New insights into island vegetation composition and species diversity—Consistent and conditional responses across contrasting insular habitats at the plot-scale

Most island-ecology studies focus on the properties of entire island communities, thus neglecting species-environment relationships operating at the habitat-level. Habitat-specific variation in the strength and sign of these relationships will conceal patterns observed on the island scale and may preclude a mechanistic interpretation of patterns and processes. Habitat-specific species-environment relationships may also depend on the descriptor of ecological communities. This paper presents a comprehensive plot-based analysis of local vegetation composition and species diversity (species richness and species evenness) of (i) rocky shore, (ii) semi-natural grassland and (iii) coniferous forest habitats in three Baltic archipelagos in Sweden. To identify differences and consistencies between habitats and descriptors, we assessed the relative contributions of the variable-sets “region”, “topography”, “soil morphology”, “soil fertility”, “soil water”, “light availability”, “distance” and “island configuration” on local vegetation composition, species richness and species evenness. We quantified the impact of “management history” on the descriptors of local grassland communities by a newly introduced grazing history index (GHI). Unlike species diversity, changes in vegetation composition were related to most of the variable-sets. The relative contributions of the variable-sets were mostly habitat-specific and strongly contingent on the descriptor involved. Within each habitat, richness and evenness were only partly affected by the same variable-sets, and if so, their relative contribution varied between diversity proxies. Across all habitats, soil variable-sets showed highly consistent effects on vegetation composition and species diversity and contributed most to the variance explained. GHI was a powerful predictor, explaining high proportions of variation in all three descriptors of grassland species communities. The proportion of unexplained variance was habitat-specific, possibly reflecting a community maturity gradient. Our results reveal that species richness alone is an incomplete representation of local species diversity. Finally, we stress the need of including habitat-based approaches when analyzing complex species-environment relationships on islands.

[1]  Michael J. Crawley,et al.  The R book , 2022 .

[2]  U. Gärdenfors Biodiversity in Sweden , 2018, Global Biodiversity.

[3]  Per B. Brockhoff,et al.  lmerTest Package: Tests in Linear Mixed Effects Models , 2017 .

[4]  M. Jafari,et al.  Vegetation composition differentiation and species-environment relationships in the northern part of Isfahan Province, Iran , 2017, Journal of Arid Land.

[5]  R. Corlett Plant diversity in a changing world: Status, trends, and conservation needs , 2016, Plant diversity.

[6]  R. Ricklefs,et al.  Region effects influence local tree species diversity , 2016, Proceedings of the National Academy of Sciences.

[7]  A. Kallimanis,et al.  Comparing the two Greek archipelagos plant species diversity and endemism patterns highlight the importance of isolation and precipitation as biodiversity drivers , 2014, Journal of Biological Research-Thessaloniki.

[8]  U. Bergström,et al.  Shoreline development and degradation of coastal fish reproduction habitats , 2014, AMBIO.

[9]  Jonathan William Gillard,et al.  The R book, second edition , 2014 .

[10]  Shinichi Nakagawa,et al.  A general and simple method for obtaining R2 from generalized linear mixed‐effects models , 2013 .

[11]  Shurong Zhou,et al.  The Relationship between Species Richness and Evenness in Plant Communities along a Successional Gradient: A Study from Sub-Alpine Meadows of the Eastern Qinghai-Tibetan Plateau, China , 2012, PloS one.

[12]  R. Lindborg,et al.  Landscape context and management regime structure plant diversity in grassland communities , 2012 .

[13]  S. Cousins,et al.  Landscape structure and land use history influence changes in island plant composition after 100 years , 2012 .

[14]  Stefan Sperlich,et al.  Explaining grassland biomass – the contribution of climate, species and functional diversity depends on fertilization and mowing frequency , 2011 .

[15]  M. Numers Sea Shore Plants of the SW Archipelago of Finland — Distribution Patterns and Long-Term Changes during the 20th Century , 2011 .

[16]  J. Thiele,et al.  Competitive displacement or biotic resistance? Disentangling relationships between community diversity and invasion success of tall herbs and shrubs , 2010 .

[17]  A. G. Endress,et al.  Relative influence of landscape vs. local factors on plant community assembly in restored wetlands. , 2009, Ecological applications : a publication of the Ecological Society of America.

[18]  S. Cousins Landscape history and soil properties affect grassland decline and plant species richness in rural landscapes. , 2009 .

[19]  C. Gries,et al.  Frontiers inEcology and the Environment Effects of urbanization on plant species diversity in central Arizona , 2009 .

[20]  Honor C. Prentice,et al.  Small-scale plant species richness and evenness in semi-natural grasslands respond differently to habitat fragmentation. , 2009 .

[21]  Mollie E. Brooks,et al.  Generalized linear mixed models: a practical guide for ecology and evolution. , 2009, Trends in ecology & evolution.

[22]  R. Ricklefs,et al.  The Theory of Island Biogeography Revisited , 2009 .

[23]  P. Legendre,et al.  Forward selection of explanatory variables. , 2008, Ecology.

[24]  Jens-Johan Hannus,et al.  ORIGINAL ARTICLE: Vascular plant species richness in relation to habitat diversity and island area in the Finnish Archipelago , 2008 .

[25]  M. Zobel,et al.  Plant diversity in a calcareous wooded meadow – The significance of management continuity , 2008 .

[26]  M. Numers,et al.  20th century vegetation changes in an island archipelago, SW Finland , 2007 .

[27]  Eva Gustavsson,et al.  Land use more than 200 years ago explains current grassland plant diversity in a Swedish agricultural landscape , 2007 .

[28]  J. H. Bock,et al.  Relationships between species richness, evenness, and abundance in a southwestern savanna. , 2007, Ecology.

[29]  Y. Qian,et al.  Environmental factors affecting vegetation composition in the Alxa Plateau, China , 2007 .

[30]  G. Stirling,et al.  Species richness and evenness respond in a different manner to propagule density in developing prairie microcosm communities , 2007, Plant Ecology.

[31]  R. Heikkinen,et al.  Impacts of local and regional factors on vegetation of boreal semi-natural grasslands , 2007, Plant Ecology.

[32]  Johannes Isselstein,et al.  Effects of local factors on plant species richness and composition of Alpine meadows , 2007 .

[33]  Johannes Isselstein,et al.  Plant species richness and composition in managed grasslands: The relative importance of field management and environmental factors , 2007 .

[34]  Peter D. Moore,et al.  The ecology of plants, 2nd edn. , 2007 .

[35]  H. Bratli,et al.  Scale-dependent importance of environment, land use and landscape structure for species richness and composition of SE Norwegian modern agricultural landscapes , 2006, Landscape Ecology.

[36]  J. Witman,et al.  Nutrient transfer from sea to land: the case of gulls and cormorants in the Gulf of Maine. , 2006, The Journal of animal ecology.

[37]  Maohua Ma Species richness vs evenness: independent relationship and different responses to edaphic factors , 2005 .

[38]  M. Willig,et al.  Relationships Among Indices Suggest that Richness Is an Incomplete Surrogate for Grassland Biodiversity , 2005 .

[39]  R. McMaster Factors influencing vascular plant diversity on 22 islands off the coast of eastern North America , 2005 .

[40]  Tapio Suominen,et al.  A GIS-based step-wise procedure for assessing physical exposure in fragmented archipelagos , 2003 .

[41]  Johan Ehrlén,et al.  Habitat configuration, species traits and plant distributions , 2002 .

[42]  A. Löfgren,et al.  Species richness, extinction and immigration rates of vascular plants on islands in the Stockholm archipelago, Sweden, during a century of ceasing management , 2002, Folia Geobotanica.

[43]  O. Eriksson,et al.  The influence of management history and habitat on plant species richness in a rural hemiboreal landscape, Sweden , 2002, Landscape Ecology.

[44]  B. McCune,et al.  Analysis of Ecological Communities , 2002 .

[45]  S. Carpenter,et al.  Catastrophic shifts in ecosystems , 2001, Nature.

[46]  G. Stirling,et al.  Empirical Relationships between Species Richness, Evenness, and Proportional Diversity , 2001, The American Naturalist.

[47]  F. Chapin,et al.  Consequences of changing biodiversity , 2000, Nature.

[48]  Andrew F. Brown,et al.  Biodiversity: An Introduction , 1999, Biodiversity & Conservation.

[49]  Michael A. Huston,et al.  Local processes and regional patterns : appropriate scales for understanding variation in the diversity of plants and animals , 1999 .

[50]  R. Gillespie,et al.  Island Biogeography — Ecology, Evolution and Conservation , 1999, Heredity.

[51]  E. Maarel,et al.  Plant distribution patterns and ecological gradients in the Southwest Finnish archipelago , 1998 .

[52]  P. Reich,et al.  The Influence of Functional Diversity and Composition on Ecosystem Processes , 1997 .

[53]  A. Cooper Plant species coexistence in cliff habitats , 1997 .

[54]  G. Tyler Cover distributions of vascular plants in relation to soil chemistry and soil depth in a granite rock ecosystem , 1996, Vegetatio.

[55]  Benjamin Smith,et al.  A consumer's guide to evenness indices , 1996 .

[56]  M. Diekmann Use and improvement of Ellenberg's indicator values in deciduous forests of the Boreo-nemoral zone in Sweden , 1995 .

[57]  R. Wetzel,et al.  Nutrient additions by waterfowl to lakes and reservoirs: predicting their effects on productivity and water quality , 1994, Hydrobiologia.

[58]  L. Gustafsson A comparison of biological characteristics and distribution between Swedish threatened and non‐threatened forest vascular plants , 1994 .

[59]  Rune Halvorsen Økland,et al.  Canonical Correspondence Analysis with variation partitioning: some comments and an application , 1994 .

[60]  H. Ronald Pulliam,et al.  Island Biogeography: Effect of Geographical Isolation on Species Composition , 1993 .

[61]  P. Morisset,et al.  Floristic richness, area, and habitat diversity in a hemiarctic archipelago , 1988 .

[62]  R. Buckley,et al.  Distinguishing the Effects of Area and Habitat Type on Island Plant Species Richness by Separating Floristic Elements and Substrate Types and Controlling for Island Isolation , 1985 .

[63]  D. Simberloff,et al.  Environmental Determinants of Island Species Numbers in the British Isles , 1974 .

[64]  J. Braun-Blanquet,et al.  Pflanzensoziologie: Grundzuge der Vegetationskunde. , 1967 .

[65]  Vivi Fleming-Lehtinen,et al.  Recent developments in assessment methodology reveal that the Baltic Sea eutrophication problem is expanding , 2015 .

[66]  C.J.F. ter Braak,et al.  Canoco Reference Manual and User’s Guide: Software for Ordination (version 5.0) , 2012 .

[67]  E. Glazkova Ornithocoprophilous flora and vegetation of the islands in the Gulf of Finland, the Baltic Sea. , 2009 .

[68]  Mark S. Fonseca,et al.  WEMo (Wave Exposure Model): Formulation, Procedures and Validation , 2007 .

[69]  A. Solow,et al.  Measuring biological diversity , 2006, Environmental and Ecological Statistics.

[70]  Mathew A. Leibold,et al.  Metacommunities: Spatial Dynamics and Ecological Communities , 2005 .

[71]  Richard J. Hobbs,et al.  Assembly rules and restoration ecology : bridging the gap between theory and practice , 2004 .

[72]  Jessica Gurevitch,et al.  The ecology of plants , 2002 .

[73]  Petr Šmilauer,et al.  CANOCO 4.5 Reference Manual and CanoDraw for Windows User's Guide: Software for Canonical Community Ordination , 2002 .

[74]  Marc J. Mazerolle,et al.  Patch characteristics and landscape context as predictors of species presence and abundance: A review1 , 1999 .

[75]  Pol Coppin,et al.  Effects of area, age and diversity of forest patches in Belgium on plant species richness, and implications for conservation and reforestation , 1999 .

[76]  M. Diekmann,et al.  Swedish plant geography , 1999 .

[77]  B. Pavlik Species Diversity in Ecological Communities: Historical and Geographical Perspectives by R. E. Ricklefs, D. Schluter , 1995 .

[78]  D. Kohn,et al.  Plant species richness: the effect of island size and habitat diversity , 1994 .

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

[80]  Ruprecht Düll,et al.  Zeigerwerte von Pflanzen in Mitteleuropa , 1992 .

[81]  K. Johannesson The Bare Zone of Swedish Rocky Shores: Why Is It There? , 1989 .

[82]  E. Ranta,et al.  Patterns and processes in species assemblages on Northern Baltic islands , 1987 .

[83]  Berger‐Landefeldt Braun-Blanquet, J.: Pflanzensoziologie. Grundzüge der Vegetationskunde. 2. Aufl. Wien 1951 (Springer-Verlag). 2. Aufl. 631 S. mit 350 Textabb. DM 63.—, Ganzleinen DM 67.20 , 1952 .