The ecological and evolutionary implications of merging different types of networks.

Interactions among species drive the ecological and evolutionary processes in ecological communities. These interactions are effectively key components of biodiversity. Studies that use a network approach to study the structure and dynamics of communities of interacting species have revealed many patterns and associated processes. Historically these studies were restricted to trophic interactions, although network approaches are now used to study a wide range of interactions, including for example the reproductive mutualisms. However, each interaction type remains studied largely in isolation from others. Merging the various interaction types within a single integrative framework is necessary if we want to further our understanding of the ecological and evolutionary dynamics of communities. Dividing the networks up is a methodological convenience as in the field the networks occur together in space and time and will be linked by shared species. Herein, we outline a conceptual framework for studying networks composed of more than one type of interaction, highlighting key questions and research areas that would benefit from their study.

[1]  U. Dieckmann,et al.  The Dynamical Theory of Coevolution : A Derivation from Stochastic Ecological Processes , 1996 .

[2]  R. Díaz-Sierra,et al.  Positive interactions, discontinuous transitions and species coexistence in plant communities. , 2010, Theoretical population biology.

[3]  P. Klinkhamer,et al.  Size constraints and flower abundance determine the number of interactions in a plant /flower visitor web , 2006 .

[4]  Martijn Egas,et al.  Evolution Restricts the Coexistence of Specialists and Generalists: The Role of Trade‐off Structure , 2004, The American Naturalist.

[5]  Colin Fontaine,et al.  Stability of Ecological Communities and the Architecture of Mutualistic and Trophic Networks , 2010, Science.

[6]  Luciano Cagnolo,et al.  Evaluating multiple determinants of the structure of plant-animal mutualistic networks. , 2009, Ecology.

[7]  Michio Kondoh,et al.  Response to Comment on "Foraging Adaptation and the Relationship Between Food-Web Complexity and Stability" , 2003, Science.

[8]  Jordi Bascompte,et al.  Temporal dynamics in a pollination network. , 2008, Ecology.

[9]  T. M. Bezemer,et al.  Divergent composition but similar function of soil food webs of individual plants: plant species and community effects. , 2010, Ecology.

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

[11]  R. May.  Food webs. , 1983, Science.

[12]  C. Lortie,et al.  Do biotic interactions shape both sides of the humped-back model of species richness in plant communities? , 2006, Ecology letters.

[13]  Ruben H. Heleno,et al.  Evaluation of restoration effectiveness: community response to the removal of alien plants. , 2009, Ecological applications : a publication of the Ecological Society of America.

[14]  R. Brys,et al.  Low specificity and nested subset structure characterize mycorrhizal associations in five closely related species of the genus Orchis , 2010, Molecular ecology.

[15]  Paulo R. Guimarães,et al.  Interaction Intimacy Affects Structure and Coevolutionary Dynamics in Mutualistic Networks , 2007, Current Biology.

[16]  Ken Thompson,et al.  Evidence of a causal connection between anti-herbivore defence and the decomposition rate of leaves , 1996 .

[17]  D. Brooks,et al.  Evolutionary biology of parasites. , 1981, Monographs in population biology.

[18]  Edda Klipp,et al.  Biophysical properties of Saccharomyces cerevisiae and their relationship with HOG pathway activation , 2010, European Biophysics Journal.

[19]  John D. Rummel,et al.  Some differences between invasion-structured and coevolution-structured competitive communities: a preliminary theoretical analysis , 1983 .

[20]  P. Jordano.  Coevolution in Multispecific Interactions among Free-Living Species , 2010, Evolution: Education and Outreach.

[21]  J. Bascompte,et al.  Effects of phenotypic complementarity and phylogeny on the nested structure of mutualistic networks , 2007 .

[22]  C. Guyer,et al.  Nestedness of Ectoparasite-Vertebrate Host Networks , 2009, PloS one.

[23]  N. Eisenhauer.  Book reviewAboveground–Belowground Linkages: Biotic Interactions, Ecosystem Processes and Global Change, R.D. Bardgett, D.A. Wardle, Oxford University Press, Oxford (2010), 320 pp., £29.95 (paperback), £60.00 (hardback), ISBN: 978-0-19-954688-6 (paperback), 978-0-19-954687-9 (hardback) , 2011 .

[24]  Stefano Allesina,et al.  Parasites in food webs: the ultimate missing links , 2008, Ecology letters.

[25]  William F Morris,et al.  Ecological Dynamics of Mutualist/Antagonist Communities , 2003, The American Naturalist.

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

[27]  H. Müller-Schärer,et al.  Evolution in invasive plants: implications for biological control. , 2004, Trends in ecology & evolution.

[28]  Rebecca E. Irwin,et al.  Direct and ecological costs of resistance to herbivory , 2002 .

[29]  J. Travis,et al.  Facilitation in plant communities: the past, the present, and the future , 2007 .

[30]  Kevin Gross,et al.  Positive interactions among competitors can produce species-rich communities. , 2008, Ecology letters.

[31]  Jane Memmott,et al.  Global warming and the disruption of plant-pollinator interactions. , 2007, Ecology letters.

[32]  Michel Loreau,et al.  Parallel ecological networks in ecosystems , 2009, Philosophical Transactions of the Royal Society B: Biological Sciences.

[33]  Roger E Bumgarner,et al.  Integrated genomic and proteomic analyses of a systematically perturbed metabolic network. , 2001, Science.

[34]  Pedro Jordano,et al.  Patterns of Mutualistic Interactions in Pollination and Seed Dispersal: Connectance, Dependence Asymmetries, and Coevolution , 1987, The American Naturalist.

[35]  Guy Woodward,et al.  Body size in ecological networks. , 2005, Trends in ecology & evolution.

[36]  R. Gomulkiewicz,et al.  Gene flow and geographically structured coevolution , 1999, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[37]  D. L. Angelis,et al.  STABILITY AND CONNECTANCE IN FOOD WEB MODELS , 1975 .

[38]  J. Thompson,et al.  COEVOLUTIONARY ALTERNATION IN ANTAGONISTIC INTERACTIONS , 2006, Evolution; international journal of organic evolution.

[39]  Christopher Baraloto,et al.  Diversity of the Volatile Organic Compounds Emitted by 55 Species of Tropical Trees: a Survey in French Guiana , 2009, Journal of Chemical Ecology.

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

[41]  C. Fonseca,et al.  Asymmetries, compartments and null interactions in an Amazonian ant-plant community , 1996 .

[42]  Jonathan M. Chase,et al.  Trophic cascades across ecosystems , 2005, Nature.

[43]  H. Howe,et al.  INTERACTION AND COEVOLUTION. , 1984, Evolution; international journal of organic evolution.

[44]  Colin Fontaine,et al.  Are insect pollinators more generalist than insect herbivores? , 2009, Proceedings of the Royal Society B: Biological Sciences.

[45]  J. Bruno,et al.  Inclusion of facilitation into ecological theory , 2003 .

[46]  M. V. D. van der Heijden,et al.  The unseen majority: soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems. , 2008, Ecology letters.

[47]  Jordi Bascompte,et al.  Diversity in a complex ecological network with two interaction types , 2009 .

[48]  Takashi Ikegami,et al.  Food-web formation with recursive evolutionary branching. , 2006, Journal of theoretical biology.

[49]  Nicolas Loeuille,et al.  Evolutionary emergence of size-structured food webs. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[50]  D. Wardle,et al.  Ecological Linkages Between Aboveground and Belowground Biota , 2004, Science.

[51]  M. Häggblom,et al.  Fatty acids of fungi and nematodes—possible biomarkers in the soil food chain? , 2002 .

[52]  Dylan G. Fischer,et al.  COMMUNITY AND ECOSYSTEM GENETICS: A CONSEQUENCE OF THE EXTENDED PHENOTYPE , 2003 .

[53]  J. Bascompte,et al.  Ecological networks : beyond food webs Ecological networks – beyond food webs , 2008 .

[54]  A. Valiente‐Banuet,et al.  The Nested Assembly of Plant Facilitation Networks Prevents Species Extinctions , 2008, The American Naturalist.

[55]  Paulo R Guimarães,et al.  Asymmetries in specialization in ant–plant mutualistic networks , 2006, Proceedings of the Royal Society B: Biological Sciences.

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

[57]  G. Heimpel,et al.  Does floral nectar improve biological control by parasitoids , 2005 .

[58]  P. Asprelli,et al.  The Geographic Mosaic of Coevolution , 2006 .

[59]  Guy Woodward,et al.  BODY SIZE DETERMINANTS OF THE STRUCTURE AND DYNAMICS OF ECOLOGICAL NETWORKS: SCALING FROM THE INDIVIDUAL TO THE ECOSYSTEM , 2005 .

[60]  Paulo R Guimarães,et al.  Cheaters in mutualism networks , 2010, Biology Letters.

[61]  Johan van de Koppel,et al.  Reconciling complexity with stability in naturally assembling food webs , 2009, Nature.

[62]  S. Strauss,et al.  FLORAL CHARACTERS LINK HERBIVORES, POLLINATORS, AND PLANT FITNESS , 1997 .

[63]  T. Okuyama,et al.  Network structural properties mediate the stability of mutualistic communities. , 2008, Ecology letters.

[64]  N. Loeuille,et al.  Emergence of complex food web structure in community evolution models , 2009 .

[65]  Judith L Bronstein,et al.  Evolution and persistence of obligate mutualists and exploiters: competition for partners and evolutionary immunization. , 2007, Ecology letters.

[66]  T. Tscharntke,et al.  Effects of below- and above-ground herbivores on plant growth, flower visitation and seed set , 2003, Oecologia.

[67]  M. Wade.  The co-evolutionary genetics of ecological communities , 2007, Nature Reviews Genetics.

[68]  Steven D. Gaines,et al.  SOME IMPLICATIONS OF DIRECT POSITIVE INTERACTIONS FOR COMMUNITY SPECIES DIVERSITY , 1997 .

[69]  N. Loeuille.  Influence of evolution on the stability of ecological communities. , 2010, Ecology letters.

[70]  Carlos J. Melián,et al.  The nested assembly of plant–animal mutualistic networks , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[71]  É. Thébault,et al.  Does asymmetric specialization differ between mutualistic and trophic networks , 2008 .

[72]  A. Galston.  Fundamentals of Ecology , 1972, The Yale Journal of Biology and Medicine.

[73]  Jane Memmott,et al.  The restoration of parasites, parasitoids, and pathogens to heathland communities. , 2009, Ecology.

[74]  J. Thompson,et al.  The Coevolutionary Process , 1994 .

[75]  B. A. Hawkins,et al.  Parasitoid-Host Food Webs and Donor Control , 1992 .

[76]  Jordi Bascompte,et al.  A neutral‐niche theory of nestedness in mutualistic networks , 2008 .

[77]  Jeff Ollerton,et al.  The pollination ecology of an assemblage of grassland asclepiads in South Africa. , 2003, Annals of botany.

[78]  Joel E. Cohen,et al.  Food web patterns and their consequences , 1991, Nature.

[79]  Michio Kondoh,et al.  Food webs are built up with nested subwebs. , 2010, Ecology.

[80]  Owen L Petchey,et al.  Size, foraging, and food web structure , 2008, Proceedings of the National Academy of Sciences.

[81]  L. Bersier,et al.  The signature of phylogenetic constraints on food-web structure , 2008 .

[82]  Stefano Allesina,et al.  Network structure , predator-prey modules , and stability in large food webs : Electronic Supplementary Material ( ESM ) , 2007 .

[83]  Jordi Bascompte,et al.  Non-random coextinctions in phylogenetically structured mutualistic networks , 2007, Nature.

[84]  Carsten F. Dormann,et al.  Ecological networks - foodwebs and beyond , 2009 .

[85]  A. Ives,et al.  Phylogenetic Analysis of Trophic Associations , 2006, The American Naturalist.

[86]  Jane Memmott,et al.  Food webs: a ladder for picking strawberries or a practical tool for practical problems? , 2009, Philosophical Transactions of the Royal Society B: Biological Sciences.

[87]  N. Franks,et al.  Food Hoarding in Animals , 1990 .

[88]  F. J. Frank van Veen,et al.  The impact of an ant–aphid mutualism on the functional composition of the secondary parasitoid community , 2010 .

[89]  Jane Memmott,et al.  Tolerance of pollination networks to species extinctions , 2004, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[90]  J. Combs,et al.  The Effects of Above- and Belowground Mutualisms on Orchid Speciation and Coexistence , 2011, The American Naturalist.

[91]  Ricard V. Solé,et al.  On nestedness in ecological networks , 2010 .

[92]  K. Burns,et al.  Network properties of arboreal plants: Are epiphytes, mistletoes and lianas structured similarly? , 2009 .

[93]  Jordi Bascompte,et al.  The architecture of mutualistic networks minimizes competition and increases biodiversity , 2009, Nature.

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

[95]  Corinne Vacher,et al.  Architecture of an Antagonistic Tree/Fungus Network: The Asymmetric Influence of Past Evolutionary History , 2008, PloS one.

[96]  Harry Eugene Stanley,et al.  Catastrophic cascade of failures in interdependent networks , 2009, Nature.

[97]  A. Rossberg,et al.  Food webs: experts consuming families of experts. , 2005, Journal of theoretical biology.

[98]  R. Dawkins.  Burying the vehicle , 1994, Behavioral and Brain Sciences.

[99]  S. Schreiber,et al.  Why intraspecific trait variation matters in community ecology. , 2011, Trends in ecology & evolution.

[100]  H. Godfray,et al.  Food web structure of three guilds of natural enemies: predators, parasitoids and pathogens of aphids. , 2008, The Journal of animal ecology.