Plant species segregation on different spatial scales in semi-natural grasslands

Abstract Question: We studied the patterns of plant species co-occurrence on three, nested, spatial scales in semi-natural grassland communities and explored the possible ecological processes underlying the patterns. Location: Dry, semi-natural grasslands in a 4.5 km × 4.5 km area on the Baltic Island of Öland (Sweden). Methods: The study used replicated samples on three, nested, spatial scales: 50 cm × 50 cm plots (N = 516), grassland patches (N = 109) and the whole landscape (N = 6). We used a null model approach to study species co-occurrence patterns and compared the ecological amplitudes of the pairs of species contributing most to the patterns. We used linear models to search for associations between species segregation and environmental and landscape factors. Results and Conclusions: Our results support the prediction that patterns of species co-occurrence are likely to be influenced by different mechanisms on different spatial scales. On the plot scale, we interpreted the species segregation in terms of species interactions. The degree of species segregation was significantly associated with the plots' positions within the grassland patches (edge effects) and with management intensity of the grasslands – both variables can be assumed to influence species interactions. On the grassland patch scale, we interpreted the species segregation in terms of within-patch environmental heterogeneity. The degree of segregation was significantly associated with the area of the grassland patches and with management intensity – both variables that are likely to be related to environmental heterogeneity within the grasslands. Species segregation on the landscape scale was interpreted in terms of environmental heterogeneity among grassland patches and was significantly associated with land-use history. Nomenclature: Karlsson (1998).

[1]  N. Gotelli,et al.  Assembly of avian mixed-species flocks in Amazonia. , 1993, Proceedings of the National Academy of Sciences of the United States of America.

[2]  Lewi Stone,et al.  Competitive exclusion, or species aggregation? , 1992, Oecologia.

[3]  O. Eriksson,et al.  Size‐related deterioration of semi‐natural grassland fragments in Sweden , 2002 .

[4]  David Mouillot,et al.  Parasite species coexistence and limiting similarity: a multiscale look at phylogenetic, functional and reproductive distances , 2005, Oecologia.

[5]  T. Karlsson Förteckning över svenska kärlväxter , 1997 .

[6]  L. Stone,et al.  The checkerboard score and species distributions , 1990, Oecologia.

[7]  Karen H. Beard,et al.  Soil History as a Primary Control on Plant Invasion in Abandoned Agricultural Fields , 2006 .

[8]  Paul A. Keddy,et al.  Community Assembly Rules, Morphological Dispersion, and the Coexistence of Plant Species , 1998 .

[9]  F. G. Araújo,et al.  Co-occurrence of demersal fishes in a tropical bay in southeastern Brazil: A null model analysis , 2006 .

[10]  N. Gotelli Null model analysis of species co-occurrence patterns , 2000 .

[11]  S. Harrison LOCAL AND REGIONAL DIVERSITY IN A PATCHY LANDSCAPE: NATIVE, ALIEN, AND ENDEMIC HERBS ON SERPENTINE , 1999 .

[12]  M. Zobel,et al.  The relative of species pools in determining plant species richness: an alternative explanation of species coexistence? , 1997, Trends in ecology & evolution.

[13]  M. Hill,et al.  Detrended correspondence analysis: an improved ordination technique , 1980 .

[14]  H. H. Bruun,et al.  Community assembly in experimental grasslands: suitable environment or timely arrival? , 2006, Ecology.

[15]  E. Gianoli,et al.  Species richness and structure of ant communities in a dynamic archipelago: effects of island area and age , 2005 .

[16]  Martin T. Sykes,et al.  Small-scale plant species turnover in a limestone grassland: the carousel model and some comments on the niche concept. , 1993 .

[17]  U. Ekstam,et al.  If grassland management ceases - vascular plants as indicator species in meadows and pastures , 1992 .

[18]  Pedro R. Peres-Neto,et al.  Patterns in the co-occurrence of fish species in streams: the role of site suitability, morphology and phylogeny versus species interactions , 2004, Oecologia.

[19]  Honor C. Prentice,et al.  Semi-natural grassland continuity, long-term land-use change and plant species richness in an agricultural landscape on Öland, Sweden , 2008 .

[20]  George R. Parker,et al.  Relationships between landcover proportion and indices of landscape spatial pattern , 1992, Landscape Ecology.

[21]  E. Maarel,et al.  Restoration of alvar vegetation on Öland, Sweden , 2000 .

[22]  M. Hermy,et al.  The land use history (1278–1990) of a mixed hardwood forest in western Belgium and its relationship with chemical soil characteristics , 1999 .

[23]  P. Poschlod,et al.  Plant dispersal potential and its relation to species frequency and coexistence , 2005 .

[24]  Paul A. Keddy,et al.  Assembly rules, null models, and trait dispersion : new questions from old patterns , 1995 .

[25]  Miska Luoto,et al.  Plant species richness and persistence of rare plants in abandoned semi-natural grasslands in northern Europe , 2005 .

[26]  D. Franzen Plant species coexistence and dispersion of seed traits in a grassland , 2004 .

[27]  Marco A. Molina-Montenegro,et al.  Slope aspect influences plant association patterns in the Mediterranean matorral of central Chile , 2005 .

[28]  C. B. Williams Area and Number of Species , 1943, Nature.

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

[30]  M. Sykes,et al.  Time and space in the community structure of a species-rich limestone grassland , 1995 .

[31]  D. Jenkins In search of quorum effects in metacommunity structure: species co-occurrence analyses. , 2006, Ecology.

[32]  Nicholas J. Gotelli,et al.  SPECIES CO‐OCCURRENCE: A META‐ANALYSIS OF J. M. DIAMOND'S ASSEMBLY RULES MODEL , 2002 .

[33]  J. Brunet,et al.  Land use effects on soil N, P, C and pH persist over 40-80 years of forest growth on agricultural soils , 2006 .

[34]  Wolfgang Cramer,et al.  THE PLANT COMMUNITY AS A NICHE BIOASSAY: ENVIRONMENTAL CORRELATES OF LOCAL VARIATION IN GYPSOPHILA FASTIGIATA , 1990 .

[35]  R. Hobbs,et al.  Biological Consequences of Ecosystem Fragmentation: A Review , 1991 .

[36]  Graham Bell,et al.  THE CO‐DISTRIBUTION OF SPECIES IN RELATION TO THE NEUTRAL THEORY OF COMMUNITY ECOLOGY , 2005 .

[37]  Daniel Simberloff,et al.  The Assembly of Species Communities: Chance or Competition? , 1979 .

[38]  M. Diekmann,et al.  Differences in species richness and life‐history traits between grazed and abandoned grasslands in southern Sweden , 2001 .

[39]  M. Sykes,et al.  Fragmented grasslands on the Baltic Island of Öland. Plant community composition and land-use history , 2008 .

[40]  P. Grubb THE MAINTENANCE OF SPECIES‐RICHNESS IN PLANT COMMUNITIES: THE IMPORTANCE OF THE REGENERATION NICHE , 1977 .