Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

Trickle-down effects of aboveground trophic cascades on the soil food web

Trophic cascades are increasingly being regarded as important features of aboveground and belowground food webs, but the effects of aboveground cascades on soil food webs, and vice versa, remains essentially unexplored. We conducted an experiment consisting of model synthesised communities containing grassland plant and invertebrate species, in which treatments included soil only, soil+plants, soil+plants+aphids, and soil+plants+aphids+predators; predator treatments consisted of the lacewing Micromus tasmaniae and ladybird beetle Coccinella undecimpunctata added either singly or in combination. Addition of Micromus largely reversed the negative effects of aphids on plant biomass, while both of the predator species caused large changes in the relative abundances of dominant plant species. Predators of aphids also affected several components of the belowground subsystem. Micromus had positive indirect effects on the primary consumer of the soil decomposer food web (microflora), probably through promoting greater input of basal resources to the decomposer subsystem. Predator treatments also influenced densities of the tertiary consumers of the soil food web (top predatory nematodes), most likely through inducing changes in plant community composition and therefore the quality of resource input to the soil. The secondary consumers of the soil food web (microbe‐feeding nematodes) were, however, unresponsive. The fact that some trophic levels of the soil food web but not others responded to aboveground manipulations is explicable in terms of top‐down and bottom‐up forces differentially regulating different belowground trophic levels. Addition of aphids also influenced microbial community structure, promoted soil bacteria at the expense of fungi, and enhanced the diversity of herbivorous nematodes; in all cases these effects were at least partially reversed by addition of Micromus. These results in tandem point to upper level consumers in aboveground food webs as a potential driver of the belowground subsystem, and provide evidence that predator‐induced trophic cascades aboveground can have effects that trickle through soil food webs.     

Trophic redundancy among fishes in an East African nearshore seagrass community inferred from stable‐isotope analysis

Stable‐isotope analysis supplemented with stomach contents data from published sources was used to quantify the trophic niches, trophic niche overlaps and potential trophic redundancy for the most commonly caught fish species from an East African nearshore seagrass community. This assessment is an important first step in quantifying food‐web structure in a region subject to intense fishing activities. Nearshore food webs were driven by at least two isotopically distinct trophic pathways, algal and seagrass, with a greater proportion of the sampled species feeding within the seagrass food web (57%) compared with the algal food web (33%). There was considerable isotopic niche overlap among species (92% of species overlapped with at least one other species). Narrow isotopic niche widths of most (83%) species sampled, low isotopic similarity (only 23% of species exhibited no differences in δ¹³C and δ¹⁵N) and low predicted trophic redundancy among fishes most commonly caught by fishermen (15%), however, suggest that adjustments to resource management concerning harvesting and gear selectivity may be needed for the persistence of artisanal fishing in northern Tanzania. More detailed trophic studies paired with information on spatio‐temporal variation in fish abundance, especially for heavily targeted species, will assist in the development and implementation of management strategies to maintain coastal food‐web integrity.     

Back to the future? Late Holocene marine food web structure in a warm climatic phase as a predictor of trophodynamics in a warmer South‐Western Atlantic Ocean

Stable carbon and nitrogen isotope ratios in the skeletal elements of both ancient and modern marine species from the Beagle Channel were used to compare the structure of Late Holocene and modern food webs, and predict potential changes as a result of a Sea Surface Temperature (SST) increase in the region. Complementary, ancient and modern shells of limpets and mussels were isotopically analysed to explore changes in the isotopic baseline and compare marine food webs through time after an appropriate correction for baseline shifts. Results confirmed a declining pattern of marine primary productivity during the Late Holocene in the Beagle Channel. In general, the isotopic niches overlapped largely in the ancient food web in comparison to the current marine one, with the exception of that of cormorants (Phalacrocorax sp.). Our data suggest that all the species that have undergone intense human exploitation (Arctocephalus australis, Otaria flavescens and Merluccius sp.) significantly increased their trophic levels. The most important finding of this work was the very high isotopic overlap between snoek (Thyrsites atun) and hake (Merluccius sp.) during the Late Holocene. Increasing SST as a result of global warming could favour the recolonization of the southern South‐Western Atlantic Ocean by snoek from the South‐Eastern Pacific Ocean, with a potential impact on the landings of the economically important Argentine and Austral hake. These findings highlight the relevance of using zooarchaeological remains for providing predictions about marine food webs changes in the near future.     

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier‐marine habitats by developing a multi‐trophic level Bayesian three‐isotope mixing model. We utilized large gradients in stable (δ¹³C, δ¹⁵N, δ²H) and radiogenic (Δ¹⁴C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial‐marine habitats. We also compared isotope ratios between glacial‐marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic‐level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier‐nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest‐nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100–1500 years BP ¹⁴C‐age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP ¹⁴C‐age to modern). Thus terrestrial‐derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial‐marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate‐driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.     

Plant diversity has contrasting effects on herbivore and parasitoid abundance in Centaurea jacea flower heads

High biodiversity is known to increase many ecosystem functions, but studies investigating biodiversity effects have more rarely looked at multi‐trophic interactions. We studied a tri‐trophic system composed of Centaurea jacea (brown knapweed), its flower head‐infesting tephritid fruit flies and their hymenopteran parasitoids, in a grassland biodiversity experiment. We aimed to disentangle the importance of direct effects of plant diversity (through changes in apparency and resource availability) from indirect effects (mediated by host plant quality and performance). To do this, we compared insect communities in C. jacea transplants, whose growth was influenced by the surrounding plant communities (and where direct and indirect effects can occur), with potted C. jacea plants, which do not compete with the surrounding plant community (and where only direct effects are possible). Tephritid infestation rate and insect load, mainly of the dominant species Chaetorellia jaceae, decreased with increasing plant species and functional group richness. These effects were not seen in the potted plants and are therefore likely to be mediated by changes in host plant performance and quality. Parasitism rates, mainly of the abundant chalcid wasps Eurytoma compressa and Pteromalus albipennis, increased with plant species or functional group richness in both transplants and potted plants, suggesting that direct effects of plant diversity are most important. The differential effects in transplants and potted plants emphasize the importance of plant‐mediated direct and indirect effects for trophic interactions at the community level. The findings also show how plant–plant interactions critically affect results obtained using transplants. More generally, our results indicate that plant biodiversity affects the abundance of higher trophic levels through a variety of different mechanisms.     

Dietary partitioning promotes the coexistence of planktivorous species on coral reefs

Theories involving niche diversification to explain high levels of tropical diversity propose that species are more likely to co‐occur if they partition at least one dimension of their ecological niche space. Yet, numerous species appear to have widely overlapping niches based upon broad categorizations of resource use or functional traits. In particular, the extent to which food partitioning contributes to species coexistence in hyperdiverse tropical ecosystems remains unresolved. Here, we use a molecular approach to investigate inter‐ and intraspecific dietary partitioning between two species of damselfish (Dascyllus flavicaudus, Chromis viridis) that commonly co‐occur in branching corals. Species‐level identification of their diverse zooplankton prey revealed significant differences in diet composition between species despite their seemingly similar feeding strategies. Dascyllus exhibited a more diverse diet than Chromis, whereas Chromis tended to select larger prey items. A large calanoid copepod, Labidocera sp., found in low density and higher in the water column during the day, explained more than 19% of the variation in dietary composition between Dascyllus and Chromis. Dascyllus did not significantly shift its diet in the presence of Chromis, which suggests intrinsic differences in feeding behaviour. Finally, prey composition significantly shifted during the ontogeny of both fish species. Our findings show that levels of dietary specialization among coral reef associated species have likely been underestimated, and they underscore the importance of characterizing trophic webs in tropical ecosystems at higher levels of taxonomic resolution. They also suggest that niche redundancy may not be as common as previously thought.     

Diverse diet compositions among harpaline ground beetle species revealed by mixing model analyses of stable isotope ratios

1. Species diversities of some insect lineages have been attributed to differentiation of feeding habits among species. Our objective was to determine variation in diet composition among harpaline ground beetle species occurring in a riverside grassland. 2. We examined the diet compositions of 14 species from six genera in the spring and 10 species from two genera in the autumn. We performed measurements of nitrogen and carbon stable isotope ratios in consumers and in their potential food items, and estimated relative contributions of different food items with two mixing models, IsoSource and MixSIR. 3. IsoSource and MixSIR software gave similar results, but IsoSource tended to calculate higher contributions of principal food items and smaller percentile ranges than MixSIR. Among harparine beetle species, there were diverse food utilisation patterns among four food categories (detritivorous invertebrates, herbivorous invertebrates, C3 plants, and C4 plants). Detritivores comprised the main diets of abundant harpaline species in the spring, whereas abundant harpaline species in the autumn were primarily herbivores feeding on C4 plants, or omnivores feeding on herbivorous invertebrates and C3 plants. Seasonal changes in food use were related to seasonal changes in the abundance of each food resource. 4. Mixing model analysis of stable isotope ratios is a convenient and effective method for roughly estimating diets of many species with diverse food habits (such as ground beetles). This method can contribute to determining the trophic relationships of related insects in one ecosystem.     

Soil acidification reduces the effects of short‐term nutrient enrichment on plant and soil biota and their interactions in grasslands

Soil nitrogen (N) and phosphorus (P) contents, and soil acidification have greatly increased in grassland ecosystems due to increased industrial and agricultural activities. As major environmental and economic concerns worldwide, nutrient enrichment and soil acidification can lead to substantial changes in the diversity and structure of plant and soil communities. Although the separate effects of N and P enrichment on soil food webs have been assessed across different ecosystems, the combined effects of N and P enrichment on multiple trophic levels in soil food webs have not been studied in semiarid grasslands experiencing soil acidification. Here we conducted a short‐term N and P enrichment experiment in non‐acidified and acidified soil in a semiarid grassland on the Mongolian Plateau. We found that net primary productivity was not affected by N or P enrichment alone in either non‐acidified or acidified soil, but was increased by combined N and P enrichment in both non‐acidified and acidified soil. Nutrient enrichment decreased the biomass of most microbial groups in non‐acidified soil (the decrease tended to be greatest with combined N and P enrichment) but not in acidified soil, and did not affect most soil nematode variables in non‐acidified or acidified soil. Nutrient enrichment also changed plant and microbial community structure in non‐acidified but not in acidified soil, and had no effect on nematode community structure in non‐acidified or acidified soil. These results indicate that the responses to short‐term nutrient enrichment were weaker for higher trophic groups (nematodes) than for lower trophic groups (microorganisms) and primary producers (plants). The findings increase our understanding of the effects of nutrient enrichment on multiple trophic levels of soil food webs, and highlight that soil acidification, as an anthropogenic stressor, reduced the responses of plants and soil food webs to nutrient enrichment and weakened plant–soil interactions.     

Trophic structure of a coastal fish community determined with diet and stable isotope analyses

A combination of dietary guild analysis and nitrogen (δ¹⁵N) and carbon (δ¹³C) stable‐isotope analysis was used to assess the trophic structure of the fish community in Rhode Island and Block Island Sounds, an area off southern New England identified for offshore wind energy development. In the autumn of 2009, 2010 and 2011, stomach and tissue samples were taken from 20 fish and invertebrate species for analysis of diet composition and δ¹⁵N and δ¹³C signatures. The food chain in Rhode Island and Block Island Sounds comprises approximately four trophic levels within which the fish community is divided into distinct dietary guilds, including planktivores, benthivores, crustacivores and piscivores. Within these guilds, inter‐species isotopic and dietary overlap is high, suggesting that resource partitioning or competitive interactions play a major role in structuring the fish community. Carbon isotopes indicate that most fishes are supported by pelagic phytoplankton, although there is evidence that benthic production also plays a role, particularly for obligate benthivores such as skates Leucoraja spp. This type of analysis is useful for developing an ecosystem‐based approach to management, as it identifies species that act as direct links to basal resources as well as species groups that share trophic roles.     

Explaining geographical variation in the isotope composition of mouse lemurs (Microcebus)

Aim We sought to quantify geographical variation in the stable isotope values of mouse lemurs (Microcebus) and to determine whether this variation reflects trophic differences among populations or baseline isotopic differences among habitats. If the latter pattern is demonstrated, then Microcebus can become a proxy for tracking baseline habitat isotopic variability. Establishing such a baseline is crucial for identifying niche partitioning in modern and ancient communities. Location We studied five species of Microcebus from eight distinct habitats across Madagascar. Methods We compared isotopic variation in C₃ plants and Microcebus fur within and among localities. We predicted that carbon and nitrogen isotope values of Microcebus should: (1) vary as a function of abiotic variables such as rainfall and temperature, and (2) covary with isotopic values in plants. We checked for trophic differences among Microcebus populations by comparing the average difference between mouse lemur and plant isotope values for each locality. We then used multiple regression models to explain spatial isotope variation in mouse lemurs, testing a suite of explanatory abiotic variables. Results We found substantial isotopic variation geographically. Ranges for mean isotope values were similar for both Microcebus and plants across localities (carbon 3.5–4.0‰; nitrogen 10.5–11.0‰). Mean mouse lemur and plant isotope values were lowest in cool, moist localities and highest in hot, dry localities. Rainfall explained 58% of the variation in Microcebus carbon isotope values, and mean plant nitrogen isotope values explained 99.7% of the variation in Microcebus nitrogen isotope values. Average differences between mouse lemur and plant isotope values (carbon 5.0‰; nitrogen 5.9‰) were similar across localities. Main conclusions Isotopic data suggest that trophic differences among Microcebus populations were small. Carbon isotope values in mouse lemurs were negatively correlated with rainfall. Nitrogen isotope values in Microcebus and plants covaried. Such findings suggest that nitrogen isotope values for Microcebus are a particularly good proxy for tracking baseline isotopic differences among habitats. Our results will facilitate future comparative research on modern mouse lemur communities, and ecological interpretations of extinct Holocene communities.     

Overlapping trophic niches among co‐occurring amphipods from a cryptic species complex

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Determinants of parasitoid communities of willow‐galling sawflies: habitat overrides physiology,host plant and space

Studies on the determinants of plant–herbivore and herbivore–parasitoid associations provide important insights into the origin and maintenance of global and local species richness. If parasitoids are specialists on herbivore niches rather than on herbivore taxa, then alternating escape of herbivores into novel niches and delayed resource tracking by parasitoids could fuel diversification at both trophic levels. We used DNA barcoding to identify parasitoids that attack larvae of seven Pontania sawfly species that induce leaf galls on eight willow species growing in subarctic and arctic–alpine habitats in three geographic locations in northern Fennoscandia, and then applied distance‐ and model‐based multivariate analyses and phylogenetic regression methods to evaluate the hierarchical importance of location, phylogeny and different galler niche dimensions on parasitoid host use. We found statistically significant variation in parasitoid communities across geographic locations and willow host species, but the differences were mainly quantitative due to extensive sharing of enemies among gallers within habitat types. By contrast, the divide between habitats defined two qualitatively different network compartments, because many common parasitoids exhibited strong habitat preference. Galler and parasitoid phylogenies did not explain associations, because distantly related arctic–alpine gallers were attacked by a species‐poor enemy community dominated by two parasitoid species that most likely have independently tracked the gallers’ evolutionary shifts into the novel habitat. Our results indicate that barcode‐ and phylogeny‐based analyses of food webs that span forested vs. tundra or grassland environments could improve our understanding of vertical diversification effects in complex plant–herbivore–parasitoid networks.     

Drought decreases incorporation of recent plant photosynthate into soil food webs regardless of their trophic complexity

Theory suggests that more complex food webs promote stability and can buffer the effects of perturbations, such as drought, on soil organisms and ecosystem functions. Here, we tested experimentally how soil food web trophic complexity modulates the response to drought of soil functions related to carbon cycling and the capture and transfer below‐ground of recent photosynthate by plants. We constructed experimental systems comprising soil communities with one, two or three trophic levels (microorganisms, detritivores and predators) and subjected them to drought. We investigated how food web trophic complexity in interaction with drought influenced litter decomposition, soil CO₂ efflux, mycorrhizal colonization, fungal production, microbial communities and soil fauna biomass. Plants were pulse‐labelled after the drought with ¹³C‐CO₂ to quantify the capture of recent photosynthate and its transfer below‐ground. Overall, our results show that drought and soil food web trophic complexity do not interact to affect soil functions and microbial community composition, but act independently, with an overall stronger effect of drought. After drought, the net uptake of ¹³C by plants was reduced and its retention in plant biomass was greater, leading to a strong decrease in carbon transfer below‐ground. Although food web trophic complexity influenced the biomass of Collembola and fungal hyphal length, ¹³C enrichment and the net transfer of carbon from plant shoots to microbes and soil CO₂ efflux were not affected significantly by varying the number of trophic groups. Our results indicate that drought has a strong effect on above‐ground–below‐ground linkages by reducing the flow of recent photosynthate. Our results emphasize the sensitivity of the critical pathway of recent photosynthate transfer from plants to soil organisms to a drought perturbation, and show that these effects may not be mitigated by the trophic complexity of soil communities, at least at the level manipulated in this experiment.     

Mismatch in microbial food webs: predators but not prey perform better in their local biotic and abiotic conditions

Understanding how trophic levels respond to changes in abiotic and biotic conditions is key for predicting how food webs will react to environmental perturbations. Different trophic levels may respond disproportionately to change, with lower levels more likely to react faster, as they typically consist of smaller‐bodied species with higher reproductive rates. This response could cause a mismatch between trophic levels, in which predators and prey will respond differently to changing abiotic or biotic conditions. This mismatch between trophic levels could result in altered top‐down and bottom‐up control and changes in interaction strength. To determine the possibility of a mismatch, we conducted a reciprocal‐transplant experiment involving Sarracenia purpurea food webs consisting of bacterial communities as prey and a subset of six morphologically similar protozoans as predators. We used a factorial design with four temperatures, four bacteria and protozoan biogeographic origins, replicated four times. This design allowed us to determine how predator and prey dynamics were altered by abiotic (temperature) conditions and biotic (predators paired with prey from either their local or non‐local biogeographic origin) conditions. We found that prey reached higher densities in warmer temperature regardless of their temperature of origin. Conversely, predators achieved higher densities in the temperature condition and with the prey from their origin. These results confirm that predators perform better in abiotic and biotic conditions of their origin while their prey do not. This mismatch between trophic levels may be especially significant under climate change, potentially disrupting ecosystem functioning by disproportionately affecting top‐down and bottom‐up control.     

Relationships between Methylobacteria and Glyphosate with Native and Invasive Plant Species: Implications for Restoration

After removing invasive plants, whether by herbicides or other means, typical restoration design focuses on rebuilding native plant communities while disregarding soil microbial communities. However, microbial–plant interactions are known to influence the relative success of native versus invasive plants. Therefore, the abundance and composition of soil microorganisms may affect restoration efforts. We assessed the effect of herbicide treatment on phytosymbiotic pink‐pigmented facultative methylotrophic (PPFM) bacteria and the potential consequences of native and invasive species establishment post‐herbicide treatment in the lab and in a coastal sage scrub (CSS)/grassland restoration site. Lab tests showed that 4% glyphosate reduced PPFM abundance. PPFM addition to seeds increased seedling length of a native plant (Artemisia californica) but not an invasive plant (Hirschfeldia incana). At the restoration site, methanol addition (a PPFM substrate) improved native bunchgrass (Nassella pulchra) germination and size by 35% over controls. In a separate multispecies field experiment, PPFM addition stimulated the germination of N. pulchra, but not that of three invasive species. Neither PPFM nor methanol addition strongly affected the growth of any plant species. Overall, these results are consistent with the hypothesis that PPFMs have a greater benefit to native than invasive species. Together, these experiments suggest that methanol or PPFM addition could be useful in improving CSS/grassland restorations. Future work should test PPFM effects on additional species and determine how these results vary under different environmental conditions.     

Plant host and soil origin influence fungal and bacterial assemblages in the roots of woody plants

Microbial communities in plant roots provide critical links between above‐ and belowground processes in terrestrial ecosystems. Variation in root communities has been attributed to plant host effects and microbial host preferences, as well as to factors pertaining to soil conditions, microbial biogeography and the presence of viable microbial propagules. To address hypotheses regarding the influence of plant host and soil biogeography on root fungal and bacterial communities, we designed a trap‐plant bioassay experiment. Replicate Populus, Quercus and Pinus plants were grown in three soils originating from alternate field sites. Fungal and bacterial community profiles in the root of each replicate were assessed through multiplex 454 amplicon sequencing of four loci (i.e., 16S, SSU, ITS, LSU rDNA). Soil origin had a larger effect on fungal community composition than did host species, but the opposite was true for bacterial communities. Populus hosted the highest diversity of rhizospheric fungi and bacteria. Root communities on Quercus and Pinus were more similar to each other than to Populus. Overall, fungal root symbionts appear to be more constrained by dispersal and biogeography than by host availability.     

Generalist trophic ecology in a changing habitat: The case of the four‐striped mouse in a woody‐encroached savannah

Trophic generalists tolerate greater habitat change than specialists; however, few studies explore how generalist trophic ecology is affected. We established how the trophic ecology of an extreme generalist, Rhabdomys pumilio, changed in relation to a directionally changing woody‐encroached savannah in Eastern Cape, South Africa by investigating (a) foraging behaviour, (b) trophic niche and (c) feedback effects. (a) Giving‐up densities showed that R. pumilio preferred foraging in subcanopy microhabitat during the night as a result of lower thermoregulatory costs, but had similar preferences for sub‐ and intercanopy microhabitats during the day. (b) An isotope analysis revealed that the dietary composition and trophic niche occupied by R. pumilio differed among tree canopy cover levels (0%, 30% and 80%), which appeared to be related to changes in C₄ grass material and invertebrate availability. (c) Artificial seed patches suggested that R. pumilio was a potentially important postdispersal seed predator of the woody‐encroaching species, Vachellia karroo. Thus, an increase in tree canopy cover altered the trophic niche of R. pumilio by reducing foraging costs at night and providing alternative food resources in terms of availability and source. These findings demonstrate how an extreme generalist adapted to human‐induced habitat change through changes in its trophic ecology.     

Characterizing the trophic niches of stocked and resident cyprinid fishes: consistency in partitioning over time,space and body sizes

Hatchery‐reared fish are commonly stocked into freshwaters to enhance recreational angling. As these fishes are often of high trophic position and attain relatively large sizes, they potentially interact with functionally similar resident fishes and modify food‐web structure. Hatchery‐reared barbel Barbus barbus are frequently stocked to enhance riverine cyprinid fish communities in Europe; these fish can survive for over 20 years and exceed 8 kg. Here, their trophic consequences for resident fish communities were tested using cohabitation studies, mainly involving chub Squalius cephalus, a similarly large‐bodied, omnivorous and long‐lived species. These studies were completed over three spatial scales: pond mesocosms, two streams and three lowland rivers, and used stable isotope analysis. Experiments in mesocosms over 100 days revealed rapid formation of dietary specializations and discrete trophic niches in juvenile B. barbus and S. cephalus. This niche partitioning between the species was also apparent in the streams over 2 years. In the lowland rivers, where fish were mature individuals within established populations, this pattern was also generally apparent in fishes of much larger body sizes. Thus, the stocking of these hatchery‐reared fish only incurred minor consequences for the trophic ecology of resident fish, with strong patterns of trophic niche partitioning and diet specialization. Application of these results to decision‐making frameworks should enable managers to make objective decisions on whether cyprinid fish should be stocked into lowland rivers according to ecological risk.     

Trophodynamics of Southern Ocean pteropods on the southern Kerguelen Plateau

Pteropods are a group of small marine gastropods that are highly sensitive to multiple stressors associated with climate change. Their trophic ecology is not well studied, with most research having focused primarily on the effects of ocean acidification on their fragile, aragonite shells. Stable isotopes analysis coupled with isotope‐based Bayesian niche metrics is useful for characterizing the trophic structure of biological assemblages. These approaches have not been implemented for pteropod assemblages. We used isotope‐based Bayesian niche metrics to investigate the trophic relationships of three co‐occurring pteropod species, with distinct feeding behaviors, sampled from the Southern Kerguelen Plateau area in the Indian Sector of the Southern Ocean—a biologically and economically important but poorly studied region. Two of these species were gymnosomes (shell‐less pteropods), which are traditionally regarded as specialist predators on other pteropods, and the third species was a thecosome (shelled pteropod), which are typically generalist omnivores. For each species, we aimed to understand (a) variability and overlap among isotopic niches; and (b) whether there was a relationship between body size and trophic position. Observed isotopic niche areas were broadest for gymnosomes, especially Clione limacina antarctica, whose observed isotopic niche area was wider than expected on both δ¹³C and δ¹⁵N value axes. We also found that trophic position significantly increased with increasing body length for Spongiobranchaea australis. We found no indication of a dietary shift toward increased trophic position with increasing body size for Clio pyramidata f. sulcata. Trophic positions ranged from 2.8 to 3.5, revealing an assemblage composed of both primary and secondary consumer behaviors. This study provides a comprehensive comparative analysis on trophodynamics in Southern Ocean pteropod species, and supports previous studies using gut content, fatty acid and stable isotope analyses. Combined, our results illustrate differences in intraspecific trophic behavior that may be attributed to differential feeding strategies at species level.