Michelle R. Leishman

The evolutionary ecology of seed size.

Seed mass is a trait that occupies a pivotal position in the ecology of a species. It links the ecology of reproduction and seedling establishment with the ecology of vegetative growth, strategy sectors that are otherwise largely disconnected (Grime et al., 1988; Shipley et al., 1989; Leishman and Westoby, 1992). There is a startling diversity of shapes and sizes of seeds among the plant species of the world. Seeds range from the dust seeds of the Orchidaceae and some saprophytic and parasitic species (around 10 6 g), across ten orders of magnitude to the double coconut Lodoicea seychellarum (104 g) (Harper et al., 1970). Within species, seed size typically spans less than half an order of magnitude (about fourfold: Michaels et al., 1988). Most within-species variation occurs within plant rather than among plants or populations (Michaels et al., 1988; Obeso, 1993; Vaughton and Ramsey, 1998), indicating environmental effects during development rather than genetic differences between mothers. This chapter is concerned with the differences in seed size among species, and the consequences for vegetation dynamics and community composition. During the last 10–15 years, there has been considerable progress in the ecology of seed mass. Unlike many other areas of comparative plant ecology, we have substantial published information from several different scales and research styles. As well as field experiments and demographic studies with a few species at a time, we have simple experiments with larger numbers of species (ten to 50), quantification of seed mass and its correlates in whole-vegetation types (hundreds of species) and tests of consistency across different continents. The wide-scale quantification began as early as Salisbury (1942) and Baker (1972), but has been much added to and consolidated over the past 10 years (e.g. Mazer, 1989, 1990; Leishman and Westoby, 1994a; Leishman et al., 1995; Eriksson and Jakobsson, 1998). The work spanning large numbers of species is complementary to detailed experiments involving only a few species, giving a stronger sense of how widely the results from particular experiments can be generalized. Much of the literature examines how natural selection on seed size might be influenced by various environmental factors. In this context, it is at first glance surprising that seed size varies within communities across a remarkable five to six orders of magnitude (Leishman et al., 1995; Fig. 2.1). Further, there is strong overlap of seed-size distributions between quite different habitats. Within the temperate zone,

Invasion success of exotic plants in natural ecosystems: the role of disturbance, plant attributes and freedom from herbivores

Invasion of natural ecosystems by exotic species is a major threat to biodiversity globally. We assessed two alternative (but not exclusive) hypotheses to explain the success of exotic species in urban bushland on low fertility sandstone-derived soils in Sydney, Australia. These were that success of exotic species is promoted by: (1) plant attributes in particular disturbance types; and (2) freedom from herbivores. We tested these at sites subject to different types of disturbance: nutrient and water enrichment (below stormwater outlets), nutrient enrichment (riparian zones of creeks with an urban catchment) and physical disturbance (tracks), and control sites. At each site we estimated percentage cover of all species and surveyed leaves for damage by herbivores. Species were classified as native, non-invasive exotic or invasive exotic. We found that sites without any disturbance did not support exotic plants. Physically disturbed sites on low fertility soils supported only one exotic species, suggesting that nutrient enrichment is a critical prerequisite for exotic species invasion on low fertility soils. Exotic species cover was highest and native species richness most reduced in areas of highest nutrient enrichment. Both invasive exotic and non-invasive exotic species had significantly lower levels of leaf herbivory than native species, implying that release from pests alone cannot account for the success of invasive species. Specific leaf area of invasive exotic species was consistently higher than specific leaf area of non-invasive exotic and native species, regardless of disturbance type. In physically disturbed sites of higher soil fertility, exotic species were small herbs and grasses of long flowering duration and with small unassisted or wind-dispersed seeds. In sites subject to nutrient-enrichment, exotic species were more likely to be climbers, able to propagate vegetatively, and with seeds dispersed by vertebrates. Thus different plant attributes contribute to exotic species success under different disturbance types. The clearest consistent difference we found between invasive exotic and non-invasive exotic species was in specific leaf area, suggesting that large specific leaf area facilitates invasiveness.

Three keys to the radiation of angiosperms into freezing environments

Early flowering plants are thought to have been woody species restricted to warm habitats. This lineage has since radiated into almost every climate, with manifold growth forms. As angiosperms spread and climate changed, they evolved mechanisms to cope with episodic freezing. To explore the evolution of traits underpinning the ability to persist in freezing conditions, we assembled a large species-level database of growth habit (woody or herbaceous; 49,064 species), as well as leaf phenology (evergreen or deciduous), diameter of hydraulic conduits (that is, xylem vessels and tracheids) and climate occupancies (exposure to freezing). To model the evolution of species’ traits and climate occupancies, we combined these data with an unparalleled dated molecular phylogeny (32,223 species) for land plants. Here we show that woody clades successfully moved into freezing-prone environments by either possessing transport networks of small safe conduits and/or shutting down hydraulic function by dropping leaves during freezing. Herbaceous species largely avoided freezing periods by senescing cheaply constructed aboveground tissue. Growth habit has long been considered labile, but we find that growth habit was less labile than climate occupancy. Additionally, freezing environments were largely filled by lineages that had already become herbs or, when remaining woody, already had small conduits (that is, the trait evolved before the climate occupancy). By contrast, most deciduous woody lineages had an evolutionary shift to seasonally shedding their leaves only after exposure to freezing (that is, the climate occupancy evolved before the trait). For angiosperms to inhabit novel cold environments they had to gain new structural and functional trait solutions; our results suggest that many of these solutions were probably acquired before their foray into the cold.

The Role of Large Seed Size in Shaded Conditions: Experimental Evidence

Large seeds are thought to provide an adaptive advantage to seedlings growing in shaded environments. Four hypotheses that could account for the possible mechanisms underlying this advantage were tested using 23 species ranging in seed size from 0.04 to 22.2 mg. The hypotheses were: seedlings from large seeds are able to survive longer in shade than seedlings from small seeds; seedlings from large seeds divert relatively more resources to shoot development when grown under shaded conditions compared to seedlings from small seeds; seedlings from large seeds always have larger shoots than small-seeded seedlings growing under shaded conditions due to the large initial reserve size of large seeds and the reduction in growth rate differences between small and large seeds under growth-limiting conditions (...)

Correlates of Seed Size Variation: A Comparison Among Five Temperate Floras

1 Five temperate floras were studied to assess to what extent seed size correlations with other plant attributes are consistent across floras. The floras were from three continents: Australia (semiarid woodlands of western New South Wales, arid woodlands of Central Australia, and the Sydney region), North America (Indiana Dunes) and Europe (Sheffield region, UK). The plant attributes used were growth form, plant height, perenniality and dispersal mode. We used general linear models to consider not only the primary correlations between seed size and each other attribute, but also the overlap patterns among correlations to determine if each correlation could be interpreted as a secondary effect via a third variable. 2 Plant height and growth form were consistently correlated with the largest proportion of log seed mass variation (up to 37% in Central Australia). Although there was strong overlap in the amount of log seed mass variation explained by the two attributes (6-22%), each could explain small but significant variation after the other in all floras. The strong association between growth form/plant height and seed size was found not only among unassisted and wind-adapted species, but also among species dispersed by other means. 3 In all floras, dispersal mode was also able to account for significant variation in log seed mass independently of growth form and plant height. The association between plant perenniality and seed size could be explained as a secondary correlation of growth form and plant height with both seed size and perenniality. 4 There were significant differences in log seed mass among the five floras. However, seed size ranged over at least five orders of magnitude in each flora. Differences between floras could account for relatively little (4%) of the variation in seed size between species, compared to the attributes growth form (20%), plant height (20%) and dispersal mode (29%), despite the quite different soils and climates of the five floras. This suggests that seed size is more strongly associated with other plant attributes than with the environmental conditions for establishment. It appears that within any one community, plants have found a diversity of possible solutions to the problems of seedling establishment, resulting in a wide range of log seed mass.

Predicting Dispersal Spectra: A Minimal Set of Hypotheses Based on Plant Attributes

1 The dispersal mode adopted by a plant species is frequently associated with other attributes of the plant and its habitat. In this paper we review these associations and present a set of hypotheses which, when considered together, make a probabilistic prediction of the dispersal mode adopted by a plant species. When applied to a species list, the hypotheses can be used to generate a prediction of its dispersal spectrum, i.e. the percentages of different dispersal modes that have been adopted. 2 The formulation of such a set of hypotheses has several purposes: (i) to summarize existing knowledge about dispersal adaptations and their interrelations with other attributes of plants and their habitats; (ii) to couch that knowledge in such a way that falsifiable predictions can be made; (iii) to arrive at provisional conclusions about which factors are the most important in shaping the evolution of dispersal mode in different plants or different environments. 3 The review of relationships between dispersal mode and other attributes of plants and their habitats lead to the following provisional conclusions; (i) seeds larger than 100 mg tend to be adapted for dispersal by vertebrates while those smaller than 0.1 mg tend to be unassisted; most seeds, however, are between 0.1 and 100 mg, and in this range all of the dispersal modes are feasible; (ii) plant growth form and stature (sometimes in relation to the canopy height of the vegetation) seem to exclude certain dispersal modes; (iii) the availability of specific dispersal vectors seems rarely to be an important determinant of dispersal mode; (iv) attributes of the physical environment also seem rarely to be important, except indirectly through their influence on plant stature and seed size.

Classifying Plants into Groups on the Basis of Associations of Individual Traits--Evidence from Australian Semi-Arid Woodlands

A multivariate analysis of 43 traits of 300 species from semi-arid woodlands in western New South Wales, Australia is described. The 43 traits encompass vegetative, life-history, phenological and seed-biology characters. Five main groups of species were produced which corresponded largely to growth form. These groups were perennial forbs and C3 grasses, subshrubs of the family Chenopodiaceae, perennial C4 grasses, trees and shrubs and annual forbs and grasses. The traits associated with these groups were vegetative, life-history and phenological. Seed-biology traits were poorly associated with the groups (...)

Is plant ecology more siliceous than we realise

Although silicon occurs in all plants, it is an element that is largely overlooked by many plant ecologists and most plant-related research on silicon comes from agronomy, archaeology, palaeontology and biogeochemistry. Plant silicon has many functions, acting biochemically as silicic acid and physically as amorphous silica. It contributes to cell and plant strength and enables plants to respond adaptively to environmental stresses. Consequently, plant silicon can increase plant fitness in many fundamental aspects of ecology, including plant-herbivore interactions, light interception, pathogen resistance and alleviation of abiotic stresses. Here, we provide an ecological perspective to research outcomes from diverse disciplines, showing that silicon is an important element in plant ecology that is worthy of greater attention.

Inside the "Black Box" of River Restoration: Using Catchment History to Identify Disturbance and Response Mechanisms to Set Targets for Process-Based Restoration

Many river restoration projects fail. Inadequate project planning underpins many of the reasons given for failure (such as setting overly ambitious goals; selecting inappropriate sites and techniques; losing stakeholder motivation; and neglecting to monitor, assess, and document projects). Another major problem is the lack of an agreed guiding image to direct the activities aimed at restoring the necessary biophysical and ecological processes within the logistic constraints of on-ground works. Despite a rich literature defining the components of restoration project planning, restoration ecology currently lacks an explicit and logical means of moving from the initial project vision through to on-ground strategies. Yet this process is fundamental because it directly links the ecological goals of the project to the on-ground strategies used to achieve them. We present a planning process that explicitly uses an interdisciplinary mechanistic model of disturbance drivers and system responses to build from the initial project vision to the implementation of on-ground works. A worked example on the Upper Hunter River in southeastern Australia shows how understanding catchment history can reveal disturbance and response mechanisms, thus facilitating process-based restoration.

Volatile isoprenoid emissions from plastid to planet

Approximately 1-2% of net primary production by land plants is re-emitted to the atmosphere as isoprene and monoterpenes. These emissions play major roles in atmospheric chemistry and air pollution-climate interactions. Phenomenological models have been developed to predict their emission rates, but limited understanding of the function and regulation of these emissions has led to large uncertainties in model projections of air quality and greenhouse gas concentrations. We synthesize recent advances in diverse fields, from cell physiology to atmospheric remote sensing, and use this information to propose a simple conceptual model of volatile isoprenoid emission based on regulation of metabolism in the chloroplast. This may provide a robust foundation for scaling up emissions from the cellular to the global scale.