Soil Chemical Analysis

There is growing recognition that classifying terrestrial plant species on the basis of their function (into 'functional types') rather than their higher taxonomic identity, is a promising way forward for tackling important ecological questions at the scale of ecosystems, landscapes or biomes. These questions include those on vegetation responses to and vegetation effects on, environmental changes (e.g. changes in climate, atmospheric chemistry, land use or other disturbances). There is also growing consensus about a shortlist of plant traits that should underlie such functional plant classifications, because they have strong predictive power of important ecosystem responses to environmental change and/or they themselves have strong impacts on ecosystem processes. The most favoured traits are those that are also relatively easy and inexpensive to measure for large numbers of plant species. Large international research efforts, promoted by the IGBP–GCTE Programme, are underway to screen predominant plant species in various ecosystems and biomes worldwide for such traits. This paper provides an international methodological protocol aimed at standardising this research effort, based on consensus among a broad group of scientists in this field. It features a practical handbook with step-by-step recipes, with relatively brief information about the ecological context, for 28 functional traits recognised as critical for tackling large-scale ecological questions.

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A handbook of protocols for standardised and easy measurement of plant functional traits worldwide

Biological invasions into wholly new regions are a consequence of a far reaching but underappreciated component of global environmental change, the human-caused breakdown of biogeographic barriers to species dispersal . Human activity moves species from place to place both accidentally and deliberately-and it does so at rates that are without precedent in the last tens of millions of years. As a result , taxa that evolved in isolation from each other are being forced into contact in an instant of evolutionary time. This human-caused breakdown of barriers to dispersal sets in motion changes that may seem less important than the changing composition of the atmosphere , climate change , or tropical deforestation-but they are significant for several reasons. First , to date , biological invasions have caused more species extinctions than have resulted from human-caused climatic change or the changing composition of the atmosphere . Only land use change probably has caused more extinction, and (as we later discuss) land use change interacts strongly with biological invasions. Second, the effects of human-caused biological invasions are long-term: changes in climate, the atmosphere, and land use may be reversible in hundreds to thousands of years, but the breakdown of biogeographic barriers has resulted in self-maintaining and evolving

Biological invasions by exotic grasses, the grass/fire cycle, and global change

Plant functional traits are the features (morphological, physiological, phenological) that represent ecological strategies and determine how plants respond to environmental factors, affect other trophic levels and influence ecosystem properties. Variation in plant functional traits, and trait syndromes, has proven useful for tackling many important ecological questions at a range of scales, giving rise to a demand for standardised ways to measure ecologically meaningful plant traits. This line of research has been among the most fruitful avenues for understanding ecological and evolutionary patterns and processes. It also has the potential both to build a predictive set of local, regional and global relationships between plants and environment and to quantify a wide range of natural and human-driven processes, including changes in biodiversity, the impacts of species invasions, alterations in biogeochemical processes and vegetation–atmosphere interactions. The importance of these topics dictates the urgent need for more and better data, and increases the value of standardised protocols for quantifying trait variation of different species, in particular for traits with power to predict plant- and ecosystem-level processes, and for traits that can be measured relatively easily. Updated and expanded from the widely used previous version, this handbook retains the focus on clearly presented, widely applicable, step-by-step recipes, with a minimum of text on theory, and not only includes updated methods for the traits previously covered, but also introduces many new protocols for further traits. This new handbook has a better balance between whole-plant traits, leaf traits, root and stem traits and regenerative traits, and puts particular emphasis on traits important for predicting species’ effects on key ecosystem properties. We hope this new handbook becomes a standard companion in local and global efforts to learn about the responses and impacts of different plant species with respect to environmental changes in the present, past and future.

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New handbook for standardised measurement of plant functional traits worldwide

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

We explored the relationship between leaf specific mass (LSM) and its two components, leaf density and thickness. These were assessed on the leaves of (a) the moderately sclerophyllous tree Arbutus menziesii distributed along a natural nutrient/moisture gradient in California, (b) eight sclerophyllous shrub species on four substrates in south-western Australia, and (c) seedlings of two morphologically contrasting Hakea species grown under varying soil nutrient, moisture and light regimes in a glasshouse experiment. Leaf area, mass, LSM, density and thickness varied greatly between leaves on the same plant, different species, and with different nutrient, moisture and light regimes. In some cases, variations in LSM were due to changes in leaf density in particular or thickness or both, while in others, density and thickness varied without a net effect on LSM. At lower nutrient or moisture availabilities or at higher light irradiances, leaves tended to be smaller, with higher LSM, density and thickness. Under increased stress, the thickness (diameter) of needle leaves decreased despite an increase in LSM. We concluded that, while LSM is a useful measure of sclerophylly, its separation into leaf density and thickness may be more appropriate as they often vary independently and appear to be more responsive to environmental gradients than LSM.

Leaf specific mass confounds leaf density and thickness

All individuals of all known populations of Banksia goodii were assessed for seed production. Small populations produced no or only a few seeds per unit canopy area. Effects of population size on seed production per unit area and seed production per plant were present over the whole range of population sizes, indicating that even in large populations seed production may still not be at its maximum. Resource differences could not explain this disproportionate decrease in seed production with decline in population size, because there were no differences in soil properties and understorey or overstorey cover between the small and large populations. Although plants in small and large populations were similar in size, seed production per plant was much lower in small populations. This was not because plants in small populations produced fewer cones but because the fraction of these cones that was fertile was much lower. Five of the nine smallest populations (<200 m2) produced no fertile cones over the last 10 years. The number of seeds per fertile cone did not depend on population size. The results are discussed in relation to pollination biology.

Population fragmentation may reduce fertility to zero in Banksia goodii — a demonstration of the Allee effect

We explore the effect of post—fire microsites on seed and seedling distribution and hence their potential role in community restoration. A summer wildfire and control burn in a sclerophyll shrubland in mediterranean Australia produced mosaics of physically and chemically contrasting microsites of litter and sand. Most seeds (>75%) of all species released from the burnt canopies fell, or were redispersed by wind, into the litter patches after both fires. Data on microsite characteristics and wind exposure (fire intensity), height of fruits, time of release, and seed properties were required to interpret relative distribution between the litter and sand patches. Seeds remained equally viable (up to 100%) over summer—autumn in the litter and sand and had equally high rates and levels (up to 100%) of subsequent winter germination. However, seedlings were 2—3 times less likely to survive in the litter and survivors were 35% smaller than those in the sand by the end of the first summer. Banksia hookeriana was particularly vulnerable to microsite properties, whereas the needle—leaved Hakea polyathema showed only minor responses. Pre—summer thinning of seedlings in the litter increased survival of the remainder by 2 times and size of the survivors by 31%. The fire—sensitive, small—seeded B. hookeriana had 17 times more seeds in the backburn litter than the resprouting, larger—seeded B. attenuata, which more than compensated for its 3 times greater seedling mortality levels over the dry summer. Recruitment of species prone to density—dependent mortality in the litter was enhanced by the retention of some seeds in the sand where competition for water was minimal, as indicated by the 2.2 times greater stomal conductance of their seedlings in early summer.

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Post-fire litter microsites: Safe for seeds, unsafe for seedlings

(1) Production, nitrogen and phosphorus return, and decomposition of leaf litter of the invasive alien, Acacia saligna, was compared with that of the indigenous sclerophyllous shrub, Leucospermum parile, in sand-plain lowland fynbos with acid soils low in P during the early stages of alien invasion. The same was done for A. cyclops and Pterocelastrus tricuspidatus in strandveld with alkaline soils high in P. (2) The Acacia spp. had twice the leaf N concentrations of the indigenous species, whereas P concentrations were highest in the strandveld species. (3) The Acacia spp. tended to produce more litter, with three times the N content of that of the indigenous species. No significant differences in P return were found, between the acacias and indigenous species in either vegetation. (4) Decomposition turnover times were longer in the fynbos species than those of the strandveld. Nitrogen was immobilized in the leaf litter of the indigenous species, while the N contents of the acacias varied little. Phosphorus was immobilized in the fynbos species compared with a release of about 50% from P. tricuspidatus after 2 years. (5) Soil N concentrations and litter-layer N contents were elevated under acacia canopies. (6) The N status of the fynbos and strandveld ecosystems is elevated by the invasion of alien acacias. The results for P cycling are equivocal and P availability does not appear to limit plant growth in the strandveld.

Ed T.F. Witkowski

Papers

Leaf specific mass confounds leaf density and thickness

Population fragmentation may reduce fertility to zero in Banksia goodii — a demonstration of the Allee effect

Post-fire litter microsites: Safe for seeds, unsafe for seedlings

Effects of invasive alien acacias on nutrient cycling in the coastal lowlands of the cape fynbos

Regeneration by coppicing (resprouting) of miombo (African savanna) trees in relation to land use