Herbivore Density Research Articles

Livestock density affects species richness and community composition of butterflies: A nationwide study

Extensively managed grasslands are globally recognized for their high biodiversity value. Over the past century, a continuous loss and degradation of grassland habitats has been observed across Europe that is mainly attributable to agricultural intensification and land abandonment. Particularly insects have suffered from the loss of grassland habitats due to land-use change and the decrease in habitat quality, either due to an increase in livestock density, higher mowing frequency, and an increase in nitrogen fertilization, or by abandonment. However, only a few studies have used nationwide datasets to analyse the effects of land cover and land-use intensity on insects. It further remains largely unexplored how these effects are modulated by species traits, i.e. habitat specialisation and mobility. Using nationwide butterfly data originating from the German Butterfly Monitoring Scheme, we investigated the effect of three indicators related to land cover and agricultural land-use intensity on species richness as well as trait composition of butterfly communities. Based on agricultural census data at the municipality scale, we calculated the share of permanent grasslands (measure of habitat availability), the total livestock density (proxy for organic fertilization) and the livestock density of domestic herbivores (proxy for management intensity in grasslands) within a 2 km buffer surrounding each butterfly transect. To analyse the relationships between butterflies and indicators of land cover and land-use intensity, we applied generalised linear mixed effect models. We found a negative relationship between butterfly species richness and the livestock density of domestic herbivores. Further, the ratio of butterfly generalist to specialist species shifted towards generalists and the size of butterflies increased with higher herbivore livestock density, indicating a shift in communities towards mobile habitat generalists. Our results are in accordance with previous studies carried out across smaller geographic extents, highlighting the importance of low herbivore livestock densities to halt the loss of pollinating insects and safeguard biodiversity and associated ecosystem services in agricultural landscapes. We here demonstrate that indicators based on livestock distribution data at the municipality scale can provide insights into processes and spatial diversity patterns of butterflies at the national level. Further, we highlight potentials and limitations of using agricultural census data to quantify and assess effects of land cover and land-use intensity on butterflies, and make recommendations for further research needs.

Grazing by semi‐feral cattle and horses supports plant species richness and uniqueness in grasslands

AbstractQuestionHow does naturalistic grazing (trophic rewilding with large herbivores), in contrast to mowing and free succession (no grazing), affect plant community composition and species richness in temperate grassland grazed by semi‐feral cattle and horses?LocationMols Laboratory, Denmark.MethodsWe investigated grazing exclosures in the rewilding area of the Mols Laboratory, four years after its establishment. We focused on moist to dry grassland vegetation, that is, excluding scrub and woodland. Each experimental block consisted of five 5 × 9 m plots, representing four fenced treatments, that is, summer‐only grazing, winter‐only grazing, full exclosure with annual autumn mowing and full exclosure with passive succession. The matrix (the fifth treatment) was grazed by large herbivores at close‐to‐natural densities, that is, regulated bottom‐up by the carrying capacity of the area. Hence, even the seasonal grazing treatments were grazed at close‐to‐natural animal density. Quantitative plant community composition was assessed using the point‐intercept method in 25 × 25 cm quadrats, supplemented with biomass calibration models based on additional quadrats, in which above‐ground plant biomass was harvested after recording and the material sorted to species and weighed. Uniqueness was assessed as the sum of inverse range sizes for constituent species (unicity).ResultsWe found an appreciably higher plant species richness in grazing treatments than under both annual mowing and full exclosure, but only minor differences between seasonal grazing treatments. Uniqueness was highest in year‐round and winter‐only grazing and lowest in summer‐only grazing. The forb:graminoid ratio tended to be high in the winter‐only grazing treatment, whereas annual mowing was associated with dominance of graminoids over forbs. Full‐exclosure plots had accumulation of litter and the lowest species richness. Initial heterogeneity between plots within blocks and a systematic difference between blocks in moist and dry grasslands may have diluted treatment effects at this early point after the onset of the experiment. Data analysis using the biomass estimates derived from the calibration models yielded only minor differences in the patterns described above, when compared to the results obtained using the raw number of intercepts.ConclusionsNaturalistic grazing is a goal in itself in ecological restoration, but also proposed as an efficient management tool to promote conservation of grassland plants and communities. We found both plant species richness and the prevalence of regionally rarer species (uni) to be higher with grazing than mowing or abandonment. Similarly, the tendency for forbs to prevail under grazing may translate into enhanced floral resources for anthophilous insects. Summer‐only grazing at low density of large herbivores was not significantly different from winter‐only and year‐round grazing, but this treatment was much closer to natural grazing than intensive summer grazing typical of agri‐environmental practices.

Pyric Herbivory and the Nexus Between Forage, Fire and Native and Introduced Large Grazing Herbivores in Australian Tropical Savannas

Earth’s tropical savannas typically support high biomass of diverse grazing herbivores that depend on a highly fluctuating resource: high-quality forage. An annual wet–dry cycle, fire and herbivory combine to influence forage quality and availability throughout the year. In the savannas of northern Australia, a depauperate suite of large native (marsupial) herbivores (wallaroos [Osphranter spp.] and the agile wallaby [Notamacropus agilis]) compete for resources with non-native large herbivores introduced in the late nineteenth century, particularly bovines (feral and managed cattle [Bos spp.] and feral water buffalo [Bubalus bubalis]) that now dominate the landscape. Anecdotal reports of recent population declines of large macropods and negative impacts of bovines highlight the need to better understand the complex relationship between forage, fire and abundance of native and introduced large herbivores. The pyric herbivory conceptual model, which posits complex feedbacks between fire and herbivory and was developed outside Australia, predicts that native and introduced large herbivores will both respond positively to post-fire forage production in Australian savannas where they co-occur. We used grazing exclosures, forage biomass and nutrient analyses and motion-sensor camera-trapping to evaluate the overall robustness of the pyric herbivory model in the Australian context, specifically whether forage quantity and quality are impacted by herbivory, season and fire activity, and which forage attributes most influence large grazing herbivore abundance. Forage quantity, as measured by live, dead and total herbaceous biomass and proportion of biomass alive, was higher inside herbivore exclosures, even at relatively low densities of herbivores. Forage quality, as measured by fibre content, was not affected by herbivory, however, crude protein content of live herbaceous biomass was greater outside herbivore exclosures. Recent fire was an important predictor of all measures of forage quantity and quality. Recent fire occurrence decreased overall quantity (biomass) but increased quality (decreased fibre content and increased crude protein content); late dry season fires resulted in forage with the highest crude protein content. The predictions of the pyric herbivory conceptual model are consistent with observations of the feeding behaviour of introduced bovines and some large macropods in northern Australian savannas, lending support to the global generality of pyric herbivory in fire-prone grassy biomes.

Association between arthropod densities suggests dominance of top-down control of predator-prey food-webs on pear trees during winter

Although winter in the temperate zone is considered to be the period of arthropod quiescence, some pests and their natural enemies remain active and interact in simplified food-webs. Limited information exists about the relative importance of top-down and bottom-up processes regulating arthropod food-webs and their spatio-temporal dynamics during winter. This information is essential for the development of effective conservation biocontrol methods. We investigated how the pest management of pear orchards (integrated [IPM] vs organic) and tree location within orchards (margin vs centre) influence the association between densities of winter-active spiders, insect herbivores and winter-inactive predators (spiders, insects) from October to April. We installed carboard bands on trunks and branches of pear trees in four organic and four IPM orchards to collect bark-dwelling arthropods. We then modelled relationships between the densities of four functional arthropod groups, namely winter-active spiders, winter-inactive spiders, winter-inactive insect predators, and herbivores. In early winter, we found a hump-shaped relationship between the densities of winter-active spiders and herbivores. This agrees with the top-down model of the ecosystem exploitation hypothesis, predicting that predators are first bottom-up limited, then accumulate with prey densities, and exert top-down control when they become sufficiently abundant. The densities of herbivores strongly declined during winter, a phenomenon which may be partly, along with other causes of natural mortality, attributable to predation by winter active spiders. The association between winter-active spiders and winter-inactive arthropod predators switched from positive to none in organic orchards or even to a negative association in IPM orchards. Negative intraguild interactions seem to be intensified due to the declining availability of alternative prey. Overall, the investigated food-webs seem to be mostly top-down regulated during winter, and IPM seems to intensify intraguild interactions.

Impact of Transgenic Cry1Ab/2Aj Maize on Abundance of Non-Target Arthropods in the Field.

Transgenic Bacillus thuringiensis (Bt) maize has broad prospects for application in China. Before commercialization, it is necessary to assess possible ecological impacts, including impacts on non-target arthropods (NTAs) in the field. In the present study, transgenic Bt maize expressing cry1Ab/2Aj and its corresponding non-transformed near isoline were planted under the same environmental and agricultural conditions, and arthropods in the field were collected during the three main growth stages of maize. In a one year trial, the results showed the composition of NTA communities in the transgenic and control maize fields were similar. There were no significant differences for community-level parameters of species richness (S), Shannon–Wiener diversity index (H′), evenness index (J) and Simpson’s dominant concentration (C) between the two types of maize fields. Likewise, a Bray–Curtis dissimilarity and distance analysis showed that Cry1Ab/2Aj toxin exposure did not increase community dissimilarities between Bt and non-Bt maize plots and that the structure of the NTAs community was similar on the two maize varieties. Furthermore, planting of the transgenic cry1Ab/2Aj maize did not affect the density or composition of non-target decomposers, herbivores, predators, parasitoids and pollinator guilds. In summary, our results showed that planting of Bt maize producing Cry1Ab/Cry2Aj proteins do not adversely affect population dynamics and diversity of NTAs.

Enhanced top-down control of herbivore population growth on plants with impaired defences.

Herbivore densities can be regulated by bottom-up and top-down forces such as plant defences and natural enemies, respectively. These forces can interact with each other to increase plant protection against herbivores; however, how much complementarity exists between bottom-up and top-down forces still remains to be fully elucidated. Particularly, because plant defences can hinder natural enemies, how these interactions affect herbivore performance and dynamics remains elusive.To address this topic, we performed laboratory and greenhouse bioassays with herbivorous mite pests and predatory mites on mutant tomato plants that lack defensive hairs on stems and leaves. Particularly, we investigated the behaviour and population dynamics of different phytophagous mite species in the absence and presence of predatory mites.We show that predatory mites do not only perform better on tomatoes lacking defensive hairs but also that they can suppress herbivore densities better and faster on these hairless plants. Hence, top-down control of herbivores by natural enemies more than compensated the reduced bottom-up herbivore control by plant defences.Our results lead to the counter-intuitive insight that removing, instead of introducing, plant defence traits can result in superior protection against important pests through biological control. Read the free Plain Language Summary for this article on the Journal blog.

Chronic browsing by an introduced mammalian herbivore in a tropical island alters species composition and functional traits of forest understory plant communities

AbstractMammalian herbivores have large‐scale impacts on vegetation, altering structure, and species composition, especially in tropical grasslands and savannas. However, there is limited understanding of the potential impacts of mammalian herbivores in tropical wet forests, where they are typically less abundant. We investigated the effects of an introduced mammalian herbivore chital (Axis axis) on vegetation structure, composition and leaf functional traits of tropical evergreen forests of the Andaman Islands, India. Across seven islands, representing a gradient of herbivore densities, increasing chital presence was associated with decreased understory richness, understory density and adult tree richness, but was not related to adult tree density or size class distributions. We also found a significant decrease in community level leaf palatability traits (specific leaf area decreased and leaf thickness increased) with increasing chital habitat use. This community level shift in leaf trait values was better explained by intraspecific variation in leaf traits across islands rather than changes in species composition. In summary, we show persistent long‐term impacts of an introduced mammalian herbivore on understory tropical tree communities although there is little impact on adult tree communities. Our results also show that functional traits of species can be altered in response to novel herbivory, even at herbivore densities where there are no detectable impacts on adult forest structure or composition. Such altered functional traits may potentially alter ecosystem functioning in these forests even without changes in vegetation structure.Abstract in Hindi is available with online material.

Insectivorous birds reduce herbivory but do not increase mangrove growth across productivity zones.

Top-down effects of predators and bottom-up effects of resources are important drivers of community structure and function in a wide array of ecosystems. Fertilization experiments impose variation in resource availability that can mediate the strength of predator impacts, but the prevalence of such interactions across natural productivity gradients is less clear. We studied the joint impacts of top-down and bottom-up factors in a tropical mangrove forest system, leveraging fine-grained patchiness in resource availability and primary productivity on coastal cays of Belize. We excluded birds from canopies of red mangrove (Rhizophoraceae: Rhizophora mangle) for 13 months in zones of phosphorus-limited, stunted dwarf mangroves, and in adjacent zones of vigorous mangroves that receive detrital subsidies. Birds decreased total arthropod densities by 62%, herbivore densities more than fivefold, and reduced rates of leaf and bud herbivory by 45% and 52%, respectively. Despite similar arthropod densities across both zones of productivity, leaf and bud damage were 2.0 and 4.3 times greater in productive stands. Detrital subsidies strongly impacted a suite of plant traits in productive stands, potentially making leaves more nutritious and vulnerable to damage. Despite consistently strong impacts on herbivory, we did not detect top-down forcing that impacted mangrove growth, which was similar with and without birds. Our results indicated that both top-down and bottom-up forces drive arthropod community dynamics, but attenuation at the plant-herbivore interface weakens top-down control by avian insectivores.

Top-down regulation of hemlock woolly adelgid (Adelges tsugae) in its native range in the Pacific Northwest of North America.

The density of insect herbivores is regulated by top-down factors (e.g., natural enemies), bottom-up effects (e.g., plant defenses against herbivory), or a combination of both. As such, understanding the relative importance of these factors can have important implications for the establishment of effective management options for invasive species. Here, we compared the relative importance of top-down and bottom-up factors on the abundance of hemlock woolly adelgid (HWA), Adelges tsugae. HWA is invasive in eastern North America, but its native range includes the Pacific Northwest of North America where it has co-evolved with western hemlock, Tsuga heterophylla. Eastern hemlock, Tsuga canadensis, can also be found planted in city and park settings in the Pacific Northwest and the presence of both host species allowed us to directly compare the importance of predators (top-down) and host plant resistance (bottom-up) on HWA abundance by placing mesh exclusion bags on branches of both species and monitoring HWA abundance over two years. We found no evidence for bottom-up control of HWA on western hemlock (a native host). HWA established more readily on that species than on eastern hemlock on which it is a major pest in eastern North America. We found strong evidence for top-down control in that both summer and winter-active predators significantly reduced HWA densities on the branches of both tree species where predators were allowed access. These findings support the validity of the biological control program for HWA, the goal of which is to reduce outbreak populations of HWA in eastern North America.

A stronger role for long-term moisture change than for CO2 in determining tropical woody vegetation change.

Anthropogenically elevated CO2 (eCO2) concentrations have been suggested to increase woody cover within tropical ecosystems through fertilization. The effect of eCO2 is built into Earth system models, although testing the relationship over long periods remains challenging. Here, we explore the relative importance of six drivers of vegetation change in western Africa over the past ~500,000 years (moisture availability, fire activity, mammalian herbivore density, temperature, temperature seasonality, CO2) by coupling past environmental change data from Lake Bosumtwi (Ghana) with global data. We found that moisture availability and fire activity were the most important factors in determining woody cover, whereas the effect of CO2 was small. Our findings suggest that the role of eCO2 effects on tropical vegetation in predictive models must be reconsidered.

A multiproxy approach to long-term herbivore grazing dynamics in peatlands based on pollen, coprophilous fungi and faecal biomarkers

Herbivory plays a significant role in regulating many contemporary terrestrial plant ecosystems, but remains an imperfectly understood component of past ecosystem dynamics because the diagnostic capability of methods is still being tested and refined. To understand the efficacy of a multiproxy approach, we compare the sensitivity of pollen and coprophilous fungal spores (CFS) to changes in grazing intensity over the last 100–150 years in six peat cores from three UK upland areas, and apply faecal lipid biomarkers to two of the cores, using agricultural census data to calculate an independent record of herbivore density. Rising sheep density adversely affected moorland ecology over the last century, which therefore provides a suitable period to test the sensitivity of these proxies. In particular, we assess whether CFS can be used to track variations in large herbivore densities over time, since this has received less attention than their ability to identify high grazing levels. At selected sites, we test whether faecal lipid biomarkers can be used to identify which herbivore species were present. Our results highlight the differential sensitivity of each proxy, demonstrating on peat- and moorlands (i) that peak CFS abundance is a more consistent indicator of ecologically influential (high) herbivore levels than variations in animal density through time; (ii) when recorded with high CFS values, the decline or disappearance of grazing-tolerant pollen taxa is a reliable indicator of high herbivory; and (iii) at low herbivore densities, faecal lipid biomarkers are not an effective indicator of herbivore presence or identity. Quantitative reconstructions of past herbivory and identifying grazer species therefore remain challenging. However, our findings indicate that pollen and CFS provide complementary evidence for high intensity grazing, and emphasise that studies using CFS should aim to define ‘high’ herbivore levels in terms of the grazing sensitivity of the ecosystem, rather than relative animal abundance.

Population density and plant availability interplay to shape browsing intensity by roe deer in a deciduous forest

Browsing damage in forests relies on a complex interaction between herbivore density and forest understory composition and relative availability. Although variation in the amount of browsed twigs is sometimes used to assess abundance of large herbivores, the potential confounding effect of resource availability on this relationship has not been investigated yet. To fill the gap, we measured how browsing intensity of the woody plants varied in response to changes in both roe deer (Capreolus capreolus) abundance and vegetation availability from an intensive long-term monitoring. We estimated plant availability and consumption by roe deer from a modified Aldous method throughout a 14 year-long period during which we experimentally manipulated population density. The functional response was strongly non-linear and density-dependent. When plant availability was low (<12.5%), browsing intensity strongly increased with plant availability with an increasing rate of roe deer density, whereas beyond this threshold, browsing intensity slightly increased with both plant availability and population density in an additive way. Thus, forest susceptibility to browsing increases with increasing competition for food, especially when plant availability is low. Moreover, the interplay between browsing intensity and population density at low plant availability prevents the use of browsing intensity to monitor roe deer abundance when plant availability is low. Our findings provide clear evidence that relying on key ecological concepts such as functional responses improves the accuracy of management tools when monitoring changes of the herbivore-plant system over time.

A legacy of fire emerges from multiple disturbances to most shape microbial and nitrogen dynamics in a deciduous forest

Healthy forests are vital components of terrestrial ecosystems for their raw materials, high biodiversity, cycling of nutrients, and potential to sequester carbon. However, these ecosystems are sensitive to disturbances, and anthropogenic activities pose a serious threat to forest ecosystems globally. For example, human activities have dramatically altered multiple historical disturbance regimes in forests, including suppressing fire, increasing the density of large herbivores, and reducing the size of canopy gaps, among other disturbances. Such disturbances can have dramatic impacts on microbially-mediated forest soil functions, but more research is needed to determine the collective impacts of these disturbances. In this study, we investigated the interactive effects of disturbances, namely the legacies of fire, large herbivore densities, and canopy gap creation, in a deciduous forest soil. We determined that forest floor and mineral soil carbon and nitrogen pools were shaped by multiple disturbances, but fire was more influential than the other disturbances. The abundance of several functionally-relevant microbial taxa were significantly impacted by fire, and the effect was more pronounced in the mineral soil than in the forest floor. Together, these findings demonstrate that multiple disturbances, especially a legacy of fire, exerts long-term control over soil carbon, nitrogen and microbial dynamics in a deciduous forest system.

Browsing herbivores improve the state and functioning of savannas: A model assessment of alternative land-use strategies.

Changing climatic conditions and unsustainable land use are major threats to savannas worldwide. Historically, many African savannas were used intensively for livestock grazing, which contributed to widespread patterns of bush encroachment across savanna systems. To reverse bush encroachment, it has been proposed to change the cattle‐dominated land use to one dominated by comparatively specialized browsers and usually native herbivores. However, the consequences for ecosystem properties and processes remain largely unclear. We used the ecohydrological, spatially explicit model EcoHyD to assess the impacts of two contrasting, herbivore land‐use strategies on a Namibian savanna: grazer‐ versus browser‐dominated herbivore communities. We varied the densities of grazers and browsers and determined the resulting composition and diversity of the plant community, total vegetation cover, soil moisture, and water use by plants. Our results showed that plant types that are less palatable to herbivores were best adapted to grazing or browsing animals in all simulated densities. Also, plant types that had a competitive advantage under limited water availability were among the dominant ones irrespective of land‐use scenario. Overall, the results were in line with our expectations: under high grazer densities, we found heavy bush encroachment and the loss of the perennial grass matrix. Importantly, regardless of the density of browsers, grass cover and plant functional diversity were significantly higher in browsing scenarios. Browsing herbivores increased grass cover, and the higher total cover in turn improved water uptake by plants overall. We concluded that, in contrast to grazing‐dominated land‐use strategies, land‐use strategies dominated by browsing herbivores, even at high herbivore densities, sustain diverse vegetation communities with high cover of perennial grasses, resulting in lower erosion risk and bolstering ecosystem services.

Warming Alters the Relationship Between Benthic Cover and Herbivores on Hawaiian Reefs

Increases in sea surface temperature impact animal metabolism, which in turn could influence benthic structure and resulting algal-coral balance. We utilized a long-term coral reef dataset from the west coast of Hawai‘i Island to investigate impacts of annual positive and negative sea surface temperature anomalies (SSTA) on benthic cover [algal turf, macroalgae, crustose coralline algae (CCA), and coral], herbivore density (sea urchins, grazers, browsers, and scrapers) and the relationship between benthic cover and herbivore density. Results showed significantly lower coral cover, but higher CCA cover with positive SSTA. Additionally, the density of sea urchins, grazers and browsers increased with increasing SSTA. Warming disrupted the normal relationship between herbivores and benthic cover on reefs, particularly for grazers where higher densities were coupled with lower algal turf cover only during negative SSTA. The direction of the relationship between benthic cover and herbivore type changed with positive SSTA, where increased algal turf cover was associated with increased herbivore density. Here, herbivores are likely responding accordingly to increases in food availability due to increased metabolism under warming. Despite herbivore populations increasing in density over the past two decades, algal turf cover remains on an upward trajectory. These results indicate that warming can alter herbivore-algal dynamics, where greater herbivore densities may be required to cause a reduction in algal turf cover. Protection of herbivores in addition to reducing nutrient input onto reefs will be essential in driving a reduction in algal turf cover on Hawaiian reefs.

Host plant limitation of butterflies in highly fragmented landscapes

Insect herbivores can be limited by host plants in two ways: density-dependent competition for food resources or density-independent search time limitation. Our understanding of density-dependent host plant limitation is relatively well developed and well integrated into conservation plans for at-risk insects. Search time limitation, a density-independent process, is much less well developed. Here, we explore both mechanisms using empirically based models of monarch butterfly population dynamics. These mechanisms differ fundamentally in their predictions: resource competition leads to matching of herbivore densities to host plant densities, and visible competition via consumption of host plants. Search time limitation leads to changes in population growth rate that can cause herbivore numbers to decline when host plant densities are constant. Search time limitation also implies that host plants can limit herbivores, even when many individual plants are uneaten. For monarch butterflies, our calculations suggest that many parts of North America have host plant densities below the threshold for search time limitation, which contrasts with the typical assumption of resource competition. More generally, incorporating search time limitation into conservation plans is important for reframing our understanding of how host plants limit insect herbivores in highly fragmented landscapes.

Functional traits of alpine plant communities show long‐term resistance to changing herbivore densities

AbstractHerbivores shape vegetation by suppressing certain plant species while benefitting others. By thus modifying plant species functional composition, herbivores affect carbon cycling, albedo, vegetation structure and species' interactions. These effects have been suggested to be able to counteract the effects of increasing temperatures on vegetation in alpine environments. Managing the dominant large ungulates in these ecosystems could thus provide a tool to mitigate climate change effects. However, it is possible that legacy effects of past grazing will dampen ungulate impacts on vegetation. We shed a light on this topic by investigating the short‐ and long‐term effects of varying sheep densities on the plant trait composition in the Norwegian alpine tundra with centuries‐long of intensive grazing history. In the first part of our study, we quantified the effects of sheep on the plant community functional trait composition at different elevations and under moderate and low productivity in. We combined data from two long‐term (14 and 19 yr) sheep fence experiments and showed that differences in sheep densities did not affect plant trait composition, irrespective of productivity. However, in the second part of our study, we showed that the plant trait composition in mainland (that has been grazed for centuries) differed from vegetation on islands which have been herbivore‐free. Taken together, these results suggest that sheep have an effect on the alpine plant communities on historical time scales covering centuries, but that the resulting sheep grazing resistant/tolerant communities may not respond to shorter‐term (14 and 19 yr) changes in sheep densities, that is, at temporal scales relevant for ecosystem management. Furthermore, we showed that the plant trait composition at the site with low productivity had gone through a temporal trait change independent of sheep treatment, potentially due to increased temperatures and precipitation, suggesting that sheep may not be able to counteract climatic impacts in the areas with centuries‐long grazing history.

Overabundant native herbivore impacts on native plant communities in south‐eastern Australia

SummaryNative herbivores can affect the structure and function of plant communities. Most studies of overabundant native herbivore effects on native plant communities in south‐eastern Australia are observational or compare, at most, two levels of herbivory (ambient vs. none), and typically do not separate out the impacts of multiple herbivores. Several studies show that high densities of herbivores had mostly negative effects on plant regeneration and growth. These effects were usually attributed to palatability differences amongst plant species. Surprisingly, few studies have quantified vegetation responses across a gradient of herbivore density, and impacts on plant communities (i.e. species richness, composition, beta diversity) have not been well studied. Hence, there remains a need to better understand how native herbivores impact plant communities at different densities.

LeopardPanthera pardusdensity and survival in an ecosystem with depressed abundance of prey and dominant competitors

AbstractThe leopardPanthera pardusis in range-wide decline, and many populations are highly threatened. Prey depletion is a major cause of global carnivore declines, but the response of leopard survival and density to this threat is unclear: by reducing the density of a dominant competitor (the lionPanthera leo) prey depletion could create both costs and benefits for subordinate competitors. We used capture–recapture models fitted to data from a 7-year camera-trap study in Kafue National Park, Zambia, to obtain baseline estimates of leopard population density and sex-specific apparent survival rates. Kafue is affected by prey depletion, and densities of large herbivores preferred by lions have declined more than the densities of smaller herbivores preferred by leopards. Lion density is consequently low. Estimates of leopard density were comparable to ecosystems with more intensive protection and favourable prey densities. However, our study site is located in an area with good ecological conditions and high levels of protection relative to other portions of the ecosystem, so extrapolating our estimates across the Park or into adjacent Game Management Areas would not be valid. Our results show that leopard density and survival within north-central Kafue remain good despite prey depletion, perhaps because (1) prey depletion has had weaker effects on preferred leopard prey compared to larger prey preferred by lions, and (2) the density of dominant competitors is consequently low. Our results show that the effects of prey depletion can be more complex than uniform decline of all large carnivore species, and warrant further investigation.

Ecological and environmental context shape the differential effects of a facilitator in its native and invaded ranges.

Invasive species often exhibit disproportionately strong negative effects in their introduced range compared to their native range, and much research has been devoted to understanding the role of shared evolutionary history, or lack thereof, in driving these differences. Less studied is whether introduced species, particularly those that are important as facilitators in their native range, have persistent positive effects in their invaded range despite a lack of a shared evolutionary history with the invaded community. Here, we manipulated the density of a habitat-forming facilitator, the high intertidal acorn barnacle Balanus glandula, factorially with herbivore density in its native range (Bluestone Point, British Columbia, Canada) and invaded range (Punta Ameghino, Chubut Province, Argentina) to determine how this facilitator differentially affects associated species at these two locations. Given that high intertidal species at Punta Ameghino (PA) are evolutionarily naïve to barnacles, we predicted that the positive effects of B. glandula at PA would be absent or weak compared to those at Bluestone Point (BP). However, we found that B. glandula had an equally positive effect on herbivore biomass at PA compared to BP, possibly because the moisture-retaining properties of barnacle bed habitats are particularly important in seasonally dry Patagonia. Barnacle presence indirectly decreased ephemeral algal cover at BP by increasing grazer pressure, but barnacles instead facilitated ephemeral algae at PA. In contrast, B. glandula increased perennial algal cover at BP, but generally decreased perennial algal cover at PA, likely due to differences in dominant algal morphology. Though our experiment was limited to one location on each continent, our results suggest that shared evolutionary history may not be a prerequisite for strong facilitation to occur, but rather that the nature and strength of novel species interactions are determined by the traits of associated species and the environment in which they occur.