Rob W. Brooker

Partitioning net interactions among plants along altitudinal gradients to study community responses to climate change

Summary Altitudinal gradients provide a useful space-for-time substitution to examine the capacity for plant competition and facilitation to mediate responses to climate change. Decomposing net interactions into their facilitative and competitive components, and quantifying the performance of plants with and without neighbours along altitudinal gradients, may prove particularly informative in understanding the mechanisms behind plant responses to environmental change. To decouple the inherent responses of species to climate from the responses of plant–plant interactions to climate, we conducted a meta-analysis. Using data from 16 alpine experiments, we tested if changes in net interactions along altitudinal gradients were due to a change in the performance of target species without neighbours (i.e. environmental severity effects only) or with neighbours (neighbour trait mediated effects). There was a global shift from competition to facilitation with increasing altitude driven by both environmental severity and neighbour trait effects. However, this global pattern was strongly influenced by the high number of studies in mesic climates and driven by competition at low altitude in temperate climates (neighbour trait effect), and facilitation at high altitude in arctic and temperate climates (environmental severity effect). In Mediterranean systems, there was no significant effect of competition, and facilitation increased with decreasing altitude. Changes in facilitation with altitude could not unambiguously be attributed to either neighbour trait effects or environmental severity effects, probably because of the opposing stress gradients of cold and aridity in dry environments. Partitioning net interactions along altitudinal gradients led to the prediction that climate change should decrease the importance of facilitation in mesic alpine communities, which might in turn exacerbate the negative effects of climate change in these regions. In xeric climates, the importance of facilitation by drought-tolerant species should increase at low altitudes which should mitigate the negative effect of climate change. However, the importance of facilitation by cold-tolerant species at high altitudes may decrease and exacerbate the effects of climate change.

Don't diss integration: a comment on Ricklefs's disintegrating communities.

Ricklefs’s recent call to investigate ecological processes at large scales helps focus ecologists’ attention on an undoubtedly important topic. However, we believe that some of his accompanying arguments for the primacy of such work and, in particular, for the need to “disintegrate” the local community concept are flawed. We revisit Ricklefs’s main tenets and demonstrate that research on local communities is a vital part of understanding processes and diversity across a range of spatial and temporal scales. The integration of research across spatial scales expands our horizons and understanding of ecology and evolution, and this should not be unnecessarily constrained to one extreme or the other.

Differential effects of reindeer on high Arctic lichens

We studied the effects of Svalbard reindeer on the abundance of lichens in Spitsbergen. A survey was carried out in 14 areas with contrasting reindeer densities. Separate cover estimates for crustose, fructose and foliose lichens were taken in each area, and related to the density of reindeer pellet groups, a measure of reindeer density. Dominant macrolichen families were identified in 10 areas, and a full record of macrolichen species was taken in four additional areas. Varia- tion in reindeer density is partially due to past overhunting, and subsequent incomplete recovery, releasing some areas from reindeer grazing for 100-200 yr. The cover of fruticose lichens was negatively related to reindeer pellet group density, indicating suppression by Svalbard reindeer. This makes their impact comparable to other members of the Rangifer genus around the northern hemisphere. The generally recorded low abundance of lichens in the diet of Svalbard reindeer compared to other Rangifer species, therefore, was interpreted as the depletion of fruticose lichens in Spitsbergen, and a subsequent switch to alternative foods. Of all fruticose lichens, Stereocaulon spp. appeared least sensitive to grazing. Crustose and foliose lichen cover was independent of reindeer pellet group density. The cover of crustose lichens was significantly related to latitude, with greater cover in more northern areas. Foliose lichens were more abundant in places where moss cover was high. We conclude that the impact of Svalbard reindeer on lichens is dependent on growth form, with fruticose lichens suffering from grazing, whereas foliose lichens might indirectly benefit from higher densities of reindeer or, like crustose lichens, be controlled by other factors.

Facilitation and competition in the high Arctic: the importance of the experimental approach

Abstract In the last decade, plant ecologists have focussed more on the occurrence of positive plant–plant interactions than ever before. Especially in severe environments, such as the Arctic, species removal experiments tend to find facilitative rather than competitive effects, casting doubt on the importance of competition under extreme growing conditions. Two approaches to measure plant–plant interactions presented here reveal that competition affects plant growth even in the high Arctic. Luzula confusa and Salix polaris show a reduced growth in mixed stands compared with pure. This competition effect is not detected in a removal experiment, which inevitably also alters site microclimate. Indeed, in the latter experiment, facilitative effects of Luzula on Salix were found. Evidently, both facilitation and competition are acting and important. Causes and implications are discussed using a conceptual model derived from that of Brooker and Callaghan (1998) .

Importance versus intensity of ecological effects: why context matters

In any ecological study, target organisms are usually impacted by multiple environmental drivers. In plant interaction research, recent debate has focussed on the importance of competition; that is, its role in regulating plant success relative to other environmental drivers. Despite being clearly and specifically defined, the apparently simple concept of the importance of competition has been commonly overlooked, and its recognition has helped reconcile long-running debates about the dependence of competition on environmental severity. In this review, we argue that extending this formalised concept of importance to other aspects of ecology would be beneficial. We discuss approaches for measuring importance, and provide examples where explicit acknowledgement of this simple concept might promote understanding and resolve debate.

Trait assembly in plant assemblages and its modulation by productivity and disturbance

Understanding how communities assemble is a key challenge in ecology. Conflicting hypotheses suggest that plant traits within communities should show divergence to reflect strategies to reduce competition or convergence to reflect strong selection for the environmental conditions operating. Further hypotheses suggest that plant traits related to productivity show convergence within communities, but those related to disturbance show divergence. Data on functional diversity (FDvar) of 12 traits from 30 communities ranging from arable fields, mown and grazed grasslands to moorland and woodland were employed to test this using randomisations tests and correlation and regression analysis. No traits showed consistent significant convergence or divergence in functional diversity. When correlated to measures of the environment, the most common pattern was for functional diversity to decline (7 out of 12 traits) and the degree of convergence (7 out of 12 traits) to increase as the levels of productivity (measured as primary productivity, soil nitrogen release and vegetation C:N) and disturbance increased. Convergence or a relationship between functional diversity and the environment was not seen for a number of important traits, such as LDMC and SLA, which are considered as key predictors of ecosystem function. The analysis indicates that taking into account functional diversity within a system may be a necessary part of predicting the relationship between plant traits and ecosystem function, and that this may be of particular importance within less productive and less disturbed systems.

Can soil temperature direct the composition of high arctic plant communities

Low temperatures exert a primary constraint on the growth of high arctic vascular plants. However, investigations into the impact of temperature on high arctic plants rarely separate out the role of air and soil temperatures, and few data exist to indicate whether soil temperatures alone can significantly influence the growth of high arctic vascular plants in a manner that might direct community composition. We examined the response of high arctic plants of three functional types (grasses, sedges/rushes and non-graminoids) to manipulated soil temperature under common air temperature conditions. Target plants, within intact soil cores, were placed in water baths at a range of temperatures between 4.9 and 15.3 oC for one growing season. Grasses responded most rapidly to increased soil temperature, with increased total live plant mass, above-ground live mass and total below-ground live mass, with non-graminoids having the lowest, and sedges/ rushes an intermediate degree of response. The ratio of aboveground live mass to total live mass increased in all growth forms. Grasses, in particular, responded to enhanced soil temperatures by increasing shoot size rather than shoot number. In all growth forms the mass of root tissue beneath the moss layer increased significantly and to a similar extent with increasing soil temperature. These results clearly indicate that different growth forms, although collected from the same plant community, respond differently to changes in soil temperature. As a consequence, factors influencing soil temperature in high arctic ecosystems, such as global climate change or herbivory (which leads to reduced moss depth and increased soil temperatures), may also direct changes in vascular plant community composition.

Plant ecology's guilty little secret: understanding the dynamics of plant competition

Summary Plant competition has been studied for decades. Yet, it is still an elusive concept that means different things to different people, is resistant to direct study and is shrouded in semantic and statistical complexity. We still lack basic information about many competitive mechanisms, processes and outcomes and their relationship to other ecological processes, and about how local interactions between individuals are propagated through communities. We suggest here that two critical issues have been overlooked in previous studies. First, there is a need for direct measurements of the process of competition as opposed to indirect mechanisms of competitive outcomes. Biomass has become the ‘industry standard’ for measuring competition, but we suggest that biomass cannot provide unambiguous insights into plant competition because it is the product of too great a range of factors and processes. Second, the use of a single measure of competition at an arbitrarily assigned end point of an experiment misses much of the complexity of dynamic interactions between competing plants and can lead to erroneous interpretations. Here, we suggest approaches to handle these difficulties, using new techniques or the application of well-known methods in a novel way. We also provide examples of systems or questions where the improved understanding these approaches could bring would be of particular benefit. Ultimately, we suggest the need for a major shift in the way in which we consider and measure plant competition to identify broadly agreed rules for variation in its importance, its role in different communities and habitats, and how and whether it influences or drives patterns of species diversity and abundance.

The value of stress and limitation in an imperfect world: A reply to Körner

Abstract A recent perspective paper offered by Körner essentially argued that ‘limitation’ and ‘stress’ are functionally useless terms for ecology except perhaps within limited contexts such as plant physiology or agriculture. We strongly disagree, and to this end argue that, although stress is not as precise as other concepts in ecology and is probably more difficult to apply to communities than to individuals, if ecologists want to communicate in a meaningful and interesting way about the distribution and abundance of species, we have to use multi-purpose terminology that allows us to scale from reductionistic, strictly quantifiable levels of analysis to more general conceptual levels. Here, we revisit the main arguments presented against these concepts and use three lines of counter-argument to support our conclusion that limitation and stress are necessary concepts for organizing and integrating general ecological inquiry. We discuss (1) the role of interactions between individuals in changing the limitation experienced by a species, (2) the importance of delineating whether stress is being applied to individuals or to the community, and (3) the evolutionary argument that fitness is never perfect since even adapted species are likely limited to some degree by the environment.

Between migration load and evolutionary rescue: dispersal, adaptation and the response of spatially structured populations to environmental change

The evolutionary potential of populations is mainly determined by population size and available genetic variance. However, the adaptability of spatially structured populations may also be affected by dispersal: positively by spreading beneficial mutations across sub-populations, but negatively by moving locally adapted alleles between demes. We develop an individual-based, two-patch, allelic model to investigate the balance between these opposing effects on a populations evolutionary response to rapid climate change. Individual fitness is controlled by two polygenic traits coding for local adaptation either to the environment or to climate. Under conditions of selection that favour the evolution of a generalist phenotype (i.e. weak divergent selection between patches) dispersal has an overall positive effect on the persistence of the population. However, when selection favours locally adapted specialists, the beneficial effects of dispersal outweigh the associated increase in maladaptation for a narrow range of parameter space only (intermediate selection strength and low linkage among loci), where the spread of beneficial climate alleles is not strongly hampered by selection against non-specialists. Given that local selection across heterogeneous and fragmented landscapes is common, the complex effect of dispersal that we describe will play an important role in determining the evolutionary dynamics of many species under rapidly changing climate.