The habitat preferences of invasive raccoon dog imply elevated risks for wetland-associated prey species.

Habitat preferences of invasive predators determine where and for which prey species they pose a threat upon. This is crucial information for the conservation of endangered prey species because invasive predators pose additional predation on top of that caused by natural predators. In large parts of Europe, the most common invasive mesopredator is the raccoon dog (Nyctereutes procyonoides). To understand the risk that the raccoon dog poses for wetland-associated species, we collected information about its habitat preferences near these habitats. We used data on 24 GPS-collared raccoon dogs from three landscape types in Finland, to study their spatial and temporal habitat preferences. We first determined their home ranges, within which we then examined habitat use and preferences. Raccoon dogs showed generalist habitat use, which was evident in their wide range of used habitats. However, in spring and summer, during the breeding seasons of waterfowl and amphibians, they preferred wetlands and peatlands. They also preferred shorelines and the edges of forests and agricultural fields. During autumn and winter, raccoon dogs did not prefer wetlands. These findings support the conclusion that the raccoon dog's habitat preferences pose a particular threat to wetland-associated species, such as nesting waterfowl and amphibians. The species' habitat preferences coupled with high numbers of this invasive mesopredator pose additional predation for endangered wetland-associated species on top of that of native predators.

Mercury bioaccumulation and its relationship with trophic biomarkers in a Mediterranean elasmobranch mesopredator.

Human activity has led to increased concentrations of mercury (Hg) in the world's oceans. Mercury can bioaccumulate and biomagnify in animal tissues via trophic transfer, thus, becoming most pronounced in larger and older predators. Here, we measured Hg concentrations and their relationship with stable isotopes-based proxies of trophic level (δ13C and δ15N values) in multiple tissues of Mustelus spp. from the Mediterranean Sea. We found higher Hg concentrations in muscle than in liver and fin tissues. The relationship between Hg concentrations and δ15N values in muscle suggested repeated foraging for low trophic level and Hg-poor prey, and biomagnification of Hg at higher trophic levels. Seasonal variations in δ13C values could indicate shifts in primary production sources and/or in local prey availability. The HBVSe index suggested no risk to human health, however the safe meal limit recommendations are 4.5 and 2.2 portions per month for adults and children, respectively.

Crowding in the middle of marine food webs: A focus on Raja asterias and other mediterranean batoids.

The Mediterranean Sea is among the three biodiversity hotspots of the world where elasmobranchs are severely threatened. Elasmobranchs act as apex or meso-predators within marine food webs and the loss/decline of apex predators determines the mesopredator release, leading in turn to increased predation on smaller prey. However, also several mesopredators (including rays, skates and small sharks) are intensively fished, being of commercial interest, or by-caught, and thus mesopredators increase could not be so evident. We analysed the trophic ecology of an endemic Mediterranean ray, the starry ray Raja asterias, at a seasonal scale from the Adriatic basin, one of the most intensively exploited area of the Mediterranean, by means of stomach contents and stable isotopes analyses. Our results evidenced that starry rays rely on benthic sources including species of local commercial values, such as swimming crabs, small cephalopods, and stomatopods and share the same trophic position with other elasmobranchs (rays, skates, and small sharks) and other mesopredators (e.g., common soles, Norway lobsters and mullets). As all mesopredators are overexploited, as well as their benthic prey are affected by intense trawl-fishing, the whole food webs are disrupted and neither the classical trophic cascade nor the mesopredator release hypothesis could be verified. Conservation measures for these species, such as the release after capture or the application of exclusion grids to the net, should be applied in areas where populations are strongly impacted by trawling.

Visitation of artificial watering points by the red fox (Vulpes vulpes) in semiarid Australia.

The introduced red fox (Vulpes vulpes) now occupies most of the Australian continent outside the tropics, including arid and semiarid ecosystems. Information on the water requirements of foxes is scant, but free water is not thought to be required if adequate moisture-containing food is available. The frequency and duration of visits by foxes fitted with GPS collars to known artificial watering points in semiarid Australia were recorded for 22 individual foxes across four austral seasons between October 2015 and November 2017, providing >93,000 location fixes. We modeled home range and the distance traveled by range-resident foxes beyond their home range to reach known water sources. We used recurse analysis to determine the frequency of visitation and step-selection functions to model the speed and directionality of movement inside and outside the home range. Our study demonstrates that some foxes in this semiarid environment utilize free-standing water. The findings suggest that artificial watering points can be used as a focal point for conducting strategic fox control in arid and semiarid environments. Additionally, strategies that restrict access to water by foxes may reduce their duration of occupancy and/or long-term abundance in parts of the landscape, thus providing benefits for conservation and agriculture.

Revisiting the paradigm of shark-driven trophic cascades in coral reef ecosystems.

Global overfishing of higher-level predators has caused cascading effects to lower trophic levels in many marine ecosystems. On coral reefs, which support highly diverse food webs, the degree to which top-down trophic cascades can occur remains equivocal. Using extensive survey data from coral reefs across the relatively unfished northern Great Barrier Reef (nGBR), we quantified the role of reef sharks in structuring coral reef fish assemblages. Using a structural equation modeling (SEM) approach, we explored the interactions between shark abundance and teleost mesopredator and prey functional group density and biomass, while explicitly accounting for the potentially confounding influence of environmental variation across sites. Although a fourfold difference in reef shark density was observed across our survey sites, this had no impact on either the density or biomass of teleost mesopredators or prey, providing evidence for a lack of trophic cascading across nGBR systems. Instead, many functional groups, including sharks, responded positively to environmental drivers. We found reef sharks to be positively associated with habitat complexity. In turn, physical processes such as wave exposure and current velocity were both correlated well with multiple functional groups, reflecting how changes to energetic conditions and food availability, or modification of habitat affect fish distribution. The diversity of species within coral reef food webs and their associations with bottom-up drivers likely buffers against trophic cascading across GBR functional guilds when reef shark assemblages are depleted, as has been demonstrated in other complex ecosystems.

Culling recolonizing mesopredators increases livestock losses: Evidence from the South African Karoo.

Populations of adaptable mesopredators are expanding globally where passive rewilding and natural recolonization are taking place, increasing the risk of conflict with remaining livestock farmers. We analysed data from two social surveys of farmers in the Karoo, South Africa, where black-backed jackals (Canis mesomelas) and caracals (Caracal caracal) have re-emerged as a threat to sheep farms in the context of falling agricultural employment and the expansion of natural areas. We show that irrespective of measurement approach, lethal control of mesopredators in this fragmented socio-economic landscape was associated with increased livestock losses the following year. Terrain ruggedness was positively, and number of farmworkers negatively, associated with livestock losses. Our study provides further evidence that lethal control of mesopredators in this context is probably counter-productive and supports calls to develop, share and financially support a range of non-lethal methods to protect livestock, especially where natural recolonization of mesopredators is occurring. A graphical abstract can be found in Electronic supplementary material.

Novel predators and anthropogenic disturbance influence spatio-temporal distribution of forest antelope species.

Understanding the effects of anthropogenic disturbance on species' behaviour is crucial for conservation planning, considering the extent of habitat loss. We investigated the influence of anthropogenic disturbances including agriculture, urbanisation, protected areas, and the presence of novel predators, on the temporal and spatial behaviour of sympatric forest antelope (Tragelaphus scriptus, Philantomba monticola, Sylvicapra grimmia, and Cephalophus natalensis) in an anthropogenic matrix containing forest fragments in the Indian Ocean Coastal Belt of South Africa. We integrated land-use types with camera-trap data and compared activity patterns using circular statistics and occupancy modelling. Antelope species overlapped in temporal and spatial distribution and exhibited diurnal activity for 50% of the time. All species exhibited nocturnal activity for ∼25-33% of all observations, except for C. natalensis. Nocturnal activity varied between species and land-use types. The predators Canis familiaris, C. mesomelas and Caracal caracal negatively influenced occupancy of P. monticola, S. grimmia and C. natalensis. Humans negatively influenced temporal activity of P. monticola, and spatial distribution of T. scriptus and S. grimmia. C. familiaris had an overall negative influence on S. grimmia. We found spatial, and to a lesser extent temporal, segregation between species. However, plasticity in activity patterns existed, which varied in response to anthropogenic effects and novel predators.

Top carnivore decline has cascading effects on scavengers and carrion persistence.

Top carnivores have suffered widespread global declines, with well-documented effects on mesopredators and herbivores. We know less about how carnivores affect ecosystems through scavenging. Tasmania's top carnivore, the Tasmanian devil (Sarcophilus harrisii), has suffered severe disease-induced population declines, providing a natural experiment on the role of scavenging in structuring communities. Using remote cameras and experimentally placed carcasses, we show that mesopredators consume more carrion in areas where devils have declined. Carcass consumption by the two native mesopredators was best predicted by competition for carrion, whereas consumption by the invasive mesopredator, the feral cat (Felis catus), was better predicted by the landscape-level abundance of devils, suggesting a relaxed landscape of fear where devils are suppressed. Reduced discovery of carcasses by devils was balanced by the increased discovery by mesopredators. Nonetheless, carcasses persisted approximately 2.6-fold longer where devils have declined, highlighting their importance for rapid carrion removal. The major beneficiary of increased carrion availability was the forest raven (Corvus tasmanicus). Population trends of ravens increased 2.2-fold from 1998 to 2017, the period of devil decline, but this increase occurred Tasmania-wide, making the cause unclear. This case study provides a little-studied potential mechanism for mesopredator release, with broad relevance to the vast areas of the world that have suffered carnivore declines.

Multiple predator effects on juvenile prey survival.

Predicting multiple predator effects (MPEs) on shared prey remains one of the biggest challenges in ecology. Empirical evidence indicates that interactions among predators can alter predation rates and modify any expected linear effects on prey survival. Knowledge on predator density, identity and life-history traits is expected to help predict the behavioral mechanisms that lead to non-linear changes in predation. Yet, few studies have rigorously examined the effects of predator-predator interactions on prey survival, particularly with marine vertebrate predators. Using an additive-substitutive design, we experimentally paired reef piscivores with different hunting mode [active predator, Pseudochromis fuscus (F); ambush predators, Cephalopholis boenak (B), Epinephelus maculatus (M)] to determine how behavioral interactions modified their combined impacts on damselfish prey. Results showed that behavioral patterns among predators matched those predicted from their hunting mode. However, it was the identity of the predators what determined the strength of any positive or negative interactions, and thus the nature and magnitude of MPEs on prey survival (i.e., risk-enhancing effects: treatments BB, MM and FM; risk-reducing: BM; and linear effects: FF, FB). Given the specificity of predator-predator interactions, none of the predators were fully functionally redundant. Even when two species seemed substitutable (i.e., predators F and M), they led to vastly diverse effects when paired with additional predator species (i.e., B). We concluded that knowledge of the identity of the predator species and the behavioral interactions among them is crucial to successfully predict MPEs in natural systems.

Agroforestry system as a buffer zone in a Brazilian Atlantic Forest conservation unit: an artificial nest predation perspective / Sistema Agroflorestal como zona de amortecimento em uma Unidade de Conservação da Mata Atlântica brasileira: uma perspectiva de predação de ninho artificial

Abstract Here we inferred about the effectiveness of a buffer zone composed by an Agroforestry System (AS) for providing reproductive conditions for Atlantic Forest birds by comparing artificial nest predation rates between a buffer zone, a primary forest area, and a set of small and isolated Atlantic Forest fragments. We have used 237 nests throughout the three areas, of which 100 (42.2%) were depredated. Predation frequencies were 28.6% in the continuous forest, 100% in the AS, and 51.9% in the fragments, differing significantly. We could detect nest predators in 48 of the 100 depredated nests (48%) using camera traps. Differences between areas may be attributed to changes in nest predator species. We support previous evidences that the conservation of bird communities may not rely on buffer zones as an extension of their breeding habitats.

Resumo O presente estudo teve como objetivo avaliar a efetividade de uma zona tampão composta por um sistema agroflorestal (SA) para a reprodução de aves da Mata Atlântica. Para isto, as porcentagens de predação em ninhos artificiais foram comparadas entre as zonas tampão, uma área de floresta primária e em um conjunto de fragmentos de Mata Atlântica pequenos e isolados. Um total de 237 ninhos foi utilizado nas três áreas e 100 deles (42,2%) foram predados. As porcentagens de predação foram 28,6% na área de floresta primária, 100% no SA e 51,9% nos fragmentos, havendo diferença significativa. Os predadores puderam ser identificados em 48 das 100 predações (48%) com o uso de câmeras trap, o que gerou evidências de que as diferenças possam ser atribuídas a mudanças na composição de espécies de predadores entre as áreas. Esta análise dá suporte a estudos prévios de que as zonas tampão não devam ser consideradas como extensão dos hábitats naturais para a conservação de comunidades de aves.

Seed dispersal of Diospyros virginiana in the past and the present: Evidence for a generalist evolutionary strategy.

Several North American trees are hypothesized to have lost their co-evolved seed disperser during the late-Pleistocene extinction and are therefore considered anachronistic. We tested this hypothesis for the American persimmon (Diospyros virginiana) by studying the effects of gut passage of proposed seed dispersers on seedling survival and growth, natural fruiting characteristics, and modern animal consumption patterns. We tested gut passage effects on persimmon seeds using three native living species, the raccoon (Procyon lotor), Virginia opossum (Didelphis virginiana), and coyote (Canis latrans), and two Pleistocene analogs; the Asian elephant (Elephas maximus) and alpaca (Vicugna pacos). Persimmon seeds excreted by raccoons, coyotes, and elephants survived gut transit. Gut passage did not affect sprouting success, but did tend to decrease time to sprout and increase seedling quality. Under field conditions, persimmon fruits were palatable on the parent tree and on the ground for an equal duration, but most fruits were consumed on the ground. Seven vertebrate species fed upon persimmon fruits, with the white-tailed deer (Odocoileus virginianus)-a species not capable of dispersing persimmon seeds-comprising over 90% of detections. Conversely, potential living seed dispersers were rarely detected. Our results suggest the American persimmon evolved to attract a variety of seed dispersers and thus is not anachronistic. However, human-induced changes in mammal communities could be affecting successful seed dispersal. We argue that changes in the relative abundance of mammals during the Anthropocene may be modifying seed dispersal patterns, leading to potential changes in forest community composition.

Insectivorous birds disrupt biological control of cereal aphids.

Insect-feeding birds may interfere with trophic interactions in plant-insect food webs, which may be particularly important in agroecosystems. Here, we studied how Eurasian Tree Sparrows (Passer montanus) affect aphids and their predators in cereal fields using bird exclusion experiments. The Tree Sparrows fed their nestlings with aphid antagonists. Hoverflies and ladybird beetles accounted for 77% of the food for the nestlings during peak aphid density. When birds were excluded, densities of hoverfly larvae, which were the most abundant aphid predator group in the cereals, were 4% higher in wheat and 45% higher in oat, while aphid densities were 24% lower in wheat and 26% lower in oat. The demonstrated disruption of biological control by mesopredators through bird predation may be a common phenomenon in cropping systems characterized by small-sized and abundant pest species. Management of biotic interactions such as biological control needs a broad food-web perspective, even in simplified agroecosystems.

Cascading effects of predation risk determine how marine predators become terrestrial prey on an oceanic island.

Apex predators can suppress the foraging activity of mesopredators, which may then result in cascading benefits for the prey of those mesopredators. We studied the interactions between a top predator, the Barn Owl (Tyto alba), and their primary prey, an island endemic deer mouse (Peromyscus maniculatus elusus), which in turn consumes the eggs of seabirds nesting on Santa Barbara Island in California. Scripps's Murrelets (Synthliboramphus scrippsi), a threatened nocturnal seabird, arrive annually to breed on this island, and whose first egg is particularly vulnerable to predation by mice. We took advantage of naturally occurring extreme variations in the density of mice and owls on the island over 3 years and predicted that (1) mouse foraging would decrease with increasing predation risk from owls and moonlight and (2) these decreases in foraging would reduce predation on murrelet eggs. We measured the giving up densities of mice with experimental foraging stations and found that mice were sensitive to predation risk and foraged less when owls were more abundant and less during the full moon compared to the new moon. We also monitored the fates of 151 murrelet eggs, and found that murrelet egg predation declined as owl abundance increased, and was lower during the full moon compared to the new moon. Moreover, high owl abundance suppressed egg predation even when mice were extremely abundant. We conclude that there is a behaviorally mediated cascade such that owls on the island had a positive indirect effect on murrelet egg survival. Our study adds to the wider recognition of the strength of risk effects to structure food webs, as well as highlighting the complex ways that marine and terrestrial food webs can intersect.

Food web structure shaped by habitat size and climate across a latitudinal gradient.

Habitat size and climate are known to affect the trophic structure and dynamics of communities, but their interactive effects are poorly understood. Organisms from different trophic levels vary in terms of metabolic requirements and heat dissipation. Indeed, larger species such as keystone predators require more stable climatic conditions than their prey. Likewise, habitat size disproportionally affects large-sized predators, which require larger home ranges and are thus restricted to larger habitats. Therefore, food web structure in patchy ecosystems is expected to be shaped by habitat size and climate variations. Here we investigate this prediction using natural aquatic microcosm (bromeliad phytotelmata) food webs composed of litter resources (mainly detritus), detritivores, mesopredators, and top predators (damselflies). We surveyed 240 bromeliads of varying sizes (water retention capacity) across 12 open restingas in SE Brazil spread across a wide range of tropical latitudes (-12.6° to -27.6°, ca. 2,000 km) and climates (Δ mean annual temperature = 5.3°C). We found a strong increase in predator-to-detritivore mass ratio with habitat size, which was representative of a typical inverted trophic pyramid in larger ecosystems. However, this relationship was contingent among the restingas; slopes of linear models were steeper in more stable and favorable climates, leading to inverted trophic pyramids (and top-down control) being more pronounced in environments with more favorable climatic conditions. By contrast, detritivore-resource and mesopredator-detritivore mass ratios were not affected by habitat size or climate variations across latitudes. Our results highlight that the combined effects of habitat size, climate and predator composition are pivotal to understanding the impacts of multiple environmental factors on food web structure and dynamics.

Incorporating anthropogenic effects into trophic ecology: predator-prey interactions in a human-dominated landscape.

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth's terrestrial surface.

Feeding ecology and trophic comparisons of six shark species in a coastal ecosystem off southern Brazil.

The diets of six shark species, Sphyrna lewini, Sphyrna zygaena, Carcharhinus obscurus, Carcharhinus limbatus, Rhizoprionodon lalandii and Galeocerdo cuvier, were investigated in a subtropical coastal ecosystem of southern Brazil. Stomach content data were obtained to assess foraging niche segregation and ontogenetic shifts in the diets of these sharks. Five of the shark species off the Paraná coast were ichthyophagous, with the exception of S. zygaena, which was teutophagous. With the exception of G. cuvier, which had a generalist diet, the other five species displayed specialization in their feeding. Ontogenetic shifts were observed in C. obscurus and S. lewini with large individuals consuming elasmobranchs. Owing to the diet overlap between C. obscurus and S. lewini, C. obscurus and C. limbatus and R. lalandii and C. limbatus, future studies on the spatial and temporal distributions of these species are needed to understand the extent of competitive interactions.

Bats and birds increase crop yield in tropical agroforestry landscapes.

Human welfare is significantly linked to ecosystem services such as the suppression of pest insects by birds and bats. However, effects of biocontrol services on tropical cash crop yield are still largely unknown. For the first time, we manipulated the access of birds and bats in an exclosure experiment (day, night and full exclosures compared to open controls in Indonesian cacao agroforestry) and quantified the arthropod communities, the fruit development and the final yield over a long time period (15 months). We found that bat and bird exclusion increased insect herbivore abundance, despite the concurrent release of mesopredators such as ants and spiders, and negatively affected fruit development, with final crop yield decreasing by 31% across local (shade cover) and landscape (distance to primary forest) gradients. Our results highlight the tremendous economic impact of common insectivorous birds and bats, which need to become an essential part of sustainable landscape management.