Revising the Role of Species Mobility in Maintaining Biodiversity in Communities with Cyclic Competition

One of the most crucial tasks faced by biologists today is revealing the mechanisms which account for biodiversity, yet we are still far from a full understanding of these mechanisms, and in particular the role of spatially heterogeneous population distributions. Recently, the spatially heterogeneous coexistence seen in cyclic competition models—in which species compete as in the game rock-paper-scissors—has brought them to the fore as a paradigm for biodiversity. Research into cyclic competition has so far been focused almost exclusively on stochastic lattice models with discrete space, which ignore several key dynamical aspects. In particular, such models usually assume that species disperse at the same speed. This paper aims to extend our understanding of cyclic competition by applying a reaction–diffusion Lotka–Volterra scheme to the problem, which allows us to vary the mobility of each species, and lets us take into account cyclic competition with more complex underlying mechanisms. In this paper we reveal an entirely new kind of cyclic competition—‘conditional’ cyclic competition, with a different underlying mechanism to ‘classic’ cyclic competition—and we show that biodiversity in communities with cyclic competition in fact depends heavily on the ratios between the species mobilities. Furthermore, we show that this dependence can be completely different for conditional and classic cyclic competition. We also present a wide range of spatiotemporal patterns which are formed in the system, including spiral and target waves, spiralling patches, and irregular chaotic patches. We show that the previously unknown case of conditional cyclic competition is host to a scenario of patchy co-invasion, where the spread of the population front takes place via the formation, splitting and propagation of patches of high species density. This is also an example of invasional meltdown because one competitor facilitates the invasion of the other, but unlike well-known cases of invasional meltdown the co-invaders in this system are not mutualists but antagonistic competitors, and the overall result mitigates, rather than amplifies, the damage done to the native ecosystem.

[1]  F. A. Davidson,et al.  Travelling waves in near-degenerate bistable competition models , 2010 .

[2]  R. May,et al.  Nonlinear Aspects of Competition Between Three Species , 1975 .

[3]  P. Chesson Mechanisms of Maintenance of Species Diversity , 2000 .

[4]  Peter W. W. Lurz,et al.  Modelling the spatial dynamics of parapoxvirus disease in red and grey squirrels: a possible cause of the decline in the red squirrel in the UK? , 2000 .

[5]  Hilla Peretz,et al.  Ju n 20 03 Schrödinger ’ s Cat : The rules of engagement , 2003 .

[6]  P. Levin,et al.  COMMUNITY-WIDE EFFECTS OF NONINDIGENOUS SPECIES ON , 2002 .

[7]  J. Gallas,et al.  How community size affects survival chances in cyclic competition games that microorganisms play. , 2010, Physical review. E, Statistical, nonlinear, and soft matter physics.

[8]  J. Moen Diffuse Competition: A Diffuse Concept , 1989 .

[9]  P. S. Lake,et al.  Invasional 'meltdown' on an oceanic island , 2003 .

[10]  P. Amarasekare Competitive coexistence in spatially structured environments: a synthesis , 2003 .

[11]  D. Simberloff Confronting introduced species: a form of xenophobia? , 2003, Biological Invasions.

[12]  Pan-Jun Kim,et al.  Spatio-temporal dynamics in the origin of genetic information , 2004, q-bio/0406035.

[13]  Bingtuan Li,et al.  Spreading speed and linear determinacy for two-species competition models , 2002, Journal of mathematical biology.

[14]  Margaret A. Riley,et al.  Antibiotic-mediated antagonism leads to a bacterial game of rock–paper–scissors in vivo , 2004, Nature.

[15]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[16]  Maarten C. Boerlijst,et al.  Spatial gradients enhance persistence of hypercycles , 1995 .

[17]  M. Wonham,et al.  Positive effects of a dominant invader on introduced and native mudflat species , 2005 .

[18]  Bai-lian Li,et al.  On the importance of dimensionality of space in models of space-mediated population persistence. , 2007, Theoretical population biology.

[19]  D. Simberloff,et al.  Positive Interactions of Nonindigenous Species: Invasional Meltdown? , 1999, Biological Invasions.

[20]  M. Nowak,et al.  Habitat destruction and the extinction debt , 1994, Nature.

[21]  C. Paquin,et al.  Relative fitness can decrease in evolving asexual populations of S. cerevisiae , 1983, Nature.

[22]  Joydev Chattopadhyay,et al.  Towards a resolution of ‘the paradox of the plankton’: A brief overview of the proposed mechanisms , 2007 .

[23]  Sergei Petrovskii,et al.  Spatiotemporal complexity of patchy invasion in a predator-prey system with the Allee effect. , 2006, Journal of theoretical biology.

[24]  D. Richard,et al.  Thermogenic Capacity in Gray and Black Morphs of the Gray Squirrel, Sciurus carolinensis , 1989, Physiological Zoology.

[25]  J. W. Thomas Numerical Partial Differential Equations: Finite Difference Methods , 1995 .

[26]  M Rejmánek,et al.  Plant invasions — the role of mutualisms , 2000, Biological reviews of the Cambridge Philosophical Society.

[27]  Daniel M. Tompkins,et al.  Ecological replacement of native red squirrels by invasive greys driven by disease , 2003 .

[28]  M. Feldman,et al.  Local dispersal promotes biodiversity in a real-life game of rock–paper–scissors , 2002, Nature.

[29]  R. Nisbet,et al.  How should we define 'fitness' for general ecological scenarios? , 1992, Trends in ecology & evolution.

[30]  György Károlyi,et al.  Rock-scissors-paper game in a chaotic flow: the effect of dispersion on the cyclic competition of microorganisms. , 2005, Journal of theoretical biology.

[31]  C. Cosner,et al.  Spatial Ecology via Reaction-Diffusion Equations , 2003 .

[32]  Ben-Naim,et al.  Spatial organization in cyclic Lotka-Volterra systems. , 1996, Physical review. E, Statistical physics, plasmas, fluids, and related interdisciplinary topics.

[33]  Marcus Frean,et al.  Rock–scissors–paper and the survival of the weakest , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[34]  Y. Lai,et al.  Cyclic competition of mobile species on continuous space: pattern formation and coexistence. , 2010, Physical review. E, Statistical, nonlinear, and soft matter physics.

[35]  Marten Scheffer,et al.  Why plankton communities have no equilibrium: solutions to the paradox , 2004, Hydrobiologia.

[36]  Tao Zhou,et al.  Effects of competition on pattern formation in the rock-paper-scissors game. , 2011, Physical review. E, Statistical, nonlinear, and soft matter physics.

[37]  N. Shigesada,et al.  Biological Invasions: Theory and Practice , 1997 .

[38]  F. A. Davidson,et al.  Reversing invasion in bistable systems , 2012, Journal of mathematical biology.

[39]  B. Sinervo,et al.  The rock–paper–scissors game and the evolution of alternative male strategies , 1996, Nature.

[40]  R. Durrett,et al.  Spatial aspects of interspecific competition. , 1998, Theoretical population biology.

[41]  Jonathan Silvertown,et al.  Cellular Automaton Models of Interspecific Competition for Space--The Effect of Pattern on Process , 1992 .

[42]  Bernd Blasius,et al.  A graphical theory of competition on spatial resource gradients. , 2011, Ecology letters.

[43]  Y. Hosono,et al.  The minimal speed of traveling fronts for a diffusive Lotka-Volterra competition model , 1998 .

[44]  S. Pacala,et al.  Modeling and analysis of stochastic invasion processes , 2000, Journal of mathematical biology.

[45]  M A Lewis,et al.  Spread rate for a nonlinear stochastic invasion , 2000, Journal of mathematical biology.

[46]  György Szabó,et al.  Evolutionary prisoner's dilemma games with voluntary participation. , 2002, Physical review. E, Statistical, nonlinear, and soft matter physics.

[47]  G. Gauze The struggle for existence, by G. F. Gause. , 1934 .

[48]  I. Hanski Spatial scale, patchiness and population dynamics on land , 1994 .

[49]  H. Takahara,et al.  PATCHY INVASION AND THE ORIGIN OF A HEMLOCK–HARDWOODS FOREST MOSAIC , 1998 .

[50]  Sergei Petrovskii,et al.  Allee effect makes possible patchy invasion in a predator-prey system. , 2002 .

[51]  P. Alpert,et al.  Effects of Nitrogen and Salinity on Growth and Competition Between a Native Grass and an Invasive Congener , 2003, Biological Invasions.

[52]  J. Sherratt,et al.  Periodic travelling waves in cyclic populations: field studies and reaction–diffusion models , 2008, Journal of The Royal Society Interface.

[53]  S. Carpenter,et al.  Stability and Diversity of Ecosystems , 2007, Science.

[54]  G. F. Gause,et al.  EXPERIMENTAL ANALYSIS OF VITO VOLTERRA'S MATHEMATICAL THEORY OF THE STRUGGLE FOR EXISTENCE. , 1934, Science.

[55]  Joseph H. Connell,et al.  Species coexistence, keystone species, and succession: a sensitivity analysis , 1994 .

[56]  Jonathan A. Sherratt,et al.  Periodic travelling waves in cyclic predator–prey systems , 2001 .

[57]  Janis Antonovics,et al.  Parasite–grass–forb interactions and rock–paper– scissor dynamics: predicting the effects of the parasitic plant Rhinanthus minor on host plant communities , 2009 .

[58]  Kimberly A. With The Landscape Ecology of Invasive Spread , 2002 .

[59]  D. Tilman Resource competition and community structure. , 1983, Monographs in population biology.

[60]  N. Shigesada,et al.  Diffusive waves, dynamical stabilization and spatio-temporal chaos in a community of three competitive species , 2001 .

[61]  S. Merino Cyclic competition of three species in the time periodic and diffusive case , 1996 .

[62]  G. E. Hutchinson,et al.  The Balance of Nature and Human Impact: The paradox of the plankton , 2013 .

[63]  A. Ōkubo,et al.  On the spatial spread of the grey squirrel in Britain , 1989, Proceedings of the Royal Society of London. B. Biological Sciences.

[64]  Daniel Simberloff,et al.  Invasional meltdown 6 years later: important phenomenon, unfortunate metaphor, or both? , 2006, Ecology letters.

[65]  Robert A Laird,et al.  Does local competition increase the coexistence of species in intransitive networks? , 2008, Ecology.

[66]  Alfred W. Crosby,et al.  Ecological Imperialism: The Biological Expansion of Europe, 900-1900 , 1988 .

[67]  William Gurney,et al.  Circles and spirals: population persistence in a spatially explicit predator-prey model , 1998 .

[68]  G. Hardin The competitive exclusion principle. , 1960, Science.

[69]  M. Gilpin Limit Cycles in Competition Communities , 1975, The American Naturalist.

[70]  Maarten C. Boerlijst,et al.  Evolutionary consequences of spiral waves in a host—parasitoid system , 1993, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[71]  L. Buss,et al.  Competitive Networks: Nontransitive Competitive Relationships in Cryptic Coral Reef Environments , 1979, The American Naturalist.

[72]  G D Ruxton,et al.  Spatial self-organisation in ecology: pretty patterns or robust reality? , 1997, Trends in ecology & evolution.

[73]  Janneke HilleRisLambers,et al.  The importance of niches for the maintenance of species diversity , 2009, Nature.

[74]  Kei-ichi Tainaka,et al.  Paradoxical effect in a three-candidate voter model , 1993 .

[75]  T. Reichenbach,et al.  Mobility promotes and jeopardizes biodiversity in rock–paper–scissors games , 2007, Nature.

[76]  J. Huisman,et al.  Towards a solution of the plankton paradox : the importance of physiology and life history , 2001 .

[77]  L. Buss,et al.  Alleopathy and spatial competition among coral reef invertebrates. , 1975, Proceedings of the National Academy of Sciences of the United States of America.

[78]  C. Hauert,et al.  Replicator dynamics for optional public good games. , 2002, Journal of theoretical biology.

[79]  Benjamin Kerr,et al.  THE EVOLUTION OF RESTRAINT IN BACTERIAL BIOFILMS UNDER NONTRANSITIVE COMPETITION , 2008, Evolution; international journal of organic evolution.

[80]  P. Levin,et al.  COMMUNITY‐WIDE EFFECTS OF NONINDIGENOUS SPECIES ON TEMPERATE ROCKY REEFS , 2002 .