Exclosure Experiment Research Articles

Airborne laser scanning reveals increased growth and complexity of boreal forest canopies across a network of ungulate exclosures in Norway

AbstractLarge herbivores are often classed as ecosystem engineers, and when they become scarce or overabundant, this can alter ecosystem states and influence climate forcing potentials. This realization has spurred a call to integrate large herbivores in earth system models. However, we lack a good understanding of their net effects on climate forcing, including carbon and energy exchange. A possible solution to this lies in harmonizing data across the myriad of large herbivore exclosure experiments around the world. This is challenging due to differences in experimental designs and field protocols. We used airborne laser scanning (ALS) to describe the effect of herbivore removal across 43 young boreal forest stands in Norway and found that exclusion caused the canopy height to increase from 1.7 ± 0.2 to 2.5 ± 0.2 m (means ± se), and also causing a marked increase in vertical complexity and above‐ground biomass. We then go on to discuss some of the issues with using ALS; we propose ALS as an approach for studying the effects of multiple large herbivore exclosure experiments simultaneously, and producing area‐based estimates on canopy structure and forest biomass in a cheap, efficient, standardized and reproducible way. We suggest that this is a vital next step towards generating biome‐wide predictions for the effects of large herbivores on forest ecosystem structure which can both inform both local management goals and earth system models.

Flood‐stimulated herbivory drives range retraction of a plant ecosystem

Abstract Climate change is generating extreme climate events, affecting ecosystem integrity and function directly through increases in abiotic stress and disturbance and indirectly through changes in the strength of biotic interactions. As consumers play an essential role in ecosystem functioning and have been shown to be highly sensitive to climate conditions, improved understanding of their role under changing environmental conditions is necessary to accurately anticipate climate change impacts on ecosystem integrity. We evaluated if prolonged periods of extreme rain, a climatic event increasing in severity in many places around the world, and coincident increases in coastal flooding duration intensify consumer control of foundational salt marsh grass structure and quantify the consequences of flooding–consumer interactions on salt marsh range extent. To achieve this, we analysed: historic trends in crab grazing; crab numbers and activity in and out of rainy years on the low marsh edge; vegetation retreat from the low marsh edge at a plot scale in a manipulative exclosure experiment; vegetation retreat at a landscape‐scale from drone image analyses; and the vertical erosion in the lowest edge of an Argentinean salt marsh. During flooded periods, crabs congregated in the low marsh, resulting in localized overgrazing of salt marsh grass and the rapid horizontal retreat of the marsh edge (98.5 cm on average). Salt marsh edge retreat resulted in a loss of ~4.5% of the total marsh area at the landscape scale. Inside crab exclusion plots, although grass cover declined slightly during the study period, the marsh edge did not retreat. Synthesis. This study provides experimental evidence that an extreme climate event can destabilize a local consumer–prey interaction, indirectly triggering the range contraction of a critical coastal habitat. This work contributes to a growing body of research demonstrating that consumers can be unleashed, rather than suppressed, by extreme climatic events. Moreover, in cases where consumer fronts form during such events, the result can be not only local (along habitat edges) but also landscape‐scale extinction of foundation species and the habitats they biogenically create. Together, this supports the general idea that models of future climate scenarios integrate the indirect effects on ecosystem‐regulating food web interactions.

Introduction of giant tortoises as a replacement “ecosystem engineer” to facilitate restoration of Santa Fe Island, Galapagos

Re‐establishment of ecosystem engineers via introduction of replacement species is increasingly being proposed to facilitate ecosystem restoration of island ecosystems around the world, yet attendant challenges and risks remain largely unknown. We evaluate the first phase (2015–2020) of a program to restore a population of giant tortoises, ecosystem engineers of terrestrial ecosystems of the Galapagos Islands, to Santa Fe Island during which a total of 551 juvenile and 31 subadult tortoises of the non‐native Española Island species (Chelonoidis hoodensis) were translocated. After 5 years, most (85%) of tortoises released remain alive, exhibiting robust vital rates comparable to those of conspecifics in their native range, having colonized 10% of the island via gradual dispersal from the point‐of‐release. Populations of an arboreal cactus (Opuntia echios)—a keystone species for many organisms on the island and intended beneficiary of tortoise restoration—and pallid land iguanas (Conolophus pallidus)—an endemic, herbivorous reptile considered potentially at risk from the introduction of tortoises, both increased coincident with the arrival of tortoises. Herbivore exclosure experiments revealed that the island's plant community manifested a response to the arrival of tortoises; however, significant engineering of the island's vegetation will require many decades. These results highlight the first stage of what appears to be a successful conservation intervention that can serve as a “proof of concept” for introduction of ecological replacements to other islands in Galapagos, and potentially for other island ecosystems around the world.

Deer exclusion unveils abiotic filtering in forest understorey plant assemblages.

The role of deer (family Cervidae) in ecosystem functioning has traditionally been neglected by forest ecologists due to the animal's scarcity in most parts of the northern hemisphere. However, the dramatic rebound in deer populations throughout the 20th century has brought deer browsing to the forefront of forest ecological questioning. Today there is ample evidence that deer affect tree regeneration, understorey plant and animal diversity, and even litter decomposition. However, the mechanisms underlying the effects of deer on forest ecosystems remain unclear. Among others, the relative role of abiotic factors versus biotic interactions (e.g. herbivory) in shaping plant assemblages remains largely unknown. We used a large-scale experiment with exclosures distributed along abiotic gradients to understand the role of black-tailed deer (Odocoileus hemionus sitchensis) on the forest understorey on the Haida Gwaii archipelago (western Canada), a unique context where most of the key ecological effects of deer presence have already been intensively studied. Our results demonstrate that 20 years of deer exclusion resulted in a clear increase in vascular plant richness, diversity and cover, and caused a decline in bryophyte cover. Exclusion also unveiled abiotic (i.e. soil water availability and fertility) filtering of plant assemblages that would otherwise have been masked by the impact of abundant deer populations. However, deer exclusion did not lead to an increase in beta diversity, probably because some remnant species had a competitive advantage to regrow after decades of over browsing. We demonstrated that long-term herbivory by deer can be a dominant factor structuring understorey plant communities that overwhelms abiotic factors. However, while exclosures prove useful to assess the overall effects of large herbivores, the results from our studies at broader scales on the Haida Gwaii archipelago suggest that exclosure experiments should be used cautiously when inferring the mechanisms at work.

Springtime grazing by Arctic-breeding geese reduces first- and second-harvest yields on sub-Arctic agricultural grasslands

Large population increases of Arctic-breeding waterfowls over recent decades have intensified the conflict with agricultural interests in both Eurasia and North America. In the spring-staging region Vesterålen in sub-Arctic Norway, sheep, dairy and meat farmers have reported reduced agricultural grassland yields due to pink-footed geese Anser brachyrhynchus and barnacle geese Branta leucopsis that rest and forage in the region for 3–4 weeks in spring on their way to their breeding grounds on Svalbard. Here, we report from an experimental exclosure design where goose access to plots at three grassland fields in Vesterålen was prevented. The experiment was conducted over 3 years between 2012 and 2014. Goose abundance varied greatly between fields and years as a function of variable spring weather and forage quantity, facilitating evaluation of longer-term impacts under contrasting grazing intensities. First and second harvest yields across fields and years were 20% and 19% higher in exclosures than in plots open for grazing, while total yields (sum of first and second harvests) were on average 27% higher. Within-year effects on harvest yields varied substantially, primarily due to highly contrasting sward development during the spring-staging periods. Cool weather (2012) led to slow sward development and little or no effects on harvest yields, warmer weather (2013) resulted in generally large effects, while variable weather (2014) led to treatment effects varying across fields, with one field experiencing 61% higher yields in exclosures while there were no significant impacts on first-harvest yields at the two other fields. Goose grazing did not increase dry weight-based proportions of weeds. Overall, the farmers' reports on yield-loss due to goose grazing were confirmed, although impacts varied substantially between years. A novel finding is that second-harvest yields were also reduced. For the most affected farmers, it is unlikely that the current subsidy scheme is sufficient to cover all the their losses.

Termite mound cover and abundance respond to herbivore-mediated biotic changes in a Kenyan savanna.

Both termites and large mammalian herbivores (LMH) are savanna ecosystem engineers that have profound impacts on ecosystem structure and function. Both of these savanna engineers modulate many common and shared dietary resources such as woody and herbaceous plant biomass, yet few studies have addressed how they impact one another. In particular, it is unclear how herbivores may influence the abundance of long‐lived termite mounds via changes in termite dietary resources such as woody and herbaceous biomass. While it has long been assumed that abundance and areal cover of termite mounds in the landscape remain relatively stable, most data are observational, and few experiments have tested how termite mound patterns may respond to biotic factors such as changes in large herbivore communities. Here, we use a broad tree density gradient and two landscape‐scale experimental manipulations—the first a multi‐guild large herbivore exclosure experiment (20 years after establishment) and the second a tree removal experiment (8 years after establishment)—to demonstrate that patterns in Odontotermes termite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15–1,000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that are significantly affected by different LMH treatments, are both positive predictors of termite mound abundance. Experimental reductions of tree densities are associated with lower abundances of termite mounds. These results reveal a richly interacting web of relationships among multiple savanna ecosystem engineers and suggest that termite mound abundance and areal cover are intimately tied to herbivore‐driven resource availability.

Summer and winter browsing affect conifer growth differently: An experimental study in a multi-species ungulate community

Ungulate browsing has been studied for several decades in the northern hemisphere. However, studies have mainly focused on just one or two ungulate species, while rarely contrasting the relative effects of summer and winter browsing. This limits our understanding of the dynamics and effects of browsing in landscapes where ungulate species diversity is increasing. We conducted a seasonal exclosure experiment on former clear-cuts in a multi-species ungulate system in Sweden, to investigate the relative impacts of summer and winter browsing on the conifers Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) at the tree species level. We tested for differences in individual conifer growth and demographic responses between Summer browsing, Winter browsing, No browsing, and Control treatment plots over a 4.5 year experimental period. We defined the demographic response as the distribution of conifers among different height classes. Individual growth rates and demographic responses of both conifer species were similar in the plots with No browsing (year-round exclosures) as in the Control plots with year-round browsing. Plots subject to Summer and Winter browsing differed in terms of their demographic response relative to plots with No browsing and Control plots; more stems reached taller height classes in the Summer and Winter browsing plots with slight differences between the conifer species. We discuss the different responses of Scots pine and Norway spruce considering their differences in palatability and their ability to tolerate plant-plant competition in a multi-species ungulate system, in light of potential associational effects. If fencing is intended to be used as a management practice to mitigate the impacts of deer browsing on conifer growth, our results suggest that a total exclusion of deer does not necessarily enhance conifer growth during the first years of regeneration.

Naïve plant communities and individuals may initially suffer in the face of reintroduced megafauna: An experimental exploration of rewilding from an African savanna rangeland.

Excluding large native mammals is an inverse test of rewilding. A 25-year exclosure experiment in an African savanna rangeland offers insight into the potentials and pitfalls of the rewilding endeavor as they relate to the native plant community. A broad theme that has emerged from this research is that entire plant communities, as well as individual plants, adjust to the absence of herbivores in ways that can ill-prepare them for the return of these herbivores. Three lines of evidence suggest that these "naïve" individuals, populations, and communities are likely to initially suffer from herbivore rewilding. First, plots protected from wild herbivores for the past 25 years have developed rich diversity of woody plants that are absent from unfenced plots, and presumably would disappear upon rewilding. Second, individuals of the dominant tree in this system, Acacia drepanolobium, greatly reduce their defences in the absence of browsers, and the sudden arrival of these herbivores (in this case, through a temporary fence break), resulted in far greater elephant damage than for their conspecifics in adjacent plots that had been continually exposed to herbivory. Third, the removal of herbivores favoured the most palatable grass species, and a large number of rarer species, which presumably would be at risk from herbivore re-introduction. In summary, the native communities that we observe in defaunated landscapes may be very different from their pre-defaunation states, and we are likely to see some large changes to these plant communities upon rewilding with large herbivores, including potential reductions in plant diversity. Lastly, our experimental manipulation of cattle represents an additional test of the role of livestock in rewilding. Cattle are in many ways ecologically dissimilar to wildlife (in particular their greater densities), but in other ways they may serve as ecological surrogates for wildlife, which could buffer ecosystems from some of the ecological costs of rewilding. More fundamentally, African savannah ecosystems represent a challenge to traditional Western definitions of "wilderness" as ecosystems free of human impacts. We support the suggestion that as we "rewild" our biodiversity landscapes, we redefine "wildness" in the 21st Century to be inclusive of (low impact, and sometimes traditional) human practices that are compatible with the sustainability of native (and re-introduced) biodiversity.

Wild and domestic savanna herbivores increase smaller vertebrate diversity, but less than additively

Abstract Cattle and other livestock graze more than a quarter of the world's terrestrial area and are widely regarded to be drivers of global biodiversity declines. Studies often compare the effects of livestock presence/absence but, to our knowledge, no studies have tested for interactive effects between large wild herbivores and livestock at varying stocking rates on small‐bodied wild vertebrates. We investigated the effects of cattle stocking rates (none/moderate/high) on the diversity of wildlife 0.05–1,000 kg using camera traps at a long‐term exclosure experiment within a semi‐arid savanna ecosystem in central Kenya. In addition, by selectively excluding wild ‘mesoherbivores’ (50–1,000 kg) and ‘megaherbivores’ (>1,000 kg; elephant and giraffe), we tested whether the presence of these two wild herbivore guilds (collectively, ‘larger wild herbivores’) mediates the effect of cattle stocking rate on habitat use and diversity of ‘smaller wildlife’ (mammals ranging between 10 and 70 cm shoulder height and birds). Our results show that cattle enhance alpha diversity of smaller wildlife (with or without larger wild herbivore presence) and of all wildlife 0.05–1,000 kg (with or without megaherbivore presence), by altering vegetation structure. However, for smaller wildlife, this effect is less pronounced in the presence of larger wild herbivores, which also shorten grass. In the absence of cattle, mesoherbivore‐accessible sites showed higher alpha diversity of smaller wildlife than sites excluding mesoherbivores. Smaller wildlife habitat use was increased by high cattle stocking rates and wild mesoherbivores more in the presence of the other. Synthesis and applications. Our findings imply that grazing, whether by livestock or wildlife, can enhance local savanna wildlife diversity. The biodiversity benefits of localised increases in herbivory are likely to be due to shortened grass and associated visibility improvements (for predator avoidance/foraging). This suggests that land managers can increase local biodiversity by shortening grass, with wild or domestic herbivores (or both), at least in patches within a taller grass matrix.

Frugivore zoogeochemistry in tropical forest ecosystems

Frugivore zoogeochemistry in tropical forest ecosystems

Assessment of Pest Control Services by Vertebrates in Nigerian Subsistence Maize Farms

Global conversion of patches of natural vegetation into agricultural land is reducing the ecosystem services provided by natural patches dwelling species to farmers. For sub-Saharan African subsistence farmers, such a reduction in pest control services by birds may be a significant disadvantage. Here we explored to what extent birds provide pest control services to the staple crop maize (Zea mays) on small subsistence farms on the Mambilla Plateau of Taraba State, Nigeria. We used exclosure experiments (maize crops with and without birds) to model how birds influenced crop yield. We found that excluding birds from maize significantly reduces crop yield, although the lack of a direct correlation between bird abundance and crop yield suggests that other taxa, such as bats, may also be important pest predators. Our results suggest that in this subsistence farming landscape, natural pest control of maize from vertebrates does occur, but further research is needed to understand the specific control agents and the role of patches of natural vegetation as habitat for them.

Are Yellow Sticky Cards and Light Traps Effective on Tea Green Leafhoppers and Their Predators in Chinese Tea Plantations?

Simple SummaryLeafhoppers are serious insect pests in agriculture across the world. Both nymphs and adults suck the sap of plant shoots and leaves with their piercing–sucking mouthparts causing damage called “hopperburn”. The tea green leafhopper, Empoasca onukii, is one of most damaging pests in tea plantations in Asia. In China, yellow sticky cards and light traps are increasingly used to control leafhoppers in tea plantations, especially the tea green leafhopper. Visually, several leafhoppers appear to be captured and killed, however, the real control efficiency and the damage to natural enemies remains unclear. In our study, a 16-week open field experiment with daily weather monitoring was designed to test the responses of tea green leafhopper, parasitoids and spiders to yellow sticky cards and light traps (cover with sticky cards) that used different light colours. An exclosure experiment was also designed to further test the influence of the three light systems (without sticky card) on the same groups of species. The results suggested that light, especially green and white, can be useful as a way to control leafhopper populations without affecting parasitoids and spiders too much.In Chinese tea plantations, yellow sticky cards and light traps are increasingly used to control insect pests, especially the tea green leafhopper Empoasca onukii. In this study, a 16-week open-field experiment with daily weather monitoring was designed to test the responses of tea green leafhopper, parasitoids and spiders to yellow sticky cards and three light traps with different wavelengths (covered with sticky cards). An exclosure experiment was also designed to further test the influence of the three light systems (without sticky card) on the same species. The results showed that all three light emitting diode (LED) light traps (white, green and yellow) and yellow sticky cards attracted many more E. onukii male adults than females during the course of the open field experiment, with less than 25% of trapped adults being females. Parasitoids and spiders were also attracted by these systems. Weather variables, especially rainfall, influenced the trapping efficiency. In the exclosure experiment, the population of leafhoppers in the yellow sticky card treatment did not decline significantly, but the number of spiders significantly decreased. The green and white light treatments without sticky cards showed a significant control of E. onukii and no obvious harm to spiders. These results suggest that yellow sticky cards and light traps have limited capacity to control tea green leafhoppers. However, light, especially green light, may be a promising population control measure for tea green leafhoppers, not as killing agents in the traps, but rather as a behavioral control system.

Lizard and frog removal increases spider abundance but does not cascade to increase herbivory

AbstractInsectivorous vertebrates, especially on islands, can exert top‐down control on herbivorous prey, which can transfer through a food chain to reduce herbivory. However, in many systems insectivorous vertebrates feed on more than one trophic level, especially consuming arthropod predators, and this intraguild predation can diminish trophic cascades. Our goal was to determine, using an exclosure experiment, the relative importance of anole lizards and coqui frogs in controlling spider and arthropod abundances as well as herbivory rates in the understory of the Luquillo Experimental Forest, Puerto Rico. We found that exclosures removing both anoles and coquis doubled spider abundance compared to exclosures with anoles and coquis at natural densities. The effect of coquis on spiders was greater and occurred more quickly than that of anoles, potentially because of the higher natural densities of coquis and removal of both vertebrates produced no interactive effects. We found support for the idea that anoles, but not coquis, reduce foliar arthropod abundances on one of the two studied plant species. However, there was also evidence that anole removal decreased herbivory, the opposite of what we would expect if there was a trophic cascade. Potential explanations include that anoles reduced predatory arthropods on foliage more than they reduced herbivorous arthropods. Results highlight that the food web in tabonuco forest is not simple and that there are complex and dynamic relationships among vertebrate insectivores, predatory arthropods, and herbivorous arthropods that do not consistently result in a trophic cascade.Abstract in Spanish is available with online material

Analyzing long-term impacts of ungulate herbivory on forest-recruitment dynamics at community and species level contrasting tree densities versus maximum heights

Herbivores are constitutive elements of most terrestrial ecosystems. Understanding effects of herbivory on ecosystem dynamics is thus a major, albeit challenging task in community ecology. Effects of mammals on plant communities are typically explored by comparing plant densities or diversity in exclosure experiments. This might over-estimate long-term herbivore effects at community levels as early life stage mortality is driven by a multitude of factors. Addressing these challenges, we established a set of 100 pairs of ungulate exclosures and unfenced control plots (25 m2) in mixed montane forests in the Alps in 1989 covering a forest area of 90 km2. Investigations ran until 2013. Analogous to the gap-maker–gap-filler approach, dynamically recording the height of the largest trees per tree species in paired plots with and without exclosures might allow for assessing herbivore impacts on those individuals with a high probability of attaining reproductive stages. We thus tested if recording maximum heights of regenerating trees would better reflect effects of ungulate herbivory on long-term dynamics of tree regeneration than recording of stem density, and if species dominance patterns would shift over time. For quantifying the effects of ungulate herbivory simultaneously at community and species level we used principle response curves (PRC). PRCs yielded traceable results both at community and species level. Trajectories of maximum heights yielded significant results contrary to trajectories of total stem density. Response patterns of tree species were not uniform over time: e.g., both Norway spruce and European larch switched in their response to fencing. Fencing explained about 3% of the variance of maximum tree heights after nine years but increased to about 10% after 24 years thus confirming the importance of long-term surveys. Maximum height dynamics of tree species, addressed in our study, can thus reflect local dominance of tree species via asymmetric plant competition. Such effects, both within and among forest patches, can accrue over time shaping forest structure and composition.

The role of browsers in maintaining the openness of savanna grazing lawns

Abstract In savannas, ruminant herbivores can have divergent impacts on tree recruitment and subsequent woody cover. Whereas heavy grazing by cattle results in woody thickening, intensive grazing by wildlife instead tends to be associated with lower woody cover. To disentangle why woody cover is low in areas heavily grazed by wildlife, we investigated (a) whether grazing lawns attract indigenous mammalian browsers, and if a preference for short‐grass habitat decreases with browser body mass as predator susceptibility decreases; and (b) whether browser attraction to grazing lawns translates into the suppression of woody plants, including seedlings and saplings, thus maintaining the openness of heavily grazed short‐grass areas. In Kruger National Park, South Africa, we contrasted browser abundance (using dung counts) on grazing lawns and on low‐herbivory sites characterised by tall grass. Additionally, a herbivore exclosure experiment was set up to investigate the combined impact of browser removal and grass height habitat type on seedling survival and sapling growth of a dominant woody plant species. Finally, in Hluhluwe‐iMfolozi Park (HiP), we examined the long‐term (10 years) impact of browser removal on the growth rates of a range of woody species, monitored across ten sites along a gradient of herbivory ranging from grazing lawn to tall grass. Steenbok and impala selected short‐ over tall grass as preferred browsing sites, while elephant preferred tall grass. Browser abundance on short grass decreased with browser body mass, indicating that predator avoidance might be a key factor driving mesoherbivores to utilise grazing lawns. Seedling survival was lowest on grazing lawns when browsers were present, with mortality occurring in two out of every three seedlings. Similarly, sapling growth was lowest on grazing lawns, although browser removal had no significant effect. Evidence for increased browser impact on grazing lawns was clearest from our long‐term herbivore exclosure experiment in HiP, which demonstrated that browsers strongly modify the growth rates of woody plants in short‐grass habitats. Synthesis. These results provide support for the hypothesis that browsers, particularly browsing mesoherbivores and mixed feeders, are attracted to short‐grass habitats, and that they help maintain grazing lawn openness by suppressing seedling survival and woody plant growth where grass is kept short by grazers.

Cool as a moose: How can browsing counteract climate warming effects across boreal forest ecosystems?

Herbivory has potential to modify vegetation responses to climatic changes. However, climate and herbivory also affect each other, and rarely work in isolation from other ecological factors, such as plant-plant competition. Thus, it is challenging to predict the extent to which herbivory can counteract, amplify, or interact with climate impacts on ecosystems. Here, we investigate how moose modify climatic responses of boreal trees by using experimental exclosures on two continents and modeling complex causal pathways including several climatic factors, multiple tree species, competition, tree height, time, food availability, and herbivore presence, density, and browsing intensity. We show that moose can counteract, that is, "cool down" positive temperature responses of trees, but that this effect varies between species depending on moose foraging preferences. Growth of preferred deciduous trees was strongly affected by moose, whereas growth of less preferred conifers was mostly driven by climate and tree height. In addition, moose changed temperature responses of rowan in Norway and balsam fir in Canada, by making fir more responsive to temperature but decreasing the strength of the temperature response of rowan. Snow protected trees from browsing, and therefore moose "cooling power" might increase should a warming climate result in decreased snow cover. Furthermore, we found evidence of indirect effects of moose via plant-plant competition: By constraining growth of competing trees, moose can contribute positively to the growth of other trees. Our study shows that in boreal forests, herbivory cooling power is highly context dependent, and in order to understand its potential to prevent changes induced by warming climate, species differences, snow, competition, and climate effects on browsing need to be considered.

Context-dependent effects of a reintroduced ungulate on soil properties are driven by soil texture, moisture, and herbivore activity.

Although there is considerable evidence that large mammalian herbivores influence ecosystem‐level processes, studies have reported such widely varying results that generalizations have remained elusive. Here, we use an 18‐year‐old exclosure experiment—stratified across a landscape heterogeneous with respect to soil texture, moisture and herbivore activity—to understand the variable effects of tule elk (Cervus canadensis nannodes), a native reintroduced herbivore, on soil properties along the coast of northern California. Elk significantly increased soil bulk density and created a compacted layer at shallow soil depth, while decreasing infiltration rate and pH. The effects of elk on bulk density, penetration resistance, and pH varied with soil type, being least pronounced in coarse, sandy loams, and greatest in loose sand. The effects of elk on nutrient availability varied along gradients of soil texture and moisture. In coarser soils, elk decreased ammonium availability, but increased it in finer soils. Elk also decreased soil moisture content, in part through their positive effect on bulk density, and this effect was most pronounced in coarser soils. Through decreasing soil moisture content, elk also decreased nitrate availability in coarser soils. At greater levels of elk activity (as measured by dung deposition), the elk effect on bulk density was amplified, and this had a corresponding negative effect on nitrate and phosphate availability. Our study has demonstrated that a better understanding of spatial variation in the effects of herbivores on ecosystems can emerge by evaluating their influences across gradients of soil texture, soil moisture, and herbivore activity. These data enabled us to evaluate several frameworks that have been developed to understand the variable effects of herbivores on ecosystems, which is a significant step in reconciling the many competing ideas put forth to explain the context‐dependent effects of large herbivores on grazed ecosystems.

Consumer-resource dynamics in Arctic ponds.

Population dynamics are shaped by species interactions with resources, competitors, enemies, and environmental fluctuations that alter the strength of these interactions. We used a food web approach to investigate the population dynamics of an abundant Arctic mosquito species, Aedes nigripes (Diptera: Culicidae). Specifically, we evaluated the importance of bottom-up variation in aquatic biofilms (food) and top-down predation from diving beetles (Colymbetes dolabratus, Coleoptera: Dytiscidae) on mosquito population performance. In spring 2018, we tracked mosquito and predator populations across eight ponds in western Greenland, measured biofilm productivity with standardized samplers, and estimated grazing pressure by invertebrate consumers with an in situ exclosure experiment. We also assessed the quality of biofilms as nutrition for mosquito larvae and evaluated pond attributes that might influence biofilm productivity and food quality. Our results indicated that mosquito population dynamics were more related to resource quality and intraspecific competition than to the density of predaceous diving beetles. Ponds with better quality biofilm tended to have more hatching larvae and those populations experienced higher per capita mortality. This aggregation of larvae in what would otherwise be the best mosquito ponds was enough to produce relatively low fitness. Thus, the landscape would support more mosquitoes if they instead distributed themselves to match predictions of the ideal free distribution. Although mortality rates were highest in ponds with the highest initial densities, the increased mortality was not generally enough to compensate for initial abundance, and 78% of the variation in the density of mosquitoes emerging from ponds was explained by the initial number of larvae in a pond. Resource quality was a strong predictor of consumer abundance, yet there was no evidence that biofilm grazing pressure was greater in ponds where mosquito density was higher. Collectively, our results suggest that mosquito ponds in western Greenland are a mosaic of source and pseudo-sink populations structured by oviposition tendencies, biofilm resource quality, and density-dependent larval mortality.

Long-term and interactive effects of different mammalian consumers on growth, survival, and recruitment of dominant tree species.

Throughout the world, numerous tree species are reported to be in decline, either due to increased mortality of established trees or reduced recruitment. The situation appears especially acute for oaks, which are dominant features of many landscapes in the northern hemisphere. Although numerous factors have been hypothesized to explain reductions in tree performance, vertebrate herbivores and granivores may serve as important drivers of these changes. Here, using data from 8‐ and 14‐year‐old exclosure experiments, we evaluated the individual and interactive effects of large and small mammalian herbivores on the performance of three widespread oak species in California—coast live oak (Quercus agrifolia), California black oak (Q. kelloggii), and Oregon white oak (Q. garryana). Although impacts varied somewhat by species and experiment, herbivory by black‐tailed deer (Odocoileus hemionus columbianus) reduced the height and survival of juvenile coast live oaks and altered their architecture, as well as reduced the abundance of black oak seedlings, the richness of woody species and the cover of nonoak woody species. Small mammals (Microtus californicus and Peromyscus maniculatus) had even more widespread effects, reducing the abundance of black oak seedlings and the height and cover of all three oak species. We also detected numerous interactions between small mammals and deer, with one herbivore having positive or negative effects on oak abundance and cover when the other herbivore was either present or absent. For example, deer often had negative effects on seedling abundance only when, or even more so when, small mammals were present. In summary, mammalian consumers play crucial roles in limiting oak recruitment by reducing seedling abundance, maintaining trees in stunted states, and preventing them from reaching sapling stages and becoming reproductive. Interactions between large and small mammals can also alter the intensity and direction of their effects on trees.

Wave exposure reduces herbivory in post-disturbed reefs by filtering species composition, abundance and behaviour\xa0of key fish herbivores

Harsh environmental conditions limit how species use the landscape, strongly influencing the way assemblages are distributed. In the wake of repeated coral bleaching mortalities in Lakshadweep, we examined how wave exposure influences herbivory in exposed and sheltered reefs. We used a combination of i. field observations of fish herbivore composition, abundance and activity across 6 exposed and 6 sheltered reefs; ii. experimental manipulations in a subset of these reefs (herbivore exclosures); and iii. opportunistic observations of fish recruitment, to determine how exposure influences herbivore biomass and herbivory. Species richness, biomass, abundance, total bite rates and species-specific per capita bite rates were lower in exposed compared to sheltered reefs, linked to strong environmental filtering of species composition, abundance and behaviour. For some critical species, this environmental filtering begins with differential recruitment and post-recruitment processes between exposures. Bite rates at sheltered sites were dominated by just a few species, most being laterally compressed surgeonfish that may find it difficult accessing or surviving in wave-battered shallow reefs. Exclosure experiments confirmed that exposed reefs were less controlled by herbivores than sheltered reefs. In post-disturbed reefs like Lakshadweep, environmental gradients appear to be key mediators of critical functions like herbivory by determining species composition, abundance and behaviour.