Diversity in a complex ecological network with two interaction types

Most studies on ecological networks consider only a single interaction type (e.g. competitive, predatory or mutualistic), and try to develop rules for system stability based exclusively on properties of this interaction type. However, the stability of ecological networks may be more dependent on the way different interaction types are combined in real communities. To address this issue, we start by compiling an ecological network in the Donana Biological Reserve, southern Spain, with 390 species and 798 mutualistic and antagonistic interactions. We characterize network structure by looking at how mutualistic and antagonistic interactions are combined across all plant species. Both the ratio of mutualistic to antagonistic interactions per plant, and the number of basic modules with an antagonistic and a mutualistic interaction are very heterogeneous across plant species, with a few plant species showing very high values for these parameters. To assess the implications of these network patterns on species diversity, we study analytically and by simulation a model of this ecological network. We find that the observed correlation between strong interaction strengths and high mutualistic to antagonistic ratios in a few plant species significantly increases community diversity. Thus, to predict the persistence of biodiversity we need to understand how interaction strength and the architecture of ecological networks with different interaction types are combined.

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

[2]  M E J Newman Assortative mixing in networks. , 2002, Physical review letters.

[3]  S. Rivas-Martínez,et al.  Donana vegetation, (Huelva, Spain). , 1980 .

[4]  F. Javier.,et al.  Biología reproductiva del matorral de Doñana , 1985 .

[5]  J. Amat Effects of wintering greylag geese Anser anser on their Scirpus food plants , 1995 .

[6]  Javier Herrera,et al.  Pollination relationships in southern Spanisch Mediterranean shrublands , 1988 .

[7]  A. Green,et al.  Passive internal transport of aquatic organisms by waterfowl in Doñana, south-west Spain , 2003 .

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

[9]  W. Wilson,et al.  COEXISTENCE OF MUTUALISTS AND EXPLOITERS ON SPATIAL LANDSCAPES , 2003 .

[10]  J. E. Cohen,et al.  Food webs and niche space. , 1979, Monographs in population biology.

[11]  Lloyd Goldwasser,et al.  SAMPLING EFFECTS AND THE ESTIMATION OF FOOD‐WEB PROPERTIES , 1997 .

[12]  M. Emmerson,et al.  Predator–prey body size, interaction strength and the stability of a real food web , 2004 .

[13]  P. Raven,et al.  BUTTERFLIES AND PLANTS: A STUDY IN COEVOLUTION , 1964 .

[14]  A. Andreu,et al.  Seed consumption and dispersal by the spur-thighed tortoise Testudo graeca , 1988 .

[15]  J. M. Smith,et al.  The Coevolution and Stability of Competing Species , 1976, The American Naturalist.

[16]  W. Fagan,et al.  Hatch Density Variation of a Generalist Arthropod Predator: Population Consequenes and Community Impact , 1994 .

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

[18]  B. Menge,et al.  Indirect Effects in Marine Rocky Intertidal Interaction Webs: Patterns and Importance , 1995 .

[19]  Pablo A. Marquet,et al.  Intraguild predation: a widespread interaction related to species biology , 2004 .

[20]  C. Herrera MEASURING THE EFFECTS OF POLLINATORS AND HERBIVORES: EVIDENCE FOR NON-ADDITIVITY IN A PERENNIAL HERB , 2000 .

[21]  J. Diamond,et al.  Ecology and Evolution of Communities , 1976, Nature.

[22]  M S Ringel,et al.  The stability and persistence of mutualisms embedded in community interactions. , 1996, Theoretical population biology.

[23]  P. Jordano Avian Fruit Removal: Effects of Fruit Variation, Crop Size, and Insect Damage. , 1987, Ecology.

[24]  James P. Grover,et al.  The interaction between predation and competition: a review and synthesis , 2002 .

[25]  Peter A. Abrams,et al.  Describing and quantifying interspecific interactions: a commentary on recent approaches , 2001 .

[26]  W. Armbruster,et al.  EXAPTATIONS LINK EVOLUTION OF PLANT–HERBIVORE AND PLANT–POLLINATOR INTERACTIONS: A PHYLOGENETIC INQUIRY , 1997 .

[27]  Jordi Bascompte,et al.  Asymmetric Coevolutionary Networks Facilitate Biodiversity Maintenance , 2006, Science.

[28]  C. Herrera Defense of Ripe Fruit from Pests: Its Significance in Relation to Plant-Disperser Interactions , 1982, The American Naturalist.

[29]  Jordi Bascompte,et al.  Interaction strength combinations and the overfishing of a marine food web. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[30]  Pedro Jordano,et al.  Interaction frequency as a surrogate for the total effect of animal mutualists on plants , 2005 .

[31]  W. K. Dodds Community structure and selection for positive or negative species interactions , 1988 .

[32]  S. Rivas-martínez,et al.  Vegetación de Doñana (Huelva, España) , 1980 .

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

[34]  Neo D. Martinez,et al.  Allometric scaling enhances stability in complex food webs. , 2006, Ecology letters.

[35]  S. Jang,et al.  Dynamics of herbivore-plant-pollinator models , 2002, Journal of mathematical biology.

[36]  J. Wootton,et al.  ESTIMATES AND TESTS OF PER CAPITA INTERACTION STRENGTH: DIET, ABUNDANCE, AND IMPACT OF INTERTIDALLY FORAGING BIRDS , 1997 .

[37]  Jordi Bascompte,et al.  SIMPLE TROPHIC MODULES FOR COMPLEX FOOD WEBS , 2005 .

[38]  John H. Lawton,et al.  Patterns of species interaction strength in assembled theoretical competition communities , 1999 .

[39]  J. Amat Information on the diet of the Stone Curlew Burhinus oedicnemus in Doñana, southern Spain , 1986 .

[40]  Rebecca E. Irwin,et al.  Ecological and Evolutionary Consequences of Multispecies Plant-Animal Interactions , 2004 .

[41]  C. Herrera Aposematic Insects as Six-Legged Fruits: Incidental Short-Circuiting of Their Defense by Frugivorous Birds , 1985, The American Naturalist.

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

[43]  Thomas P. Clausen,et al.  Do Biochemical Exaptations Link Evolution of Plant Defense and Pollination Systems? Historical Hypotheses and Experimental Tests with Dalechampia Vines , 1997, The American Naturalist.

[44]  H. F. Nijhout,et al.  Stability in Real Food Webs: Weak Links in Long Loops , 2002 .

[45]  J. Gómez Predispersal reproductive ecology of an arid land crucifer, Moricandia moricandioides: effect of mammal herbivory on seed production , 1996 .

[46]  R E Ulanowicz,et al.  Ecosystem flow networks: loaded dice? , 1991, Mathematical biosciences.

[47]  S. Shen-Orr,et al.  Network motifs: simple building blocks of complex networks. , 2002, Science.

[48]  Walter K. Dodds,et al.  Interspecific interactions : constructing a general neutral model for interaction type , 1997 .

[49]  A. García‐Ciudad,et al.  FOOD , FEEDING BEHAVIOUR AND NUTRITIONAL ECOLOGY OF WINTERING GREYLAG GEESE Anser anser , 2007 .

[50]  R. May Qualitative Stability in Model Ecosystems , 1973 .

[51]  M. Delibes,et al.  Local feeding specialization by badgers (Meles meles) in a mediterranean environment , 2004, Oecologia.

[52]  D. Janzen SEED‐EATERS VERSUS SEED SIZE, NUMBER, TOXICITY AND DISPERSAL , 1969, Evolution; international journal of organic evolution.

[53]  C. Montes,et al.  Diet of the Red Swamp Crayfish Procambarus Clarkii in Natural Ecosystems of the Doñana National Park Temporary Fresh-water Marsh (Spain) , 1998 .