Harbour porpoises respond to climate change

The effects of climate change on marine ecosystems and in particular on marine top predators are difficult to assess due to, among other things, spatial variability, and lack of clear delineation of marine habitats. The banks of West Greenland are located in a climate sensitive area and are likely to elicit pronounced responses to oceanographic changes in the North Atlantic. The recent increase in sea temperatures on the banks of West Greenland has had cascading effects on sea ice coverage, residency of top predators, and abundance of important prey species like Atlantic cod (Gadus morhua). Here, we report on the response of one of the top predators in West Greenland; the harbour porpoise (Phocoena phocoena). The porpoises depend on locating high densities of prey species with high nutritive value and they have apparently responded to the general warming on the banks of West Greenland by longer residence times, increased consumption of Atlantic cod resulting in improved body condition in the form of larger fat deposits in blubber, compared to the situation during a cold period in the 1990s. This is one of the few examples of a measurable effect of climate change on a marine mammal population.     

Can climate change jeopardize predator control of invasive herbivore species? A case study in avocado agro-ecosystems in Spain

Climate change is one of the most important factors affecting the phenology, distribution, composition and diversity of organisms. In agricultural systems many pests and natural enemies are arthropods. As poikilotherm organisms, their body temperature is highly dependent on environmental conditions. Because higher trophic levels typically have lower tolerance to high temperatures than lower trophic levels, trends towards increasing local or regional temperatures may affect the strength of predator/prey interactions and disrupt pest control. Furthermore, increasing temperatures may create climate corridors that could facilitate the invasion and establishment of invasive species originating from warmer areas. In this study we examined the effect of environmental conditions on the dynamics of an agro-ecosystem community located in southern Spain, using field data on predator/prey dynamics and climate gathered during four consecutive years. The study system was composed of an ever-green tree species (avocado), an exotic tetranychid mite, and two native species of phytoseiid mites found in association with this new pest. We also present a climatological analysis of the temperature trend in the area of study during the last 28 years, as evidence of temperature warming occurring in the area. We found that the range of temperatures with positive per capita growth rates was much wider in prey than in predators, and that relative humidity contributed to explain the growth rate variation in predators, but not in prey. Predator and prey differences in thermal performance curves could explain why natural enemies did not respond numerically to the pest when environmental conditions were harsh.     

Climate change impacts on wildlife in a High Arctic archipelago – Svalbard,Norway

The Arctic is warming more rapidly than other region on the planet, and the northern Barents Sea, including the Svalbard Archipelago, is experiencing the fastest temperature increases within the circumpolar Arctic, along with the highest rate of sea ice loss. These physical changes are affecting a broad array of resident Arctic organisms as well as some migrants that occupy the region seasonally. Herein, evidence of climate change impacts on terrestrial and marine wildlife in Svalbard is reviewed, with a focus on bird and mammal species. In the terrestrial ecosystem, increased winter air temperatures and concomitant increases in the frequency of ‘rain‐on‐snow’ events are one of the most important facets of climate change with respect to impacts on flora and fauna. Winter rain creates ice that blocks access to food for herbivores and synchronizes the population dynamics of the herbivore–predator guild. In the marine ecosystem, increases in sea temperature and reductions in sea ice are influencing the entire food web. These changes are affecting the foraging and breeding ecology of most marine birds and mammals and are associated with an increase in abundance of several temperate fish, seabird and marine mammal species. Our review indicates that even though a few species are benefiting from a warming climate, most Arctic endemic species in Svalbard are experiencing negative consequences induced by the warming environment. Our review emphasizes the tight relationships between the marine and terrestrial ecosystems in this High Arctic archipelago. Detecting changes in trophic relationships within and between these ecosystems requires long‐term (multidecadal) demographic, population‐ and ecosystem‐based monitoring, the results of which are necessary to set appropriate conservation priorities in relation to climate warming.     

Warming,eutrophication, and predator loss amplify subsidies between aquatic and terrestrial ecosystems

The exchange of organisms and energy among ecosystems has major impacts on food web structure and dynamics, yet little is known about how climate warming combines with other pervasive anthropogenic perturbations to affect such exchanges. We used an outdoor freshwater mesocosm experiment to investigate the interactive effects of warming, eutrophication, and changes in top predators on the flux of biomass between aquatic and terrestrial ecosystems. We demonstrated that predatory fish decoupled aquatic and terrestrial ecosystems by reducing the emergence of aquatic organisms and suppressing the decomposition of terrestrial plant detritus. In contrast, warming and nutrients enhanced cross‐ecosystem exchanges by increasing emergence and decomposition, and these effects were strongest in the absence of predators. Furthermore, we found that warming advanced while predators delayed the phenology of insect emergence. Our results demonstrate that anthropogenic perturbations may extend well beyond ecosystem boundaries by influencing cross‐ecosystem subsidies. We find that these changes are sufficient to substantially impact recipient communities and potentially alter the carbon balance between aquatic and terrestrial ecosystems and the atmosphere.     

Climate change modifies the size structure of assemblages of emerging aquatic insects

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Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Carrion cycling in food webs: comparisons among terrestrial and marine ecosystems

In light of current global changes to ecosystem function (e.g. climate change, trophic downgrading, and invasive species), there has been a recent surge of interest in exploring differences in nutrient cycling among ecosystem types. In particular, a growing awareness has emerged concerning the importance of scavenging in food web dynamics, although no studies have focused specifically on exploring differences in carrion consumption between aquatic and terrestrial ecosystems. In this forum we synthesize the scavenging literature to elucidate differences in scavenging dynamics between terrestrial and marine ecosystems, and identify areas where future research is needed to more clearly understand the role of carrion consumption in maintaining ecosystem function within each of these environments. Although scavenging plays a similar functional role in terrestrial and aquatic food webs, here we suggest that several fundamental differences exist in scavenging dynamics among these ecosystem types due to the unique selection pressures imposed by the physical properties of water and air. In particular, the movement of carcasses in marine ecosystems (e.g. wave action, upwelling, and sinking) diffuses biological activity associated with scavenging and decomposition across large, three‐dimensional spatial scales, creating a unique spatial disconnect between the processes of production, scavenging, and decomposition, which in contrast are tightly linked in terrestrial ecosystems. Moreover, the limited role of bacteria and temporal stability of environmental conditions on the sea floor appears to have facilitated the evolution of a much more diverse community of macrofauna that relies on carrion for a higher portion of its nutrient consumption than is present in terrestrial ecosystems. Our observations are further discussed as they pertain to the potential impacts of climate change and trophic downgrading (i.e. removal of apex consumers from ecosystems) on scavenging dynamics within marine and terrestrial ecosystems.     

Competitive displacement alters top-down effects on carbon dioxide concentrations in a freshwater ecosystem

Climate change and invasive species have the potential to alter species diversity, creating novel species interactions. Interspecific competition and facilitation between predators may either enhance or dampen trophic cascades, ultimately influencing total predator effects on communities and biogeochemical cycling of ecosystems. However, previous studies have only investigated the effects of a single predator species on CO₂ flux of aquatic ecosystems. In this study, we measured and compared the individual and joint effects of predatory damselfly larvae and diving beetles on total prey biomass, leaf litter processing, and dissolved CO₂ concentrations of experimental bromeliad ecosystems. Damselfly larvae created strong trophic cascades that reduced CO₂ concentrations by ~46 % relative to no-predator treatments. Conversely, the effects of diving beetles on prey biomass, leaf litter processing, and dissolved CO₂ were not statistically different to no-predator treatments. Relative to multiplicative null models, the presence of damselfly larvae and diving beetles together resulted in antagonistic relations that eliminated trophic cascades and top-down influences on CO₂ concentrations. Furthermore, we showed that the antagonistic interactions between predators occurred due to a tactile response that culminated in competitive displacement of damselfly larvae. Our results demonstrate that predator identity and predator–predator interactions can influence CO₂ concentrations of an aquatic ecosystem. We suggest that predator effects on CO₂ fluxes may depend on the particular predator species removed or added to the ecosystem and their interactions with other predators.     

Trophic niche segregation allows range‐extending coral reef fishes to co‐exist with temperate species under climate change

Changing climate is forcing many terrestrial and marine species to extend their ranges poleward to stay within the bounds of their thermal tolerances. However, when such species enter higher latitude ecosystems, they engage in novel interactions with local species, such as altered predator–prey dynamics and competition for food. Here, we evaluate the trophic overlap between range‐extending and local fish species along the east coast of temperate Australia, a hotspot for ocean warming and species range extensions. Stable isotope ratios (δ¹⁵N and δ¹³C) of muscle tissue and stomach content analysis were used to quantify overlap of trophic niche space between vagrant tropical and local temperate fish communities along a 730 km (6°) latitudinal gradient. Our study shows that in recipient temperate ecosystems, sympatric tropical and temperate species do not overlap significantly in their diet—even though they forage on broadly similar prey groups—and are therefore unlikely to compete for trophic niche space. The tropical and temperate species we studied, which are commonly found in shallow‐water coastal environments, exhibited moderately broad niche breadths and local‐scale dietary plasticity, indicating trophic generalism. We posit that because these species are generalists, they can co‐exist under current climate change, facilitating the existence of novel community structures.     

Predator diversity effects cascade across an ecosystem boundary

Food webs in different ecosystems are often connected through spatial resource subsidies. As a result, biodiversity effects in one ecosystem may cascade to adjacent ecosystems. I tested the hypothesis that aquatic predator diversity effects cascade to terrestrial food webs by altering a prey subsidy (biomass and trophic structure of emerging aquatic insects) entering terrestrial food webs, in turn altering the distribution of a terrestrial consumer (spider) that feeds on emerging aquatic insects. Fish presence, but not diversity, altered the trophic structure of emerging aquatic insects by strongly reducing the biomass of emerging predators (dragonflies) relative to non‐feeding taxa (chironomid midges). Fish diversity reduced emerging insect biomass through enhanced effects on the most common prey taxa: predatory dragonflies Pantala flavescens and non‐feeding chironomids. Terrestrial spiders (Tetragnathidae) primarily captured emerging chironomids, which were reduced in the high richness (3 spp.) treatment relative to the 1 and 2 species treatments. As a result, terrestrial spider abundance was lower above pools with high fish richness (3 species) than pools with 1 and 2 species. Synergistic predation effects were mostly limited to the high richness treatment, in which fish occupied each level of vertical microhabitat in the water‐column (benthic, middle, surface). This study demonstrates that predator diversity effects are not limited to the habitat of the predator, but can propagate to adjacent ecosystems, and demonstrates the utility of using simple predator functional traits (foraging domain) to more accurately predict the direction of predator diversity effects.     

Predator and detritivore niche width helps to explain biocomplexity of experimental detritus-based food webs in four aquatic and terrestrial ecosystems

In the study of food webs, the existence and explanation of recurring patterns, such as the scale invariance of linkage density, predator–prey ratios and mean chain length, constitute long-standing issues. Our study focused on litter-associated food webs and explored the influence of detritivore and predator niche width (as δ¹³C range) on web topological structure. To compare patterns within and between aquatic and terrestrial ecosystems and take account of intra-habitat variability, we constructed 42 macroinvertebrate patch-scale webs in four different habitats (lake, lagoon, beech forest and cornfield), using an experimental approach with litterbags. The results suggest that although web differences exist between ecosystems, patterns are more similar within than between aquatic and terrestrial web types. In accordance with optimal foraging theory, we found that the niche width of predators and prey increased with the number of predators and prey taxa as a proportion of total taxa in the community. The tendency was more marked in terrestrial ecosystems and can be explained by a lower per capita food level than in aquatic ecosystems, particularly evident for predators. In accordance with these results, the number of links increased with the number of species but with a significantly sharper regression slope for terrestrial ecosystems. As a consequence, linkage density, which was found to be directly correlated to niche width, increased with the total number of species in terrestrial webs, whereas it did not change significantly in aquatic ones, where connectance scaled negatively with the total number of species. In both types of ecosystem, web robustness to rare species removal increased with connectance and the niche width of predators. In conclusion, although limited to litter-associated macroinvertebrate assemblages, this study highlights structural differences and similarities between aquatic and terrestrial detrital webs, providing field evidence of the central role of niche width in determining the structure of detritus-based food webs and posing foraging optimisation constraints as a general mechanistic explanation of food web complexity differences within and between ecosystem types.     

A cross‐system meta‐analysis reveals coupled predation effects on prey biomass and diversity

Predator diversity and abundance are under strong human pressure in all types of ecosystems. Whereas predator potentially control standing biomass and species interactions in food webs, their effects on prey biomass and especially prey biodiversity have not yet been systematically quantified. Here, we test the effects of predation in a cross‐system meta‐analysis of prey diversity and biomass responses to local manipulation of predator presence. We found 291 predator removal experiments from 87 studies assessing both diversity and biomass responses. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey across ecosystems. Predation effects were highly similar between ecosystem types, whereas previous studies had shown that herbivory or decomposition effects differed fundamentally between terrestrial and aquatic systems based on different stoichiometry of plant material. Such stoichiometric differences between systems are unlikely for carnivorous predators, where effect sizes on species richness strongly correlated to effect sizes on biomass. However, the negative predation effect on prey biomass was ameliorated significantly with increasing prey richness and increasing species richness of the manipulated predator assemblage. Moreover, with increasing richness of the predator assemblage present, the overall negative effects of predation on prey richness switched to positive effects. Our meta‐analysis revealed strong general relationships between predator diversity, prey diversity and the interaction strength between trophic levels in terms of biomass. This study indicates that anthropogenic changes in predator abundance and diversity will potentially have strong effects on trophic interactions across ecosystems. Synthesis The past centuries we have experienced a dramatic loss of top–predator abundance and diversity in most types of ecosystems. To understand the direct consequences of predator loss on a global scale, we quantitatively summarized experiments testing predation effects on prey communities in a cross‐system meta‐analysis. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey, and predation effects were highly similar. However, with increasing predator richness, the overall negative effects of predation on prey richness switched to positive ones. Anthropogenic changes in predator communities will potentially have strong effects on prey diversity, biomass, and trophic interactions across ecosystems.     

Effects of experimental warming on biodiversity depend on ecosystem type and local species composition

Climatic warming is a primary driver of change in ecosystems worldwide. Here, we synthesize responses of species richness and evenness from 187 experimental warming studies in a quantitative meta‐analysis. We asked 1) whether effects of warming on diversity were detectable and consistent across terrestrial, freshwater and marine ecosystems, 2) if effects on diversity correlated with intensity, duration, and experimental unit size of temperature change manipulations, and 3) whether these experimental effects on diversity interacted with ecosystem types. Using multilevel mixed linear models and model averaging, we also tested the relative importance of variables that described uncontrolled environmental variation and attributes of experimental units. Overall, experimental warming reduced richness across ecosystems (mean log‐response ratio = –0.091, 95% bootstrapped CI: –0.13, –0.05) representing an 8.9% decline relative to ambient temperature treatments. Richness did not change in response to warming in freshwater systems, but was more strongly negative in terrestrial (–11.8%) and marine (–10.5%) experiments. In contrast, warming impacts on evenness were neutral overall and in aquatic systems, but weakly negative on land (7.6%). Intensity and duration of experimental warming did not explain variation in diversity responses, but negative effects on richness were stronger in smaller experimental units, particularly in marine systems. Model‐averaged parameter estimation confirmed these main effects while accounting for variation in latitude, ambient temperature at the sites of manipulations, venue (field versus lab), community trophic type, and whether experiments were open or closed to colonization. These analyses synthesize extensive experimental evidence showing declines in local richness with increased temperature, particularly in terrestrial and marine communities. However, the more variable effects of warming on evenness were better explained by the random effect of site identity, suggesting that effects on species’ relative abundances were contingent on local species composition. Synthesis A global research priority is to understand the consequences of climate change for biodiversity. A growing number of experimental studies have manipulated climatic drivers, in particular changes in temperature, in local communities. In the first quantitative meta‐analysis of community‐level studies across freshwater, marine and terrestrial experiments, species richness declined consistently with experimental warming. This effect was insensitive to warming intensity, duration, and multiple environmental and procedural covariates. However, evenness responses were weakly negative only in terrestrial systems and more variable across ecosystem types. Linear mixed model analyses revealed that the identity of local sites explained nearly 50% of variance in evenness effect sizes, compared to only 10% for richness. This result provides evidence that local species composition strongly constrains changes in relative species abundances in response to warming.     

The life history of the diving beetle, Lancetes angusticollis (Curtis) (Coleoptera: Dytiscidae), on sub-Antarctic South Georgia

Two populations of the world's most southerly diving beetle (Lancetes angusticollis) were studied on sub-Antarctic South Georgia between November 1995 and April 1996. Parallel observations were made on laboratory cultures of each life stage. All juvenile stages of L. angusticollis are voracious predators, preying mainly on the herbivorous copepod Boeckella poppei. Adult beetles additionally predate benthic ostracods. Laboratory predation rates and field population densities found in this study suggest that L. angusticollis has a more significant impact on its prey than recognised in recent studies of South Georgian lakes. Spring field samples contained only a few individuals of larval instar IV and adults. Young larvae were present from December onwards, with a rapid progression through juvenile instars during the summer months. Fourth instar larvae left the water for pupation between mid-December and at least mid-February. Combining these observations with known developmental threshold temperatures for each life stage implies that L. angusticollis has a complex biennial (minimum) life-cycle, with overwintering possible in three life stages [aquatic larvae, terrestrial pupae (not proven) and aquatic adults]. L. angusticollis may be a suitable indicator species in the context of climate warming studies: a small (1°C) rise in mean environmental temperatures, comparable to that already observed at several sub-Antarctic and maritime Antarctic localities, would allow completion of an annual (univoltine) life-cycle, with concomitant rapid population increase, and serious implications for trophic interactions in these simple lake ecosystems. Received: 24 November 1997 / Accepted: 12 April 1998     

Climate-driven range expansion of a critically endangered top predator in northeast Atlantic waters

Global climate change is driving rapid distribution shifts in marine ecosystems; these are well established for lower trophic levels, but are harder to quantify for migratory top predators. By analysing a 25-year sightings-based dataset, we found evidence for rapid northwards range expansion of the critically endangered Balearic shearwater Puffinus mauretanicus in northeast Atlantic waters. A 0.6 degrees C sea surface temperature increase in the mid-1990s is interpreted as an underlying controlling factor, while simultaneous northward shifts of plankton and prey fish species suggests a strong bottom-up control. Our results have important conservation implications and provide new evidence for climate-driven regime shift in Atlantic ecosystems.     

Predator diversity and density affect levels of predation upon strongly interactive species in temperate rocky reefs

Indirect effects of predators in the classic trophic cascade theory involve the effects of basal species (e.g. primary producers) mediated by predation upon strongly interactive consumers (e.g. grazers). The diversity and density of predators, and the way in which they interact, determine whether and how the effects of different predators on prey combine. Intraguild predation, for instance, was observed to dampen the effects of predators on prey in many ecosystems. In marine systems, species at high trophic levels are particularly susceptible to extinction (at least functionally). The loss of such species, which is mainly attributed to human activities (mostly fishing), is presently decreasing the diversity of marine predators in many areas of the world. Experimental studies that manipulate predator diversity and investigate the effects of this on strongly interactive consumers (i.e. those potentially capable of causing community-wide effects) in marine systems are scant, especially in the rocky sublittoral. I established an experiment that utilised cage enclosures to test whether the diversity and density of fish predators (two sea breams and two wrasses) would affect predation upon juvenile and adult sea urchins, the most important grazers in Mediterranean sublittoral rocky reefs. Changes in species identity (with sea breams producing major effects) and density of predators affected predation upon sea urchins more than changes in species richness per se. Predation upon adult sea urchins decreased in the presence of multiple predators, probably due to interference competition between sea breams and wrasses. This study suggests that factors that influence both fish predator diversity and density in Mediterranean rocky reefs (e.g. fishing and climate change) may have the potential to affect the predators' ability to control sea urchin population density, with possible repercussions for the whole benthic community structure.     

Stepping in Elton's footprints: a general scaling model for body masses and trophic levels across ecosystems

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.     

Histological evidence that diploid hybrids of <Emphasis Type="Italic">Cobitis taenia</Emphasis> and <Emphasis Type="Italic">C. elongatoides</Emphasis> (Teleostei,Cobitidae) develop into fertile females and sterile males

Rising temperatures likely affect the trophic interactions in temperate regions as global warming progresses. An open question is how a temperature rise may affect consumer pressure and plant abundance in shallow aquatic ecosystems, where most consumers are omnivorous. Interestingly, herbivory (plant-eating) is more prevalent toward low latitudes in ectotherms such as fish and aquatic invertebrates, and this may be temperature driven. We used pond snails (Lymnaea stagnalis L.) as a model aquatic ectotherm species and tested their consumption of both animal prey (Gammarus pulex L.) and plant material (Potamogeton lucens L.) at three different temperatures (15, 20, and 25°C). Higher temperatures led to higher consumption rates by the omnivore on both plant food and animal prey when fed separately. When the food was offered simultaneously, the pond snails consistently preferred animal prey over plant material at all tested temperatures. However, the omnivore did consume plant material even though they had enough animal prey available to them. Based on our experiments, we conclude that with increasing temperatures, L. stagnalis will only increase their consumption rates but not change food preference. Further studies are needed to test the generality of our findings across aquatic species to predict the effect of warming on aquatic plant consumption.     

Diversity in thermal affinity among key piscivores buffers impacts of ocean warming on predator–prey interactions

Asymmetries in responses to climate change have the potential to alter important predator–prey interactions, in part by altering the location and size of spatial refugia for prey. We evaluated the effect of ocean warming on interactions between four important piscivores and four of their prey in the U.S. Northeast Shelf by examining species overlap under historical conditions (1968–2014) and with a doubling in CO₂. Because both predator and prey shift their distributions in response to changing ocean conditions, the net impact of warming or cooling on predator–prey interactions was not determined a priori from the range extent of either predator or prey alone. For Atlantic cod, an historically dominant piscivore in the region, we found that both historical and future warming led to a decline in the proportion of prey species’ range it occupied and caused a potential reduction in its ability to exert top‐down control on these prey. In contrast, the potential for overlap of spiny dogfish with prey species was enhanced by warming, expanding their importance as predators in this system. In sum, the decline in the ecological role for cod that began with overfishing in this ecosystem will likely be exacerbated by warming, but this loss may be counteracted by the rise in dominance of other piscivores with contrasting thermal preferences. Functional diversity in thermal affinity within the piscivore guild may therefore buffer against the impact of warming on marine ecosystems, suggesting a novel mechanism by which diversity confers resilience.     

Climate change enhances the negative effects of predation risk on an intermediate consumer

Predators are a major source of stress in natural systems because their prey must balance the benefits of feeding with the risk of being eaten. Although this ‘fear’ of being eaten often drives the organization and dynamics of many natural systems, we know little about how such risk effects will be altered by climate change. Here, we examined the interactive consequences of predator avoidance and projected climate warming in a three‐level rocky intertidal food chain. We found that both predation risk and increased air and sea temperatures suppressed the foraging of prey in the middle trophic level, suggesting that warming may further enhance the top‐down control of predators on communities. Prey growth efficiency, which measures the efficiency of energy transfer between trophic levels, became negative when prey were subjected to predation risk and warming. Thus, the combined effects of these stressors may represent an important tipping point for individual fitness and the efficiency of energy transfer in natural food chains. In contrast, we detected no adverse effects of warming on the top predator and the basal resources. Hence, the consequences of projected warming may be particularly challenging for intermediate consumers residing in food chains where risk dominates predator‐prey interactions.