Integrated screening validates primary axes of specialisation in plants

Standardised procedures have been used to measure 67 traits in 43 common plants of the British flora. This paper provides an interpretation of the most consistent patterns in the resulting matrix by means of correlation, ordination and classification analyses. Only a weak coupling was observed between attributes of the regenerative and established phases of the life history. However, within each phase, attributes were strongly aggregated into sets and a high proportion of the variation between species coincided with a single axis. Attributes of the established phase displayed remarkably consistent trends, with a strong 'Axis I' being identified by three different multivariate methods. There was a marked correlation between foliar concentrations of N, P, K, Ca and Mg, high concentrations of which coincided with the capacity for rapid growth in productive conditions and an inability to sustain yield under limiting supplies of nutrients. A diverse array of other traits, less immediately involving mineral nutrients, were also entrained in Axis I; these included life history, root and shoot foraging, the morphology, longevity, tensile strength and palatability of leaves, and the decomposition rate of leaf litter. This pattern occurred in both monocotyledons and dicotyledons and appeared to reflect a tradeoff between attributes conferring an ability for high rates of resource acquisition in productive habitats and those responsible for retention of resource capital in unproductive conditions. The second axis of variation evident in the established phase was related to phylogeny and distinguished between monocotyledons and dicotyledons on the basis of a diverse set of traits including genome size, cell size, root and shoot foraging characteristics and vascular tissues. A third axis was detected in which ephemerals and perennials were separated by differences in attributes such as breeding system, leaf decomposition rate and a set of traits reflecting the small stature of many short-lived plants. In the regenerative phase, the leading axis was clearly related to the widely recognized tradeoff between seed size and seed number and was consistent with current understanding of seed banks, and with modern theories explaining species coexistence in terms of complementary responses to temporal and spatial variation in vegetation gap dynamics. The data provide strong evidence of functional integration between evolutionary specializations in root and shoot and support Donald's unified theory of competitive ability. The data are not consistent with theories of functional types based upon evolutionary tradeoffs in allocation between root and shoot. We suggest that the evidence assembled here and elsewhere in the current literature points to the existence of primary functional types, including those recognized by Ramenskii and Grime. These functional types can be reconciled with the individuality of plant ecologies in the field and provide an effective basis for interpretation and prediction at various scales from the plant community to regional floras. There are particular opportunities for prediction of successional trajectories, the role of herbivores in vegetation succession and the response of vegetation to eutrophication and extreme climatic events. It is also suggested that aspects of this investigation may provide a Darwinian underpinning for Odum's theory of ecosystem maturation.

[1]  J. Cornelissen,et al.  Functional leaf attributes predict litter decomposition rate in herbaceous plants. , 1997, The New phytologist.

[2]  Ken Thompson,et al.  Evidence of a causal connection between anti-herbivore defence and the decomposition rate of leaves , 1996 .

[3]  Roderick Hunt,et al.  Seedling growth, allocation and leaf attributes in a wide range of woody plant species and types , 1996 .

[4]  J. Cornelissen,et al.  An experimental comparison of leaf decomposition rates in a wide range of temperate plant species and types , 1996 .

[5]  Ken Thompson,et al.  A functional analysis of a limestone grassland community , 1996 .

[6]  J. Bakker,et al.  The Soil Seed Banks of North West Europe: Methodology, Density and Longevity , 1996 .

[7]  F. Kienast,et al.  Classifying the life-history strategies of trees on the basis of the Grimian model , 1994 .

[8]  K. Thompson,et al.  Seed Persistence: A Correlation Between Seed Longevity in the Soil and Ortho-Dihydroxyphenol Concentration , 1994 .

[9]  J. Hills,et al.  A Method for Classifying European Riverine Wetland Ecosystems Using Functional Vegetation Groups , 1994 .

[10]  J. P. Grime,et al.  Ellenberg numbers revisited , 1993 .

[11]  Roderick Hunt,et al.  Further responses to CO2 enrichment in British herbaceous species. , 1993 .

[12]  J. P. Grime.  Ecology Sans Frontières@@@Ecology Sans Frontieres , 1993 .

[13]  J. P. Grime,et al.  Methods in comparative plant ecology : a laboratory manual , 1993 .

[14]  O. Sala,et al.  Plant functional types and ecological strategies in Patagonian forbs , 1993 .

[15]  W. Foulds.  Nutrient concentrations of foliage and soil in South-western Australia. , 1993, The New phytologist.

[16]  Céline Boutin,et al.  A functional classification of wetland plants , 1993 .

[17]  J. P. Grime,et al.  A Comparative Study of Plasticity in Seedling Rooting Depth in Drying Soil , 1993 .

[18]  J. Hodgson.  Commonness and rarity in British butterflies , 1993 .

[19]  H. Clifford,et al.  The taxonomic and evolutionary significance of leaf longevity , 1993 .

[20]  Ken Thompson,et al.  Seed size and shape predict persistence in soil , 1993 .

[21]  J. P. Grime,et al.  Resource dynamics and vegetation processes: a deterministic model using two-dimensional cellular automata , 1993 .

[22]  Spatial pattern of seedling recruitment at two different scales in a limestone grassland , 1992 .

[23]  Michelle R. Leishman,et al.  Classifying plants into groups on the basis of associations of individual traits: evidence from Australian semi-arid woodlands , 1992 .

[24]  Martin Kent,et al.  Vegetation Description and Analysis: A Practical Approach , 1992 .

[25]  B. Shipley,et al.  The Allometry of Seed Production in Herbaceous Angiosperms , 1992, The American Naturalist.

[26]  P. Reich,et al.  Leaf Life‐Span in Relation to Leaf, Plant, and Stand Characteristics among Diverse Ecosystems , 1992 .

[27]  S. Stearns,et al.  The Evolution of Life Histories , 1992 .

[28]  P. Keddy.  A pragmatic approach to functional ecology , 1992 .

[29]  K. Reiling,et al.  The response of native, herbaceous species to ozone: growth and fluorescence screening , 1992 .

[30]  Hendrik Poorter,et al.  Inherent Variation in Growth Rate Between Higher Plants: A Search for Physiological Causes and Ecological Consequences , 1992 .

[31]  C. Stace.  New Flora Of The British Isles , 1998 .

[32]  M. Westoby,et al.  Diaspore weight, dispersal, growth form and perenniality of central australian plants , 1991 .

[33]  D. Charlesworth,et al.  Allocation of Resources to Sex Functions in Flowering Plants , 1991 .

[34]  R. Sibly.  The Life-History Approach to Physiological Ecology , 1991 .

[35]  M. Pagel,et al.  The comparative method in evolutionary biology , 1991 .

[36]  David Tilman,et al.  Allocation and the transient dynamics of succession on poor soils. , 1990 .

[37]  H. Lambers,et al.  Causes and consequences of variation in growth rate and productivity of higher plants , 1990 .

[38]  M. Sykes,et al.  Dark tolerance in plants of dunes , 1990 .

[39]  I. Washitani,et al.  A Comparative Study of the Germination Characteristics of Seeds from a Moist Tall Grassland Community , 1990 .

[40]  Bill Shipley,et al.  Regeneration and Establishment Strategies of Emergent Macrophytes , 1989 .

[41]  Roderick Hunt,et al.  Comparative Plant Ecology: A Functional Approach to Common British Species , 1989 .

[42]  D. Tilman,et al.  Growth of old field herbs on a nitrogen gradient , 1989 .

[43]  R. Sibly.  What evolution maximizes , 1989 .

[44]  W. Krzanowski,et al.  A Criterion for Determining the Number of Groups in a Data Set Using Sum-of-Squares Clustering , 1988 .

[45]  A. E. Ainsley,et al.  Genstat 5 Reference Manual , 1987 .

[46]  Thomas M. Smith,et al.  Plant Succession: Life History and Competition , 1987, The American Naturalist.

[47]  M. Hill,et al.  Data analysis in community and landscape ecology , 1987 .

[48]  David Tilman,et al.  Secondary Succession and the Pattern of Plant Dominance Along Experimental Nitrogen Gradients , 1987 .

[49]  J. P. Grime.  Dominant and subordinate components of plant communities: implications for succession, stability and diversity , 1987 .

[50]  Thomas J. Givnish,et al.  On the economy of plant form and function. , 1988 .

[51]  J. Hodgson,et al.  THE ECOLOGICAL SPECIALIZATION OF DICOTYLEDONOUS FAMILIES WITHIN A LOCAL FLORA: SOME FACTORS CONSTRAINING OPTIMIZATION OF SEED SIZE AND THEIR POSSIBLE EVOLUTIONARY SIGNIFICANCE , 1986 .

[52]  C. Braak.  Canonical Correspondence Analysis: A New Eigenvector Technique for Multivariate Direct Gradient Analysis , 1986 .

[53]  J. P. Grime.  Factors limiting the contribution of pteridophytes to a local flora , 1985 .

[54]  J. P. Grime.  THE ECOLOGY OF SPECIES, FAMILIES AND COMMUNITIES OF THE CONTEMPORARY BRITISH FLORA. , 1984, The New phytologist.

[55]  Y. Iwasa,et al.  Shoot/root balance of plants: Optimal growth of a system with many vegetative organs , 1984 .

[56]  P. Greenslade,et al.  Adversity Selection and the Habitat Templet , 1983, The American Naturalist.

[57]  P. Coley,et al.  HERBIVORY AND DEFENSIVE CHARACTERISTICS OF TREE SPECIES IN A LOWLAND TROPICAL FOREST , 1983 .

[58]  E. Newman.  Interactions Between Plants , 1983 .

[59]  Hugh G. Gauch,et al.  Multivariate analysis in community ecology , 1984 .

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

[61]  J. P. Grime,et al.  The Plant community as a working mechanism , 1982 .

[62]  David J. Hicks,et al.  The Ecology of Leaf Life Spans , 1982 .

[63]  J. P. Grime,et al.  A COMPARATIVE STUDY OF GERMINATION CHARACTERISTICS IN A LOCAL FLORA , 1981 .

[64]  F. S. Chapin,et al.  The Mineral Nutrition of Wild Plants , 1980 .

[65]  G. Pugh.  Strategies in fungal ecology , 1980 .

[66]  J. P. Grime,et al.  Plant Strategies and Vegetation Processes. , 1980 .

[67]  J. P. Grime,et al.  SEASONAL VARIATION IN THE SEED BANKS OF HERBACEOUS SPECIES IN TEN CONTRASTING HABITATS , 1979 .

[68]  T. Clutton‐Brock,et al.  Comparison and adaptation , 1979, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[69]  M. Hill.  THE DEVELOPMENT OF A FLORA IN EVEN-AGED PLANTATIONS , 1979 .

[70]  W. Westman,et al.  Measuring the Inertia and Resilience of Ecosystems , 1978 .

[71]  J. P. Grime,et al.  Evidence for the Existence of Three Primary Strategies in Plants and Its Relevance to Ecological and Evolutionary Theory , 1977, The American Naturalist.

[72]  T. R. E. Southwood,et al.  HABITAT, THE TEMPLET FOR ECOLOGICAL STRATEGIES? , 1977 .

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

[74]  Roderick Hunt,et al.  Relative growth-rate: its range and adaptive significance in a local flora. , 1975 .

[75]  J. P. Grime.  Vegetation classification by reference to strategies , 1974, Nature.

[76]  E. Newman,et al.  Competition and Diversity in Herbaceous Vegetation , 1973, Nature.

[77]  M. Hill,et al.  Reciprocal Averaging : an eigenvector method of ordination , 1973 .

[78]  G. Ledyard Stebbins,et al.  Chromosomal evolution in higher plants , 1971 .

[79]  John L. Harper,et al.  The Shapes and Sizes of Seeds , 1970 .

[80]  R. Whittaker.  The Population Structure of Vegetation , 1970 .

[81]  E. Odum.  The strategy of ecosystem development. , 1969, Science.

[82]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[83]  J. P. Grime.  Comparative Experiments as a Key to the Ecology of Flowering Plants , 1965 .

[84]  C. Donald,et al.  The interaction of competition for light and for nutrients , 1958 .

[85]  G. Stebbins.  CYTOGENETICS AND EVOLUTION OF THE GRASS FAMILY , 1956 .

[86]  A. R. Clapham.  Autecological Studies and the `Biological Flora of the British Isles' , 1956 .

[87]  E. J. Kraus.  The Reproductive Capacity of Plants , 1943, Ecology.