An important aim of plant ecology is to identify leading dimensions of ecological variation among species and to understand the basis for them. Dimensions that can readily be measured would be especially useful, because they might offer a path towards improved worldwide synthesis across the thousands of field experiments and ecophysiological studies that use just a few species each. Four dimensions are reviewed here. The leaf mass per area-leaf lifespan (LMA-LL) dimension expresses slow turnover of plant parts (at high LMA and long LL), long nutrient residence times, and slow response to favorable growth conditions. The seed mass-seed output (SM-SO) dimension is an important predictor of dispersal to establishment opportunities (seed output) and of establishment success in the face of hazards (seed mass). The LMA-LL and SM-SO dimensions are each underpinned by a single, comprehensible tradeoff, and their consequences are fairly well understood. The leaf size-twig size (LS-TS) spectrum has obvious consequences for the texture of canopies, but the costs and benefits of large versus small leaf and twig size are poorly understood. The height dimension has universally been seen as ecologically important and included in ecological strategy schemes. Nevertheless, height includes several tradeoffs and adaptive elements, which ideally should be treated separately. Each of these four dimensions varies at the scales of climate zones and of site types within landscapes. This variation can be interpreted as adaptation to the physical environment. Each dimension also varies widely among coexisting species. Most likely this within-site variation arises because the ecological opportunities for each species depend strongly on which other species are present, in other words, because the set of species at a site is a stable mixture of strategies.

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Plant Ecological Strategies: Some Leading Dimensions of Variation Between Species

Modern climate change is producing poleward range shifts of numerous taxa, communities and ecosystems worldwide. The response of species to changing environments is likely to be determined largely by population responses at range margins. In contrast to the expanding edge, the low-latitude limit (rear edge) of species ranges remains understudied, and the critical importance of rear edge populations as long-term stores of species' genetic diversity and foci of speciation has been little acknowledged. We review recent findings from the fossil record, phylogeography and ecology to illustrate that rear edge populations are often disproportionately important for the survival and evolution of biota. Their ecological features, dynamics and conservation requirements differ from those of populations in other parts of the range, and some commonly recommended conservation practices might therefore be of little use or even counterproductive for rear edge populations.

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Conserving biodiversity under climate change: the rear edge matters.

What determines the number and size of the seeds produced by a plant? How often should it reproduce them? How often should a plant produce them? Why and how are seeds dispersed, and what are the implications for the diversity and composition of vegetation? These are just some of the questions tackled in this wide-ranging review of the role of seeds in the ecology of plants. The authors bring together information on the ecological aspects of seed biology, starting with a consideration of reproductive strategies in seed plants and progressing through the life cycle, covering seed maturation, dispersal, storage in the soil, dormancy, germination, seedling establishment, and regeneration in the field. The text encompasses a wide range of concepts of general relevance to plant ecology, reflecting the central role that the study of seed ecology has played in elucidating many fundamental aspects of plant community function.

The ecology of seeds

Applications of Hutchinson’s n-dimensional niche concept are often focused on the role of interspecific competition in shaping species distribution patterns. In this paper, I discuss a variety of factors, in addition to competition, that influence the observed relationship between species distribution and the availability of suitable habitat. In particular, I show that Hutchinson’s niche concept can be modified to incorporate the influences of niche width, habitat availability and dispersal, as well as interspecific competition per se. I introduce a simulation model called NICHE that embodies many of Hutchinson’s original niche concepts and use this model to predict patterns of species distribution. The model may help to clarify how dispersal, niche size and competition interact, and under what conditions species might be common in unsuitable habitat or absent from suitable habitat. A brief review of the pertinent literature suggests that species are often absent from suitable habitat and present in unsuitable habitat, in ways predicted by theory. However, most tests of niche theory are hampered by inadequate consideration of what does and does not constitute suitable habitat. More conclusive evidence for these predictions will require rigorous determination of habitat suitability under field conditions. I suggest that to do this, ecologists must measure habitat specific demography and quantify how demographic parameters vary in response to temporal and spatial variation in measurable niche dimensions.

On the relationship between niche and distribution

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.

Long-distance seed dispersal in plant populations.

Availability of seed and microsites, respectively, are two factors that potentially may limit recruitment in plant populations. Microsites are small-scale sites suitable for germination and survival of seedlings. We discuss this dichotomy of recruitment limitation both from a theoretical and empirical point of view. Investigations of recruitment in 14 woodland species showed that 3 species were seed limited, 6 species were limited by a combination of seed and microsite availability, and 5 species were found not to be seed limited, but the limiting factor was not identified. A “combination of seed and microsite limitation” implies that recruitment is promoted by increasing both seed and microsite availability. We suggest that the importance of seed limitation in plant populations has been underestimated, and that the operating limiting factors may be dependent on spatial and temporal scale. We expect that many species, if adequately studied, will turn out to be both seed and microsite limited. Experimental field studies that incorporate a range of seed and microsite “densities” in various spatial and temporal scales are needed to examine the extent to which plant populations are seed and microsite limited.

Seed and microsite limitation of recruitment in plant populations.

Despite a long tradition in plant ecology of studies of patch dynamics, recent developments of models for large scale dynamics in source-sink and metapopulations have largely focused on animals. In contrast to mobile unitary animals, many plants resist extinction. even under conditions where only a part of the life cycle can be maintained. This model of remnant population dynamics adds to the two commonly recognized source-sink and metapopulations dynamics. A review of the literature suggests that all three types of dynamics are common in plants. Regional dynamics are related to life-cycle characteristics determining dispersal ability and longevity of life cycle stages. Short-lived or highly habitat specialized plants with good dispersal tend to build up metapopulations, i.e. systems of local populations and non-occupied but potentially suitable sites, interconnected with dispersal. and in a continuous flux of local colonization and extinction. At the other extreme, long-lived plants with clonal propagation. or plants with extensive seed banks, tend to build up remnant population systems, in which many local populations persist over periods long enough to bridge unfavourable phases of successional development, intervening periods of favourable conditions. Source-sink populations are a special case of metapopulations, in the sense that they comprise both persistent refuge populations and ephemeral populations maintained by dispersal. It is suggested that a concept of local population inertia in remnant population systems, scales to higher level phenomena of vegetation inertia, and to community stabilization (through enhanced recovery after perturbations). Such an inertia may contribute to explain cases of exceptionally high species diversity, and lack of pronounced mass extinctions of plants in the fossil record.

https://www.jstor.org/stable/pdfplus/3546063.pdf

Regional dynamics of plants : a review of evidence for remnant, source-sink and metapopulations

We examined recruitment patterns in semi-natural pastures and their relationships to disturbance and species composition at different spatial scales (0.01 m2 up to 4 m2) of both the vegetation and the seed bank. Possible associations between seedling recruitment and phenology were also studied. The study was performed in four pastures with different management history. Seedling recruitment was generally enhanced by disturbance, with the greatest effect for small-seeded species that germinate in the autumn. The local species-pool contributed to a large extent to seedling recruitment; 83 % of the recruited species were found in the vegetation, whereas 44 % of the recruited species were found in the seed bank. There were a total of 88 species found in the vegetation at different spatial scales; 61 % were recorded at the smallest spatial scale (0.01 m2). The area sampled at this scale comprised 1.25% of the area examined, indicating a small scale structuring of diversity in the pastures. The species number in the vegetation varied between 7 and 14 for the smallest scale (0.01 m2), and between 23 and 42 for the largest scale (4 m2), in the four pastures. The species number in the seed bank was less variable. Like most studies of perennial grasslands, we found no close correspondence between species distribution in the vegetation and in the seed bank. Two of the pastures had a joint management history and exhibited a close similarity of species in the seed bank, despite that one of the pastures was abandoned already in the beginning of this century.

Seedling recruitment in semi-natural pastures: the effects of disturbance, seed size, phenology and seed bank

A hypothesis is suggested for functional roles of remnant plant populations in communities and ecosystems. A remnant population is capable of persistence during extended time periods, despite a negative population growth rate, due to long-lived life stages and life-cycles, including loops that allow population persistence without completion of the whole life cycle. A list of critera is suggested to help identification of remnant plant populations. Several community and ecosystem features may result from the presence of remnant plant populations. Apart from increasing community and ecosystem resilience just by being present, remnant populations may contribute to resilience through enhancing colonization by other plant species, by providing a persistent habitat for assemblages of animals and microorganisms, and by reducing variation in nutrient cycling. It is suggested that the common ability of plants to develop remnant populations is a contributing factor to ecosystem stability. Remnant populations are important for the capacity of ecosystems to cope with the present-day impact caused by human society, and their occurrence should be recognized in surveys of threatened plant species and communities.

Functional roles of remnant plant populations in communities and ecosystems

We examined adhesive seed dispersal, recruitment and site occupancy of 17 plant species characteristic of semi-natural grasslands. The main objectives were to examine different aspects of dispersal: the potential distance of seed transport: the likelihood of recruitment at putative suitable sites along road verges: the spatial (realised) patterns of occurrence along road verges (occupancy): and associations among these aspects and seed size. Adhesive dispersal capacity was investigated experimentally using cattle. Recruitment was investigated by seed sowing experiments. Occupancy was recorded in two regional surveys. Results suggested that adhesive dispersal may be an effective mode of seed transport, even for species which lack special adhesive' structures, such as hooks or hairs. Fifteen of the 17 species successfully recruited at the experimental gaps and adjacent grassy areas. Disturbance promoted seedling recruitment. Seedling emergence was negatively associated with adhesive dispersal capacity. Spatial (occupancy) patterns were not related to recruitment features or dispersal capacity. There is some support for a trade-off between adhesive dispersal capacity and recruitment ability, but no support for a trade-off between adhesive dispersal capacity and ability to withstand competition during recruitment. The results are discussed in the context of species distribution and persistence in fragmented landscapes. We suggest that adhesive dispersal has been underestimated as a mechanism of seed transport, but that adhesive dispersal capacity per se is not the main factor determining species occupancies in fragmented grasslands.

https://www.jstor.org/stable/info/3546442

Ove Eriksson

Papers

Seed and microsite limitation of recruitment in plant populations.

Regional dynamics of plants : a review of evidence for remnant, source-sink and metapopulations

Seedling recruitment in semi-natural pastures: the effects of disturbance, seed size, phenology and seed bank

Functional roles of remnant plant populations in communities and ecosystems

Dispersal, recruitment and site occupancy of grassland plants in fragmented habitats