The measurement of plant dispersal is vital for understanding plant distribution and abundance at different scales. However, dispersal is difficult to measure and there is a lack of guidance for researchers new to the subject. In this paper we provide advice on methods for measuring dispersal in the field and approaches to experimental design. First, we encourage clear exposition of the aims of the dispersal study and the ultimate use to which the data will be put (e.g. local dynamics, invasion processes, etc). We outline the types of dispersal exhibited by plants and emphasise that many species are dispersed by multiple processes, which are not necessarily related to putative adaptations. Few studies properly address the full range of processes by which a species is dispersed. We review methods for measuring plant dispersal, summarising the type of dispersal measured and problems with each method. We then outline the major questions about effort to be considered in sampling protocols and present an optimisation algorithm for designing dispersal studies given a suite of options, and biological and resource constraints. We propose and demonstrate a simulation modelling approach to comparing the data quality obtained by alternative experimental designs. Integrating simulation models with pilot studies offers a rapid route to improved estimation methods. We then discuss functions commonly fit to dispersal data and recommend caution as none is a priori the best description of the dispersal process. Finally, we call for a better description and understanding of dispersal kernels by: a more rigorous approach to designing dispersal measurement; better targeting of dispersal studies to particular questions; and achieving a deeper understanding of the mechanisms underlying dispersal, so that we can move from descriptions of pattern to a grasp of process.
Biotic interactions are often ignored in assessments of climate change impacts. However, climate-related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co-occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad-scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population-level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.
Summary 1.Invasive species usually exhibit different spatial population dynamics in their native and invaded range. This is often attributed to demographic differences, but may be due to differences in dispersal as well. 2. Regardless of how these dispersal and demographic differences from the native range arose, studying how they contributed to increases in population spread rates will increase our understanding of what has made these species invasive. Here we investigate which vital rates and dispersal parameters of the invasive thistle Carduus nutans drive the increases in spread rate in different invaded ranges compared to that in the native range in Eurasia. 3. We construct and analyse spatial integrodifference models that combine structured, local population models with mechanistic (WALD) models of seed dispersal by wind across a homogeneous landscape. Published and new demographic and dispersal data for single populations from the native (France) and invaded (Australia, New Zealand, Kansas and Pennsylvania) ranges were used for the parameterization. 4. We developed a variance decomposition method ( c *-LTRE) to analyse the contributions of the changes in the vital rates and dispersal parameters to the increases in the invasion wave speed ( c *) estimates for the different invaded ranges compared to that for the native range. 5. The c *-LTRE analysis showed that the net contribution of the dispersal parameters to c * increases varied among the populations from 51% (Australia), to 79% (Kansas), to 80% (New Zealand) and to 85% (Pennsylvanian experiment). Escape from natural enemies that reduce seed set by floral herbivory was important in all invaded ranges. Large positive contributions were also made by increases in rapid growth of seedlings and small rosettes, increases in flowering probabilities and potential seed production, as well as by increased plant height and lower falling velocities of the plumed seeds. 6. Synthesis . By incorporating a mechanistic dispersal model with a structured population model, and by linking this joint model to field data from several continents, we demonstrate the relative importance of dispersal and demography to invasion success. This approach can be used to analyse which aspects of an invader's life history have changed most importantly from the native range.Key-words: fixed-factor life table response experiment (LTRE), integrodifference equations, matrix population models, musk or nodding thistle, native vs. invaded range, plant height, seed terminal velocity, vital rates, Wald analytical long-distance model, wind speed
Understanding and predicting population spread rates is an important problem in basic and applied ecology. In this article, we link estimates of invasion wave speeds to species traits and environmental conditions. We present detailed field studies of wind dispersal and compare nonparametric (i.e., data-based) and mechanistic (fluid dynamics model-based) dispersal kernel and spread rate estimates for two important invasive weeds, Carduus nutans and Carduus acanthoides. A high-effort trapping design revealed highly leptokurtic dispersal distributions, with seeds caught up to 96 m from the source, far further than mean dispersal distances (approx. 2 m). Nonparametric wave speed estimates are highly sensitive to sampling effort. Mechanistic estimates are insensitive to sampling because they are obtained from independent data and more useful because they are based on the dispersal mechanism. Over a wide range of realistic conditions, mechanistic spread rate estimates were most sensitive to high winds and low seed settling velocities. The combination of integrodifference equations and mechanistic dispersal models is a powerful tool for estimating invasion spread rates and for linking these estimates to characteristics of the species and the environment.
Dispersal is a critical process in ecology. It is an important biological driver of, for example, invasions, metapopulation dynamics, spatial pattern formation and pathogen movement. Much is known about the effect of environmental variability, including turbulence, on dispersal of diaspores. Here, we document experimentally the strong but under-explored influence of turbulence on the initiation of dispersal. Flower heads of two thistle species (Carduus nutans and Carduus acanthoides) with ripe seeds were exposed to series of laminar and turbulent air flows of increasing velocity in a wind tunnel. Seed release increased with wind speeds for both laminar and turbulent flows for both species. However, far more seeds were released, at significantly lower wind speeds, during turbulent flows. These results strongly suggest a need for more quantitative studies of abscission in the field, as well as dispersal models that incorporate variability in the diaspore release phase.
In recent decades we have seen rapid and co-occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi-annual landscape-wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co-occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed-species and age-heterogeneous forests with good site-matching tend to be less susceptible to large-scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.
Dispersal is a key process in biological studies of spatial dynamics, but the initiation of dispersal has often been neglected, despite strong indications that differential timing of dispersal can significantly affect dispersal distances. To investigate which plant and environmental factors determine the release of plumed seeds by the invasive thistles Carduus acanthoides and Carduus nutans, we exposed 192 flower heads of each species to increasing wind speeds in a full-factorial wind tunnel experiment with four air flow turbulence, three flower head wetness and two flower head temperature levels. The number of seed releases was highest under dry and turbulent conditions and from heads that had already lost a considerable number of seeds, but was not affected by flower head size, head angle or temperature. Inspection of the trials on video showed that higher wind speeds were needed to meet the seed release threshold in laminar flows and for C. acanthoides heads that had been wet for a longer time. Species differences were minimal, although seed release was more sensitive to lower levels of turbulence in the larger-headed and more open C. nutans heads. Knowledge of seed release biases towards weather conditions favourable for long-distance dispersal improves our understanding of the spread of invaders and allows managers to increase the efficiency of their containment strategies by applying them at crucial times.
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