Effects of competition mode on spatial pattern dynamics in plant communities

Abstract The effects of the mode of competition between individual plants (symmetric versus asymmetric) and the gap formation caused by natural disturbances on the dynamics of spatial configuration pattern of individual size were investigated theoretically based on an individual growth model incorporating competitive effects of neighbouring individuals. The degree of spatial heterogeneity in local size distribution was represented by the CV (coefficient of variation) of averages of local size distributions (CVav), the average of CVs of local size distributions (AVcv) and the CV of CVs of local size distributions (CVcv). Without gap formation, CVav, AVcv and CVcv were larger under asymmetric competition than under symmetric one, suggesting a fine-scale mosaic spatial pattern in asymmetrically competing populations. With gap formation, these statistics under symmetric competition approached those values under asymmetric competition. The spatial configuration pattern of individual size also showed the same trend in terms of the semivariogram. The patchiness index was almost the same in both gap and non-gap cases irrespective of the mode of competition. The semivariogram and patchiness index showed the presence of more uniform and larger patches under symmetric competition than asymmetric competition. Gap formation therefore increased spatial heterogeneity in local size distribution especially under symmetric competition, but there was still a difference in spatial heterogeneity between the two modes of competition even in the gap formation case. The effects of gap formation on spatial pattern dynamics were larger under symmetric competition than under asymmetric competition; under asymmetric competition, the spatial pattern dynamics were similar in both gap and non-gap cases. Therefore, against spatial disturbances (i.e. gap formation), symmetric competition brings about a more variable system than asymmetric competition. These theoretical results can explain spatial pattern dynamics of natural forests (northern coniferous and temperate hardwood forests). In conclusion, both the disturbance regime (gap formation process) and the mode of competition between individuals should be investigated to study the spatial pattern dynamics and species diversity of plant communities. The implications of the mode of between-individual competition for conservation biology are discussed. It is suggested that symmetrically competing plant communities (e.g. northern coniferous forests) should be preserved in larger areas than asymmetrically competing ones (e.g. temperate hardwood forests) if the plant communities are subject to frequent natural disturbances.

[1]  M. Yokozawa,et al.  Foliage Profile, Size Structure and Stem Diameter-Plant Height Relationship in Crowded Plant Populations , 1995 .

[2]  Tree Competition and Species Coexistence in a Sub-boreal Forest, Northern Japan , 1995 .

[3]  J. Denslow Chapter 17 – Disturbance-Mediated Coexistence of Species , 1985 .

[4]  R. Macarthur PATTERNS OF SPECIES DIVERSITY , 1965 .

[5]  Jacob Weiner,et al.  A Neighborhood Model of Annual‐Plant Interference , 1982 .

[6]  Steward T. A. Pickett,et al.  Patch dynamics and the design of nature reserves , 1978 .

[7]  D. Schneider,et al.  Analysis of scale-dependent processes with dimensionless ratios , 1994 .

[8]  Jerry F. Franklin,et al.  Gap Characteristics and Vegetation Response in Coniferous Forests of the Pacific Northwest , 1989 .

[9]  Robert M. May,et al.  Large-Scale Ecology and Conservation Biology. , 1995 .

[10]  K. Umeki,et al.  Effect of Canopy Structure on Degree of Asymmetry of Competition in Two Forest Stands in Northern Japan , 1996 .

[11]  James R. Runkle,et al.  PATTERNS OF DISTURBANCE IN SOME OLD-GROWTH MESIC FORESTS OF EASTERN NORTH AMERICA' , 1982 .

[12]  L. Lundqvist Growth and competition in partially cut sub-alpine Norway spruce forests in northern Sweden , 1994 .

[13]  Noel A Cressie,et al.  Statistics for Spatial Data. , 1992 .

[14]  Toshihiko Hara,et al.  A Canopy Photosynthesis Model for the Dynamics of Size Structure and Self-thinning in Plant Populations , 1992 .

[15]  Clayton V. Deutsch,et al.  GSLIB: Geostatistical Software Library and User's Guide , 1993 .

[16]  T. Nakashizuka REGENERATION PROCESS OF CLIMAX BEECH (FAGUS CRENATA BLUME) FORESTS : V. POPULATION DYNAMICS OF BEECH IN A REGENERATION PROCESS , 1984 .

[17]  T. Hara Mode of Competition and Size‐structure Dynamics in Plant Communities , 1993 .

[18]  Tree competition and species coexistence in a cool‐temperate old‐growth forest in southwestern Japan , 1995 .

[19]  R. Primack,et al.  Essentials of Conservation Biology , 1994 .

[20]  T. Veblen Tree Regeneration Responses to Gaps Along a Transandean Gradient , 1989 .

[21]  G. Sugihara Minimal Community Structure: An Explanation of Species Abundance Patterns , 1980, The American Naturalist.

[22]  Kunihiko Kaneko,et al.  Theory and Applications of Coupled Map Lattices , 1993 .

[23]  P. White,et al.  The Ecology of Natural Disturbance and Patch Dynamics , 1986 .

[24]  J. Connell Diversity in tropical rain forests and coral reefs. , 1978, Science.

[25]  James R. Runkle,et al.  Gap dynamics in an Ohio Acer–Fagus forest and speculations on the geography of disturbance , 1990 .

[26]  Tomáš Herben,et al.  Small‐scale spatial dynamics of plant species in a grassland community over six years , 1993 .

[27]  Makoto Kimura,et al.  GROWTH PATTERNS OF TREE HEIGHT AND STEM DIAMETER IN POPULATIONS OF ABIES VEITCHII, A. MARIESII AND BETULA ERMANII , 1991 .

[28]  J. Weiner,et al.  Asymmetric competition in plant populations. , 1990, Trends in ecology & evolution.

[29]  R. Ricklefs,et al.  Community Diversity: Relative Roles of Local and Regional Processes , 1987, Science.

[30]  T. C. Whitmore,et al.  Canopy Gaps and the Two Major Groups of Forest Trees , 1989 .

[31]  T. Kohyama Simulating Stationary Size Distribution of Trees in Rain Forests , 1991 .

[32]  H. Sakaguchi Hierarchical structures in Lotka-Volterra type equations , 1994 .

[33]  Mike Rees,et al.  5. Statistics for Spatial Data , 1993 .

[34]  Peter J. Diggle,et al.  Competition for Light in a Plant Monoculture Modelled as a Spatial Stochastic Process , 1981 .

[35]  Jacob Weiner,et al.  Growth Variation in a Naturally Established Population of Pinus Sylvestris , 1994 .

[36]  S. Pacala,et al.  Neighborhood Models of Plant Population Dynamics. I. Single-Species Models of Annuals , 1985, The American Naturalist.

[37]  J. Jackson Adaptation and Diversity of Reef CoralsPatterns result from species differences in resource use and life histories and from disturbances , 1991 .

[38]  Jacqueline McGlade,et al.  A coupled map lattice model of the growth of plant monocultures , 1996 .

[39]  T. Nakashizuka REGENERATION PROCESS OF CLIMAX BEECH (FAGUS CRENATA BLUME) FORESTS : IV. GAP FORMATION , 1984 .

[40]  T. Nakashizuka Regeneration dynamics of beech forests in Japan , 1987 .

[41]  T. Kohyama,et al.  Size-structured tree populations in gap-dynamic forest-the forest architecture hypothesis for the stable coexistence of species , 1993 .

[42]  P. White,et al.  Scale Dependence and the Species-Area Relationship , 1994, The American Naturalist.

[43]  J. Lawton,et al.  Species interactions, local and regional processes, and limits to the richness of ecological communities : a theoretical perspective , 1992 .

[44]  R. Paine,et al.  Disturbance, patch formation, and community structure. , 1974, Proceedings of the National Academy of Sciences of the United States of America.

[45]  A. Magurran,et al.  Biological diversity : the coexistence of species on changing landscapes , 1994 .

[46]  Tomasz Wyszomirski,et al.  Competitive Asymmetry Reduces Spatial Effects on Size-Structure Dynamics in Plant Populations , 1994 .

[47]  Y. Iwasa,et al.  Forest gap dynamics with partially synchronized disturbances and patch age distribution , 1995 .