Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind's most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.

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Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana, has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (40.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (o0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.

Hurlbert divides experimental ecologist into ‘those who do not see any need for dispersion (of replicated treatments and controls), and those who do recognize its importance and take whatever measures are necessary to achieve a good dose of it’. Experimental ecologists could also be divided into those who do not see any problems with sacrificing spatial and temporal scales in order to obtain replication, and those who understand that appropriate scale must always have priority over replication. If an experiment is conducted in a spatial or temporal scale, where the predictions of contesting hypotheses are convergent or ambiguous, no amount of technical impeccability can make the work instructive. Conversely, replication can always be obtained afterwards, by conducting more experiments with basically similar design in different areas and by using meta‐analysis. This approach even reduces the sampling bias obtained if resources are allocated to a small number of well‐replicated experiments. For a strict advocate of the hypothetico‐deductive method, replication is unnecessary even as a matter of principle, unless the predicted response is so weak that random background noise is a plausible excuse for a discrepancy between predictions and results. By definition, a prediction is an ‘all‐statement’, referring to all systems within a well‐defined category. What applies to all must apply to any. Hence, choosing two systems and assigning them randomly to a treatment and a control is normally an adequate design for a deductive experiment. The strength of such experiments depends on the firmness of the predictions and their a priori probability of corroboration. Replication is but one of many ways of reducing this probability. Whether the experiment is replicated or not, inferential statistics should always be used, to enable the reader to judge how well the apparent patterns in samples reflect real patterns in statistical populations. The concept ‘pseudoreplication’ amounts to entirely unwarranted stigmatization of a reasonable way to test predictions referring to large‐scale systems.

Hypotheses on trophic dynamics in terrestrial ecosystems fall into two major categories: those in which plants are assumed to be invulnerable to their consumers and those in which the build-up of plant biomass is assumed to require top-down control of folivores. The hypothesis of exploitation ecosystems (EEH) belongs to the latter category and focuses particularly on the consequences of the high energetic costs of maintenance of endotherms. Carnivorous endotherms require relatively high prey densities in order to break even. Moreover, they are dependent on folivorous prey during the limiting season, at least at high latitudes. The endotherm branch of the grazing web is thus predicted to collapse from three-link trophic dynamics (carnivores → folivores → plants → inorganic resources) to two-link dynamics (folivores → plants → inorganic resources) along gradients of decreasing primary productivity. Consequently, the vegetation of cold and unproductive areas is predicted to be under intense winter grazing pressure, which prevents the accumulation of aboveground plant biomass and excludes erect woody plants. In the most extreme habitats (e.g., polar deserts and their high alpine counterparts), even folivorous endotherms are predicted to be absent, and the scanty vegetation is predicted to be structured by preemptive competition. Within temperature-determined productivity gradients, EEH is corroborated by biomass patterns, by patterns in the structure and dynamics of carnivore, folivore, and plant communities, and by experimental results. The general idea of top-down trophic dynamics is supported for other autotroph-based systems, too, but the relevance and sufficiency of the energy constraint in explaining patterns in trophic dynamics appears to be variable. Moreover, critical empirical evidence for or against the capacity of folivorous insects to regulate plant biomass has not yet been obtained. Another open question is the ability of boreal and temperate browsers, evolved in productive environments with intense predation pressure and abundance of forage, to prevent the regeneration of the least palatable tree species. There are, thus, many open questions waiting to be answered and many exciting experiments waiting to be conducted.

In this study, we investigated the effect of reindeer grazing on tundra heath vegetation in northern Norway. Fences, erected 30 yr ago, allowed us to compare winter grazed, lightly summer grazed and heavily summer grazed vegetation at four different sites. At two sites, graminoids dominated the heavily grazed zone completely, while ericoid dwarf shrubs had almost disappeared. In the other two areas, the increase of graminoids was almost significant. At one of the sites where graminoids dominated the heavily grazed area, we also measured plant biomass, primary production and nitrogen cycling. In this site, heavy grazing increased primary production and rate of nitrogen cycling, while moderate grazing decreased primary production. These results were inconsistent with the view that the highest productivity is found at intermediate grazing pressure. These results rather support the hypothesis that intensive grazing can promote a transition of moss‐rich heath tundra into productive, graminoid‐dominated steppe‐like tundra vegetation. Moreover the results suggests that intermittent intensive reindeer grazing can enhance productivity of summer ranges.

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Olofsson, J., Stark, S. and Oksanen, L. 2004. Reindeer influence on ecosystem processes in the tundra. Á/ Oikos 105: 386 Á/396.Reindeer have been recorded to increase nutrient cycling rate and primary production in studies from fences almost 40 years old that separate areas with different grazing regimes in northern Fennoscandia. To further understand the mechanism behind the effects of herbivores on primary production, we measured the size of the major C and N pools, soil temperature, litter decomposition rate and N mineralization rate in lightly, moderately and heavily grazed areas along two of these fences.Plant N found in new biomass, indicative of plant N assimilation, was significantly higher in moderately and heavily grazed areas than in lightly grazed areas, which corresponded to a decreased amount of N in old plant parts. The amount of N found in plant litter or organic soil layer did not differ between the grazing treatments. Together with soil N concentrations and litter decomposition rates, soil temperatures were significantly higher in moderately and heavily grazed areas.We conclude that the changes in soil temperature are important for the litter decomposition rate and thus on the nutrient availability for plant uptake. However, the changes in plant community composition appear to be more important for the altered N pools and thus the enhanced primary production. The results provide some support for the keystone herbivore hypothesis, which states that intensive grazing can promote a transition from moss-rich tundra heath to productive grasslands. Grazing altered N fluxes and pools, but the total N pools were similar in all grazing treatments. Our study thus indicates that grazing can increase the primary production through enhancing the soil nutrient cycling rate, even in a long term perspective on an ecological timescale.