences of the United States of America.