Exploitation Ecosystems Hypothesis Research Articles

The role of evolutionary processes in determining trophic structure

There are two contrasting hypotheses regarding the drivers of biomass‐distribution among trophic levels within ecosystems. Energetic or bottom–up models propose control by supply of energy, either from autotrophy or from the underlying trophic level. Dynamical or top–down models propose control by predators in the overlying trophic level. Although multiple approaches have been used to reconcile these hypotheses, they have rarely considered the evolutionary pressures created by different distributions of biomass. Here, we study the effects of these evolutionary processes using an agent‐based, spatially‐explicit, eco‐evolutionary model. We demonstrate that, when ecosystems are simple and predator–prey relationships between species are fixed and do not evolve, primary productivity limits the number of trophic levels. In this case, the abundance in the top trophic level is always limited by productivity. However, productivity‐limited trophic levels subject those below them to limitation by predation, and predation‐limited trophic levels allow trophic levels below them to grow until limited by productivity. These results match the expectations of the exploitation ecosystems hypothesis (EEH), which predicts the same pattern of alternating top–down and bottom–up control on trophic levels. When species are able to evolve to adaptively adjust their trophic levels, however, selection is liable to lead to collapse of the trophic pyramid through prey switching. Under these conditions, all trophic levels experience bottom–up control. When the evolution of prey switching is restricted, higher trophic levels are evolutionarily stable and the predictions of the EEH are once again met; the stability of long food chains is thus dependent on the difficulty associated with prey switching. These results suggest that, at least under the combinations of model parameters explored in this study, evolutionary limitation is key to maintaining trophic structure.

Ants offset bottom-up control of spiders in Amazonian savanna trees

Community trophic structure is shaped by concurrent bottom-up (resources) and top-down effects (predators), but the extent to which they interact remains uncertain. The Exploitation Ecosystems Hypothesis predicts that predators should offset increases in herbivore abundance with plant productivity, which is supported by data. However, the extent to which interactions within trophic levels (e.g. competition, intraguild predation) have similar effects is less clear. Ants and spiders are abundant in vegetation and consume similar arthropod prey (occasionally each other) and plant exudates, with ants generally showing competitive dominance over spiders. We tested whether ants could shape the trophic structure of tree-dwelling macroarthropod communities by offsetting increases in spider abundance with insect prey. We performed three surveys of the macroarthropod community of 97 trees from two sites in the savanna of Northern Amazonia. Together, ants and spiders represented 74% of the sampled individuals per tree. Insect prey abundance increased with tree crown volume and crown flower cover, consistent with bottom-up limitation. Likewise, both ants and spiders increased with insect prey abundance, with ant abundance also varying with tree species, suggesting reliance on both animal and plant resources. However, as predicted, the positive relationship between spider abundance and insect prey abundance disappeared as ant abundance increased. Our results suggest that agonistic interactions within trophic levels can strongly shape community structure and size by modifying bottom-up effects as much as interactions across trophic levels.

Top-down effects have primacy over bottom-up effects on the population dynamics of a flightless desert bird

In desert ecosystems, some argue that primary productivity controls vertebrate populations while others contend that predators' top-down effects are under-appreciated. While seldom used to explain population dynamics in desert environments, the exploitation ecosystems hypothesis (EEH) unites bottom-up and top-down processes by conceptualizing how the biomass of plants and consumers vary along spatial productivity gradients in the presence and absence of predators. Here, we test predictions from the EEH along temporal primary productivity gradients, by comparing abundances of an apex predator (dingo) and a large omnivorous flightless bird (emu) on either side of Australia's dingo-proof fence over a nine year period. Where dingo populations were not suppressed by humans, their abundance increased with rainfall in the previous 12 months but did not respond to rainfall where their populations were suppressed. Conversely, emu abundance increased in response to antecedent rainfall where dingoes were rare but showed a negligible response to rainfall where dingoes were common. Our results accord with the EEH by suggesting that dingoes' top-down effects can decouple the linkage between emu populations and primary productivity. More generally, our findings show that top-down effects can have primacy over bottom-up effects for both an apex predator and an omnivorous prey species.

Apex Predators Decouple Population Dynamics Between Mesopredators and Their Prey

The mesopredator release hypothesis (MRH) predicts that the removal of apex predators should lead to increased abundance of smaller predators through relaxation of suppressive, top-down effects. However, apex predators’ effects on mesopredators are also likely to be modulated by interactions with human activities and ecosystem productivity. The exploitation ecosystems hypothesis (EEH) predicts that biomass of apex predators will scale with primary productivity but herbivore and mesopredator biomass will remain constant due to top-down control. Here, we take advantage of the manipulation of dingo abundance across Australia’s Dingo Fence to explore the primacy of top-down and bottom-up effects as drivers of feral cat abundance. Using field data collected across the Dingo Fence, we test the predictions generated by MRH and EEH that cat populations should be bottom-up controlled by prey abundance (a proxy for primary productivity) where top-down control exerted by dingoes was weak but not where it was strong. We examined dingo and cat scats to provide mechanistic support for the idea that dingoes control cats through killing and exploitative competition. Overall, cats were more abundant where dingoes were rare. Cat abundance was correlated positively with prey abundance where dingoes were rare but was not correlated with prey abundance where dingoes were common. Cat remains were present in 1% of dingo scats, and dietary overlap between cats and dingoes was high (0.75–0.82). Our study provides evidence that top-down control exerted by apex predators can decouple population dynamics between mesopredators and their prey and thus have primacy over bottom-up effects.

Large‐scale responses of herbivore prey to canid predators and primary productivity

AbstractAimThe primacy of top‐down (consumption) and bottom‐up effects (primary productivity) as forces structuring ecological communities is a controversial topic. The exploitation ecosystems hypothesis (EEH) was invoked to explain biogeographical trends in plant and consumer biomass, and differs from the top‐down/bottom‐up dichotomy by predicting that the relative strength of these processes will vary along gradients of primary productivity. Here we test the prediction of the EEH that herbivore biomass should increase with increasing primary productivity where predators are rare, but show a negligible response to primary productivity where predators are common due to population regulation by predators.LocationBoreal and temperate regions of North America and Eurasia, and deserts of Australia.Time period1970–2016.Major taxa studiedCervids and kangaroos.MethodsWe obtained abundance indices of cervids at 42 locations from the literature and conducted spotlight surveys at 27 locations to derive estimates of kangaroo abundance. For analyses, herbivore abundances were converted to biomass per km2. We tested our prediction using linear mixed effects models.ResultsHerbivore biomass showed divergent responses to increasing primary productivity and the abundance of canid predators (grey wolves, Canis lupus/dingoes, Canis dingo). The slope of the relationship between herbivore biomass and net primary productivity did not differ between Australia and the northern boreal and temperate regions. Herbivore biomass increased in response to primary productivity where canid predators were rare, but showed muted responses to increasing productivity where canid predators were common.Main conclusionsCanid predators have strong suppressive effects on herbivore biomass that scale with primary productivity. Our study shows that the EEH has wide application to canid‐predator–herbivore dynamics and may be relevant to the management of herbivores because it can provide an indication of how herbivore biomass and densities may vary in relation to ecosystem productivity and the presence and absence of canid predators.

Strength of a Trophic Cascade Between an Apex Predator, Mammalian Herbivore and Grasses in a Desert Ecosystem Does Not Vary with Temporal Fluctuations in Primary Productivity

There has long been debate regarding the primacy of bottom-up and top-down effects as factors shaping ecosystems. The exploitation ecosystems hypothesis (EEH) predicts that predators indirectly benefit plants because their top-down effects limit herbivores’ consumption of plants, and that the strength of trophic cascade increases with increasing primary productivity. However, in arid environments, pulses of primary productivity produced by irregular rainfall events could decouple herbivore–plant and predator–prey dynamics if high conversion efficiency from seed biomass to consumers allows the rapid build-up of consumer populations. Here, we test predictions of the EEH in an arid environment. We measured activity/abundances of dingoes, red kangaroos and grasses, and diet of dingoes, in landscapes where dingoes were culled or not culled over 3 years. Dingo activity was correlated with rainfall, and their tracks were less frequent at culled sites. Kangaroo abundance was greater at sites where dingoes were culled and increased with rainfall in the previous 6 months. Grass cover was greater at sites where dingoes were not culled and increased with rainfall in the previous 3 months. During a period of average rainfall, dingoes primarily consumed rodents and increased their consumption of kangaroos during a period of drier conditions. Our results are consistent with the hypothesis that suppression of an apex predator triggers a trophic cascade, but are at odds with the EEH’s prediction that the magnitude of trophic cascades should increase with primary productivity. Our study demonstrates that temporal fluctuations in primary productivity can have effects on biomasses of plants and consumers which are in many ways analogous to those observed along spatial gradients of primary productivity.

Changes in the Spatial Configuration and Strength of Trophic Control Across a Productivity Gradient During a Massive Rodent Outbreak

Understanding the determinants of spatial and temporal differences in the relative strength of consumer–resource interactions is an important endeavour in ecology. Here, we explore the necessary conditions for temporal shifts in the relative strength of rodent–plant interactions in an area characterised by profound spatial differences in trophic control, with predator–prey interactions prevailing in productive habitats and rodent–plant interactions dominating unproductive habitats of the forest–tundra ecotone. We report data obtained during the exceptionally massive rodent outbreak of 2010–2012 in northernmost Fennoscandia, including an experimental manipulation of herbivore access to vegetation plots across a large-scale productivity gradient, multiple observational measures of plant–rodent interactions linked to rodent abundance data and a large-scale survey of breeding avian predators and mammalian predator activity. Unexpectedly, rodent grazing impacts documented during the rodent outbreak were uniformly strong across the landscape, regardless of habitat productivity. The runaway response in rodent populations was facilitated by a high population growth rate in the early phase of the outbreak due to the extended absence of predators in productive habitats, concomitant with an exceptionally long-lasting lemming outbreak in unproductive habitats. Our results showed that spatio-temporal variation in trophic control also occurs in ecosystems structured according to the exploitation ecosystems hypothesis and emphasises the importance of long-term studies to capture nonlinear and stochastic features that shape ecosystem functioning. In this context, the temporary release from top–down regulation in productive habitats caused strong grazing impacts that may be crucial for the resilience of tundra ecosystems under the threat of climate change-driven shrub encroachment.

Spatial variation in vegetation damage relative to primary productivity, small rodent abundance and predation

The relative importance of top‐down and bottom‐up mechanisms in shaping community structure is still a highly controversial topic in ecology. Predatory top‐down control of herbivores is thought to relax herbivore impact on the vegetation through trophic cascades. However, trophic cascades may be weak in terrestrial systems as the complexity of food webs makes responses harder to predict. Alternatively, top‐down control prevails, but the top‐level (predator or herbivore) changes according to productivity levels. Here we show how spatial variation in the occurrence of herbivores (lemmings and voles) and their predators (mustelids and foxes) relates with grazing damage in landscapes with different net primary productivity, generating two and three trophic level communities, during the 2007 rodent peak in northern Norway. Lemmings were most abundant on the unproductive high‐altitude tundra, where few predators were present and the impact of herbivores on vegetation was strong. Voles were most common on a productive, south facing slope, where numerous predators were present, and the impacts of herbivores on vegetation were weak. The impact of herbivores on the vegetation was strong only when predators were not present, and this cannot be explained by between‐habitat differences in the abundance of plant functional groups. We thus conclude that predators influence the plant community via a trophic cascade in a spatial pattern that support the exploitation ecosystems hypothesis. The responses to grazing also differed between plant functional groups, with implications for short and long‐term consequences for plant communities.

Exploitation ecosystems and trophic cascades in non‐equilibrium systems: pasture – red kangaroo – dingo interactions in arid Australia

The exploitation ecosystems hypothesis (EEH) proposes that 1) plant biomass reflects the primary productivity of an ecosystem modified by the regulating effect of herbivory, and 2) herbivore abundance reflects the productivity of plants modified by the regulating effect of predation. Primary productivity thus determines the number of trophic levels in an ecosystem and the extent to which bottom–up and top–down regulation influence the biomass ratios of adjacent and non‐adjacent trophic levels (i.e. trophic cascading). We constructed an interactive model of plant (pasture), herbivore (red kangaroo Macropus rufus) and predator (dingo Canis lupus dingo), a system in which trophic cascades have been suggested to occur, and used it to test the effects of increasing stochastic variation in primary productivity and dingo culling on predictions of the EEH. The model contained four feedback loops: the predator–herbivore and herbivore–plant feedback loops, and the predator and plant density‐dependent feedback loops. The equilibrium conditions along the primary productivity gradient reproduced the three zones of trophic dynamics predicted by the EEH, plus an additional zone at productivities above which the maximum density of a predator is achieved due to social regulation: that zone is characterized by increasing herbivore density and decreasing plant biomass. Culling dingoes produced trophic cascades that were strongly attenuated at primary productivities below which the maximum density of dingoes was attained. Results were robust to uncertainty in kangaroo off‐take by dingoes and to the efficacy of dingo culling, but prey switching by dingoes from red kangaroos to reptiles would weaken trophic cascades. We conclude that social regulation of carnivores has important implications for expression of the EEH and trophic cascades, and that attenuation of trophic cascades increases with increasing stochasticity in primary productivity. Our model also provides a framework for understanding the conditions in which dingo‐mediated trophic cascades might be expected to occur, and generates testable predictions about the effects of higher dingo densities (e.g. by stopping culling or reintroduction to former range) on kangaroo and pasture dynamics.

Patterns in the abundance of kangaroo populations in arid Australia are consistent with the exploitation ecosystems hypothesis

The exploitation ecosystems hypothesis (EEH) makes predictions about trophic interactions along gradients of primary productivity. The EEH has been shown to apply to a wide range of terrestrial environments but its applicability to arid environments has received little attention. One reason for this is that arid environments may not satisfy the assumptions of the EEH because dearth of water may limit biological activity in both temporal and spatial contexts. The EEH predicts that herbivore biomass should increase linearly with primary productivity in the absence of predators; but when predators are present herbivore biomass will remain relatively constant due to top down regulation. We tested this prediction in an arid environment using rainfall as a proxy of primary productivity and an index of the abundance of the dominant herbivores (kangaroos Macropus spp.). We compared an index of kangaroo abundance at 18 areas situated along a gradient of mean annual rainfall in areas where a top predator (the dingo Canis lupus dingo) was rare and common. We also explored the relationship between the density of artificial water points (AWPs) and kangaroo abundance to investigate if the resource subsidy provided by AWPs allows kangaroos to persist in high numbers. Consistent with the EEH, kangaroo abundance showed a weak relationship with mean annual rainfall in the presence of dingoes but increased with increasing annual rainfall in the absence of dingoes. The density of AWPs was a poor predictor of kangaroo abundance. Our analysis of macro‐ecological patterns suggests that kangaroo populations are primarily top down regulated in the presence of dingoes, but are bottom up regulated in the absence of dingoes. Our findings provide evidence that top down regulation can prevail over bottom up regulation of herbivore populations in arid ecosystems and highlights the usefulness of the EEH as a predictor of macro‐ecological patterns of species abundance.

Large predators limit herbivore densities in northern forest ecosystems

There is a lack of scientific consensus about how top-down and bottom-up forces interact to structure terrestrial ecosystems. This is especially true for systems with large carnivore and herbivore species where the effects of predation versus food limitation on herbivores are controversial. Uncertainty exists whether top-down forces driven by large carnivores are common, and if so, how their influences vary with predator guild composition and primary productivity. Based on data and information in 42 published studies from over a 50-year time span, we analyzed the composition of large predator guilds and prey densities across a productivity gradient in boreal and temperate forests of North America and Eurasia. We found that predation by large mammalian carnivores, especially sympatric gray wolves (Canis lupus) and bears (Ursus spp.), apparently limits densities of large mammalian herbivores. We found that cervid densities, measured in deer equivalents, averaged nearly six times greater in areas without wolves compared to areas with wolves. In areas with wolves, herbivore density increased only slightly with increasing productivity. These predator effects are consistent with the exploitation ecosystems hypothesis and appear to occur across a broad range of net primary productivities. Results are also consistent with theory on trophic cascades, suggesting widespread and top-down forcing by large carnivores on large herbivores in forest biomes across the northern hemisphere. These findings have important conservation implications involving not only the management of large carnivores but also that of large herbivores and plant communities.

Cascading predator control interacts with productivity to determine the trophic level of biomass accumulation in a benthic food web

AbstractLarge‐scale exploitation of higher trophic levels by humans, together with global‐scale nutrient enrichment, highlights the need to explore interactions between predator loss and resource availability. The hypothesis of exploitation ecosystems suggests that top–down and bottom–up control alternate between trophic levels, resulting in a positive relationship between primary production and the abundance of every second trophic level. Specifically, in food webs with three effective trophic levels, primary producers and predators should increase with primary production, while in food webs with two trophic levels, only herbivores should increase. We provided short‐term experimental support for these model predictions in a natural benthic community with three effective trophic levels, where the number of algal recruits, but not the biomass of gastropod grazers, increased with algal production. In contrast, when the food web was reduced to two trophic levels by removing larger predators, the number of algal recruits was unchanged while gastropod grazer biomass increased with algal production. Predator removal only affected the consumer‐controlled early life‐stages of algae, indicating that both the number of trophic levels and the life‐stage development of the producer trophic level determine the propagation of trophic cascades in benthic systems. Our results support the hypothesis that predators interact with resource availability to determine food‐web structure.

Top predators, mesopredators and their prey: interference ecosystems along bioclimatic productivity gradients

1. The Mesopredator Release Hypothesis (MRH) suggests that top predator suppression of mesopredators is a key ecosystem function with cascading impacts on herbivore prey, but it remains to be shown that this top-down cascade impacts the large-scale structure of ecosystems. 2. The Exploitation Ecosystems Hypothesis (EEH) predicts that regional ecosystem structures are determined by top-down exploitation and bottom-up productivity. In contrast to MRH, EEH assumes that interference among predators has a negligible impact on the structure of ecosystems with three trophic levels. 3. We use the recolonization of a top predator in a three-level boreal ecosystem as a natural experiment to test if large-scale biomass distributions and population trends support MRH. Inspired by EEH, we also test if top-down interference and bottom-up productivity impact regional ecosystem structures. 4. We use data from the Finnish Wildlife Triangle Scheme which has monitored top predator (lynx, Lynx lynx), mesopredator (red fox, Vulpes vulpes) and prey (mountain hare, Lepus timidus) abundance for 17 years in a 200 000 km(2) study area which covers a distinct productivity gradient. 5. Fox biomass was lower than expected from productivity where lynx biomass was high, whilst hare biomass was lower than expected from productivity where fox biomass was high. Hence, where interference controlled fox abundance, lynx had an indirect positive impact on hare abundance as predicted by MRH. The rates of change indicated that lynx expansion gradually suppressed fox biomass. 6. Lynx status caused shifts between ecosystem structures. In the 'interference ecosystem', lynx and hare biomass increased with productivity whilst fox biomass did not. In the 'mesopredator release ecosystem', fox biomass increased with productivity but hare biomass did not. Thus, biomass controlled top-down did not respond to changes in productivity. This fulfils a critical prediction of EEH. 7. We conclude that the cascade involving top predators, mesopredators and their prey can determine large-scale biomass distribution patterns and regional ecosystem structures. Hence, interference within trophic levels has to be taken into account to understand how terrestrial ecosystem structures are shaped.

Community Ecology: How Green Is the Arctic Tundra?

The exploitation ecosystems hypothesis suggests that food chain length increases along gradients of increasing primary productivity. Recent results provide compelling new evidence for this from an arctic-alpine ecosystem.

Spatial Patterns and Dynamic Responses of Arctic Food Webs Corroborate the Exploitation Ecosystems Hypothesis (EEH)

According to the exploitation ecosystems hypothesis (EEH), productive terrestrial ecosystems are characterized by community-level trophic cascades, whereas unproductive ecosystems harbor food-limited grazers, which regulate community-level plant biomass. We tested this hypothesis along arctic-alpine productivity gradients at the Joatka field base, Finnmark, Norway. In unproductive habitats, mammalian predators were absent and plant biomass was constant, whereas herbivore biomass varied, reflecting the productivity of the habitat. In productive habitats, predatory mammals were persistently present and plant biomass varied in space, but herbivore biomass did not. Plant biomass of productive tundra scrublands declined by 40% when vegetation blocks were transferred to predation-free islands. Corresponding transfer to herbivore-free islands triggered an increase in plant biomass. Fertilization of an unproductive tundra heath resulted in a fourfold increase in rodent density and a corresponding increase in winter grazing activity, whereas the total aboveground plant biomass remained unchanged. These results corroborate the predictions of the EEH, implying that the endotherm community and the vegetation of the North European tundra behaves dynamically as if each trophic level consisted of a single population, in spite of local co-occurrence of >20 plant species representing different major taxonomic groups, growth forms, and defensive strategies.

Regional effects on competition–productivity relationship: a set of field experiments in two distant regions

We studied the effect of productivity on competition intensity and the relationship between competition intensity and community species richness, using a removal experiment with the perennial plant Solidago virgaurea. The experiment was conducted in 16 different communities from two geographically distant areas (western Estonia and northern Norway). The results were compared with the results of previous experiments with Anthoxanthum odoratum from the same areas. Removal of neighbors had a positive effect on the biomass of both Solidago and Anthoxanthum, and this response was stronger in communities with higher productivity. Thus, the corrected index of relative competition intensity, CRCI, increased with increasing community productivity. Species richness was negatively correlated with CRCI in Estonia but not in Norway and not in the case of the pooled material. The results suggest that competitive exclusion operates at least in these communities which species pool is large.Our results indicate that the relationship between competition intensity and productivity is non‐linear. In our data, competition prevails in communities where living plant biomass exceeds 200 g m−2, whereas in less productive communities, competition remains undetected and direct plant–plant relationships might at times be even mutualistic. Moreover, we found that the relationship between competition intensity and productivity is strongly dependent on regional differences and is intimately connected to a concordant variation in the intensity of grazing. The least productive communities both in Estonia and in Norway are characterized by intensive grazing, which reduces importance of competition. Hence, the contrasting results corroborates the predictions of the hypothesis of exploitation ecosystems, predicting that trophic dynamics account for the relationship between competition intensity and primary productivity.

Long‐term dynamics of voles and lemmings at the timberline and above the willow limit as a test of hypotheses on trophic interactions

We have monitored population fluctuations of microtine rodents since 1977 in two habitat complexes in Finnmark, northernmost Norway ‐ a low arctic plains landscape, with patches of willow scrubland embedded in lichen‐dwarf birch tundra, and in adjacent highlands, occupied by scrub‐free heaths, snow‐beds and bogs. In the plains landscape, voles were cyclic, with a period of five years, and with wave‐like density fluctuations. This pattern is consistent with time trajectory of prey in a predator‐prey limit cycle. Autoregression analysis implies that the prey pattern is cleanest in the most productive plains habitats, while dynamics in the prevailing heath and bog habitats are governed by two significant lags, implying that even vole‐plant interactions count. In the highlands, lemmings had two outbreaks, characterized by J‐shaped growth curves, and separated by long periods of low density. The fluctuation pattern of lemmings in highlands was consistent with the predicted time trajectory of a predator. The implications of time trajectories are corroborated by direct evidence on microtine impacts upon the vegetation and on spatial patterns in predator activity. Even the strong dispersal tendency of lemmings during population peaks is consistent with the conjecture that they are adapted to play the role of a predator in a sustained predator‐prey cycle. As a whole, the pattern supports T. Oksanen's modification of the hypothesis of exploitation ecosystems, where both local productivity and the structure of the landscape are taken in account.

Comparative Effects on Plants of Caribou/Reindeer, Moose and White-tailed Deer Herbivory

We reviewed the literature reporting negative or positive effects on vegetation of herbivory by caribou/reindeer, moose, and white-tailed deer in light of the hypothesis of exploitation ecosystems (EEH), which predicts that most of the negative impacts will occur in areas where wolves were extirpated. We were able to list 197 plant taxa negatively affected by the three cervid species, as opposed to 24 that benefited from their herbivory. The plant taxa negatively affected by caribou/reindeer (19), moose (37), and white-tailed deer (141) comprised 5%, 9%, and 11% of vascular plants present in their respective ranges. Each cervid affected mostly species eaten during the growing season: lichens and woody species for caribou/reindeer, woody species and aquatics for moose, and herbs and woody species for white-tailed deer. White-tailed deer were the only deer reported to feed on threatened or endangered plants. Studies related to damage caused by caribou/reindeer were scarce and often concerned lichens. Most reports for moose and white-tailed deer came from areas where wolves were absent or rare. Among the three cervids, white- tailed deer might damage the most vegetation because of its smaller size and preference for herbs.

Effects of grazing on above‐ground biomass on a mountain snowbed, NW Finland

The effects of grazing by the Norwegian lemming (Lemmus lemmus) and reindeer (Rangifer tarandus) on biomass and vegetation structure in a mountain snowbed habitat were studied using several grazer‐exclosures kept for 5 or 15 yr. During the 5‐yr exclusion experiment, the total biomass of mosses in particular, the main winter‐food of lemmings, increased within fenced areas as compared to open control areas. In the 15‐yr exclosures, Polytrichaceae mosses still dominated, but graminoids (Anthoxanthum odoratum and Carex bigelowii) had also increased. In these longer‐term exclosures, dead plant material, consisting of Polytrichaceae moss and graminoid necromass and other plant litter, had increased. By contrast, Kiaeria moss and lichens were absent or had low biomass after 15 yr of grazer exclosure, probably as a result of competitive exclusion. The results suggest that the plant biomass in such low productivity mountain snowbeds can be reduced to a low level by grazing. Grazing also determines the dominance relationships in the plant community and prevents accumulation of plant litter. The observed effects of grazing are inconsistent with hypotheses suggesting that herbivory is of little importance in low productivity environments, but rather support the exploitation ecosystems hypothesis predicting strong grazing impact in such low productivity environments.

The Logic and Realism of the Hypothesis of Exploitation Ecosystems.

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.