Evidence for diet partitioning among three coexisting native freshwater fishes in South Africa's Cape Fold Ecoregion

The partitioning of limited resources commonly explains how different species can coexist within the same ecological community. In this 2010 study, the diets of three coexisting freshwater fishes (Cape galaxias Galaxias zebratus, n = 27; Cape kurper Sandelia capensis, n = 60; Breede River redfin Pseudobarbus burchelli, n = 77) were characterised and compared in three headwater streams in South Africa's Cape Fold Ecoregion using gut contents and stable isotope analyses. These data were analysed to ascertain whether the three species exploit distinct trophic niches. Both approaches provided evidence that these species occupy different trophic niches, though with some overlap. However, dietary differences among sites were not consistent and were probably influenced by site-specific factors like resource availability. Pseudobarbus burchelli had a broader niche breadth at Tierkloof Stream than the other two species, but not at Waaihoek or Tierstel Streams. Our results also suggest that P. burchelli consumed a more omnivorous diet than do the other two species, whereas S. capensis occupied a higher trophic position than the other two species and consumed vertebrates. Our findings suggest that these species occupy non-equivalent feeding niches in Cape Fold Ecoregion headwater streams, and that diet partitioning might facilitate their coexistence in these systems.     

Resource partitioning among top predators in a Miocene food web

The exceptional fossil sites of Cerro de los Batallones (Madrid Basin, Spain) contain abundant remains of Late Miocene mammals. From these fossil assemblages, we have inferred diet, resource partitioning and habitat of three sympatric carnivorous mammals based on stable isotopes. The carnivorans include three apex predators: two sabre-toothed cats (Felidae) and a bear dog (Amphicyonidae). Herbivore and carnivore carbon isotope (δ¹³C) values from tooth enamel imply the presence of a woodland ecosystem dominated by C₃ plants. δ¹³C values and mixing-model analyses suggest that the two sabre-toothed cats, one the size of a leopard and the other the size of a tiger, consumed herbivores with similar δ¹³C values from a more wooded portion of the ecosystem. The two sabre-toothed cats probably hunted prey of different body sizes, and the smaller species could have used tree cover to avoid encounters with the larger felid. For the bear dog, δ¹³C values are higher and differ significantly from those of the sabre-toothed cats, suggesting a diet that includes prey from more open woodland. Coexistence of the sabre-toothed cats and the bear dog was likely facilitated by prey capture in different portions of the habitat. This study demonstrates the utility of stable isotope analysis for investigating the behaviour and ecology of members of past carnivoran guilds.     

Trophic ecology of three stingrays (Myliobatoidei: Dasyatidae) off the Brazilian north-eastern coast: Habitat use and resource partitioning

Understanding the ecological role of species with overlapping distributions is central to inform ecosystem management. Here we describe the diet, trophic level and habitat use of three sympatric stingrays, Hypanus guttatus, H. marianae and H. berthalutzae, through combined stomach content and stable isotope (δ¹³C and δ¹⁵N) analyses. Our integrated approach revealed that H. guttatus is a mesopredator that feeds on a diverse diet of benthic and epibenthic marine and estuarine organisms, principally bivalve molluscs, Alpheus shrimp and teleost fishes. Isotopic data supported movement of this species between marine and estuarine environments. H. berthalutzae is also a marine generalist feeder, but feeds primarily on teleost fishes and cephalopods, and consequently occupies a higher trophic level. In contrast, H. marianae is a mesopredator specialized on shrimps and polychaetas occurring only in the marine environment and occupying a low niche breadth. While niche overlap occurred, the three stingrays utilized the same prey resources at different rates and occupied distinct trophic niches, potentially limiting competition for resources and promoting coexistence. These combined data demonstrate that these three mesopredators perform different ecological roles in the ecosystems they occupy, limiting functional redundancy.     

Diet and trophic position of two mackerel species in the archipelago of Madeira,Portugal

The Atlantic chub mackerel Scomber colias and the blue jack mackerel Trachurus picturatus are two abundant species in the Macaronesia region which includes the archipelago of Madeira, Portugal. Both are key species in the trophic web, being important prey for several local top predators, such as seabirds and marine mammals. Nonetheless, little is known about their feeding ecology in oceanic environments. In this study, the authors describe the seasonal variation in the diet of S. colias and T. picturatus in the oceanic region of Madeira throughout a year. Visual inspection of stomach contents revealed that S. colias fed on a broader range of prey groups than T. picturatus, but for both species, zooplankton (particularly calanoid copepods) and fish were the most important food items. The diet of S. colias included a higher proportion of fish, namely Atlantic saury Scomberesox saurus and S. colias, than that of T. picturatus, that included mostly the longspine snipefish Macroramphosus scolopax. T. picturatus consumed a higher proportion of decapods and other copepods. Seasonal variation was found in the diet of both species, with zooplanktonic species being more important in colder months (February to April) for S. colias and during warm months (May to October) for T. picturatus. Their diet in other seasons was dominated by fish. Although they consume similar prey, carbon and nitrogen stable isotope analysis of muscle of S. colias and T. picturatus showed little overlap in their diets, and T. picturatus showed higher δ¹⁵N and a narrower isotopic niche.     

Food partitioning between coexisting Atlantic salmon and brook trout in the Sainte‐Marguerite River ecosystem, Quebec

Food resource partitioning between similar‐sized, sympatric Atlantic salmon Salmo salar and brook trout Salvelinus fontinalis was examined as a possible mechanism enabling their coexistence in a stream (Allaire) of the Sainte‐Marguerite River ecosystem, Quebec, Canada. Fish stomach contents and invertebrate drift were collected concurrently during three diel cycles in August to September 1996. The food and feeding habits of an allopatric brook trout population in a nearby stream (Epinette) were studied for comparison. The diel feeding rhythms of the two coexisting fish species were similar. The composition of their diet, however, showed significant differences. Atlantic salmon predominantly (60–90%) fed on aquatic insects, mainly Ephemeroptera (35–60% of the diet). The brook trout mostly (50–80%) fed upon the allochthonous terrestrial insects (mainly adults of Coleoptera, Hymenoptera and Diptera) which comprised 5–40% of the stream drift. The allopatric brook trout fed opportunistically on the more abundant aquatic insects and terrestrial insects rarely formed 25% of its diet. The allopatric trout fed nearly twice as much as the sympatric brook trout during a day. The results suggest that the differences in feeding by brook trout in the two streams (with and without Atlantic salmon) are the result of inter‐specific interaction with Atlantic salmon and are not related to the differences in food availability between the two streams. Food resource partitioning between Atlantic salmon and brook trout may be viewed as an adaptive response resulting in a greater exploitation of available resources and coexistence.     

Spatial,temporal, age,and sex related variation in the diet of South American sea lions in southern Chile

We analyzed spatial, temporal, and age/sex variation in the diet of the South American sea lions (SASL, Otaria byronia) in southern Chile via δ¹³C and δ¹⁵N isotope analysis, with emphasis on exotic farmed and feral salmonids as a source of prey for this generalist predator. Isotope mixing models show that the primary prey consumed by SASL were demersal and benthopelagic fishes. Individuals sampled in summer and in the outer coast had higher δ¹⁵N values than those from winter or from the inner coast due to consumption of high trophic level demersal fish. We also found evidence of an ontogenetic shift in isotopic niche width as measured by δ¹³C and δ¹⁵N standard ellipse areas, with juveniles showing a restricted isotopic niche width (1.0‰²) in comparison to subadult (2.2‰²) and adult males (1.9‰²). The proportion of salmonids in the diet of the SASL was negligible and lower than reported in previous studies, which may be related to a recent drop in the production of salmon aquaculture in the region. Our study suggests that the SASL are able to adapt to shifts in spatiotemporal variations in the abundance of potential prey, including feral and farmed salmonids.     

Trawl codend design (44 mm diamond PE mesh) and the effect on selectivity for Pagellus erythrinus and Pagellus acarne, two species with different morphometrics

The effects of different body shapes on size selectivity were analysed. Relationships between total length and fork length, height, width and girth were estimated and the selectivity parameters of a 44 mm PE diamond mesh codend determined for common pandora (Pagellus erythrinus) and axillary sea bream (Pagellus acarne). Two data sets were collected during demersal trawl trials carried out in Izmir Bay (Aegean Sea) between May 1996 and February 1997. Regression analyses were used to determine the relationships between total length and other measured dimensions of the species. However, selectivity parameters were estimated from pooled data by using the logistic equation with the maximum likelihood method. Fork length, height, width and girth were found to increase linearly with total length for both species (R² > 0.90, except TL–W for axillary sea bream, 0.76). Ratios of average body thickness (W/H) were 0.45 (±0.002) for common pandora and 0.52 (±0.002) for axillary sea bream. L₅₀ and SR values were found as 12.4 (±0.44) and 2.2 (±0.51) from the eight valid hauls for common pandora and as 13.6 (±0.13) and 1.9 (±0.26) from the three valid hauls for axillary sea bream, respectively. The difference between the size selectivity of the two species in the same family can be explained by the body shape and fish behaviour distinctions.     

Seasonal and habitat variations in diet of the invasive driftwood catfish Trachelyopterus galeatus in a Neotropical river basin,Brazil

The characteristics of successful invaders often include generalist traits that enable adaptation to new environments through plastic responses, including their diet. The use of trophic resources of invasive driftwood catfish Trachelyopterus galeatus of the Upper Paraná River basin, Brazil, were studied with diet analysis and stable isotopic niche metrics based on δ¹⁵N and δ¹³C to test differences between a free-from-dam and damming population, and between wet and dry season. Stomach content analyses revealed significant differences between the populations. The diet of the riverine free-flowing river population was macroinvertebrate dominated, with Coleoptera and Lepidoptera prominent. In the damming population, diet was largely plant based, although Coleoptera was also prominent. Trophic niche breadth comparisons revealed a larger niche in the free-from-dam population versus the damming population that was independent of season. In both sites had dietary differences between the wet and dry season according to stomach contents analyses, although these were less prominent according to stable isotope metrics. Therefore, the diet of this invader is relatively general and plastic, enabling their exploitation of the varying availability of food resources between free-from-dam and damming river sections, and between wet and dry season.     

Diet and Food Web Support of the White Perch, <Emphasis Type="Italic">Morone americana</Emphasis>, in the Hackensack Meadowlands of New Jersey

Synopsis I analyzed stomach contents of white perch, Morone americana, caught throughout the year in the Hackensack River in northern New Jersey. The dominant prey items were amphipods, with shrimp and fish being less important. Stable isotope analysis indicated that the primary producers of importance in the food web leading to white perch were benthic or planktonic microalgae and secondarily Phragmites australis, the dominant macrophyte in the Hackensack Meadowlands. Despite their large size, white perch do not generally appear to feed any higher in the food web than killifish, Fundulus heteroclitus.     

Trophic niche partitioning between two prey and their incidental predators revealed various threats for an endangered species

Documenting trophic niche partitioning and resource use within a community is critical to evaluate underlying mechanisms of coexistence, competition, or predation. Detailed knowledge about foraging is essential as it may influence the vital rates, which, in turn, can affect trophic relationships between species, and population dynamics. The aims of this study were to evaluate resource and trophic niche partitioning in summer/autumn between the endangered Atlantic‐Gaspésie caribou (Rangifer tarandus caribou) population, moose (Alces americanus) and their incidental predators, the black bear (Ursus americanus) and coyote (Canis latrans), and to quantify the extent to which these predators consumed caribou. Bayesian isotopic analysis showed a small overlap in trophic niche for the two sympatric ungulates suggesting a low potential for resource competition. Our results also revealed that caribou occupied a larger isotopic niche area than moose, suggesting a greater diversity of resources used by caribou. Not surprisingly, coyotes consumed mainly deer (Odocoileus virginianus), moose, snowshoe hare (Lepus americanus), and occasionally caribou, while bears consumed mainly vegetation and, to a lesser extent, moose and caribou. As coyotes and bears also feed on plant species, we documented trophic niche overlap between caribou and their predators, as searching for similar resources can force them to use the same habitats and thus increase the encounter rate and, ultimately, mortality risk for caribou. Although the decline in the Gaspésie caribou population is mostly driven by habitat‐mediated predation, we found evidence that the low level of resource competition with moose, added to the shared resources with incidental predators, mainly bears, may contribute to jeopardize the recovery of this endangered caribou population. Highlighting the trophic interaction between species is needed to establish efficient conservation and management strategies to insure the persistence of endangered populations. The comparison of trophic niches of species sharing the same habitat or resources is fundamental to evaluate the mechanisms of coexistence or competition and eventually predict the consequences of ecosystem changes in the community.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Prey composition and ontogenetic shift in coastal populations of longnose gar Lepisosteus osseus

Longnose gar Lepisosteus osseus were collected from May 2012 to July 2013 in the Charleston Harbor and Winyah Bay estuaries (SC, U.S.A.). This study examined trends in stomach fullness, described major prey components and their importance in the diet of L. osseus, compared stomach content‐based trophic level estimates with the stable‐isotope‐based proxy: δ¹⁵N and tested for the occurrence of an ontogenetic diet shift using stomach content analysis and stable C and N isotopes (δ¹³C and δ¹⁵N). Dominant prey families were Clupeidae, Sciaenidae, Penaeidae, Fundulidae and Mugilidae, with the highest consumption rates in autumn. Trophic levels calculated using stomach contents did not correspond to δ¹⁵N (P > 0·05). Stomach contents and stable‐isotope signatures indicate ontogenetic prey composition shifts from low trophic level benthic prey (fundulids) to higher trophic level pelagic prey (clupeids) as the fish grow between 400 and 600 mm in standard length. Due to their biomass, abundance and top predator status, L. osseus play a significant ecological role in the estuarine community composition, although this effect has often been overlooked by past researchers and should be considered in future estuarine community studies.     

Stable-isotope analyses reveal the importance of seagrass beds as feeding areas for juveniles of the speckled worm eel Myrophis punctatus (Teleostei: Ophichthidae) in Florida

The feeding habits and habitats of the speckled worm eel Myrophis punctatus were studied on the mangrove edge of the Indian River Lagoon (IRL, Florida) using gut-content and stable-isotope analyses of carbon (δ(13) C) and nitrogen (δ(15) N). Four taxa were identified through analyses of gut contents, and the index of relative importance suggested that amphipods, microphytobenthos and annelids are the most important food sources in the fish's diet. To assess the feeding habits of the fish after their recruitment to the IRL, these food sources were collected from mangroves and nearby seagrass beds for isotope analyses. Stable isotopes constituted a powerful tool for discriminating fish prey items from mangroves (mean ± s.d.δ(13) C = -20·5 ± 0·6‰) and those from seagrass beds (mean ± s.d.δ(13) C = -16·9 ± 0·6‰), thus providing good evidence of food source origins. The 56 M. punctatus collected [10·0 < total length (L(T) ) < 16·2 cm] had average isotopic signatures of δ(13) C = -16·7 ± 0·2‰ and δ(15) N = 8·2 ± 0·1‰. A significant depletion in (13) C was observed for larger juveniles (15·0 < L(T) < 16·2 cm), suggesting that they found a portion of their food in mangroves. Estimation of the trophic level from stable isotopes (T(Liso)) was similar among different size groups of juvenile fish (T(Liso) = 3·2-3·5); therefore, M. punctatus was considered a secondary consumer, which is consistent with its zoobenthic diet. The concentration-dependent mixing Stable Isotope Analysis in R (SIAR) model revealed the importance of food sources from seagrass beds as carbon sources for all the fish collected, with a significant increase in mangrove prey contributions, such as annelids, in the diet of larger juveniles. This study highlights the importance of seagrass beds as feeding habitats for juveniles of M. punctatus after their recruitment to coastal waters.     

Landscape-mediated variation in diet is associated with egg size and maculation in a generalist forager

Human impacts alter landscapes with consequences for the distribution and availability of high-quality food resources to populations inhabiting those landscapes, which may impact on the reproductive output of individuals in those populations. The sensitivity of wild populations to changes in food resources may vary among stages of the annual cycle. For example, in birds, effects are likely to be greater during costly stages such as egg production. Here we compare assimilated diet (from stable isotope analysis of chick feathers) and egg traits (egg size, shape, eggshell colour and maculation, using pattern-analysis software) in Herring Gulls Larus argentatus, across seven colonies in southwest Scotland and Northern Ireland. The Herring Gull is an opportunistic, generalist forager on both marine and terrestrial resources which frequently exploits anthropogenic food sources such as fishery discards and human refuse. We found that larger eggs were laid in colonies where females consumed either a higher proportion of marine resources or terrestrial resources; smaller eggs were laid in colonies where females had an intermediate diet. In colonies where females consumed more marine items, they also laid eggs with higher maculation (intensity and size of spots) compared with colonies where females mainly consumed terrestrial food. We also found smaller and more pointed eggs, suggestive of resource shortages, in larger colonies. Generalist foragers are often thought to have the capacity to buffer themselves against changes in the food web, provided that enough alternative food is available. However, this study highlights that specializing on the most profitable or available resources has consequences for egg traits even in an opportunistic generalist forager exploiting a large range of habitats. If variation in egg traits is related to reproductive output, then understanding the impact of assimilated diet on reproduction early in the breeding season can provide important insights into how populations will respond to landscapes altered by human impact.     

Seasonal variation in prey preference,diet partitioning and niche breadth in a rich large carnivore guild

Large carnivore community structure is affected by direct and indirect interactions between intra-guild members. Co-existence between different species within a carnivore guild may occur through diet, habitat or temporal partitioning. Since carnivore species are highly dependent on availability and accessibility of prey, diet partitioning is potentially one of the most important mechanisms in allowing carnivores to co-exist. Intra-guild interactions may vary over time as carnivore prey preference and diet overlap can change due to seasonal changes in resource availability. We conducted scat analysis to compare the seasonal changes in prey preference, diet partitioning and niche breadth of four large carnivore species, namely leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea and wild dog Lycaon pictus in central Tuli, Botswana. Large carnivores in central Tuli display a high dietary overlap, with spotted hyena and brown hyena displaying almost complete dietary overlap and the other carnivore species displaying slightly lower but still significant dietary overlap. Dietary niche breadth for both hyena species was high possibly due to their flexible foraging strategies, including scavenging, while leopard and wild dog showed a relatively low niche breadth, suggesting a more specialised diet. High dietary overlap in central Tuli is possibly explained by the high abundance of prey species in the area thereby reducing competition pressure between carnivore species. Our research highlights the need to assess the influence of diet partitioning in structuring large carnivore communities across multiple study sites, by demonstrating that in prey rich environments, the need for diet partitioning by carnivores to avoid competition may be limited.     

The temporal scale of diet and dietary proxies

Diets estimated from different proxies such as stable isotopes, stomach contents, and dental microwear often disagree, leading to nominally well‐supported but greatly differing estimates of diet for both extinct and extant species that complicate our understanding of ecology. We show that these perceived incongruences can be caused by proxies recording diet over vastly different timescales. Field observations reveal a diet averaged over minutes or hours, whereas dental morphology may reflect the diet of a lineage over millions of years of evolution. Failing to explicitly consider the scale of proxies and the potentially large temporal variability in diet can cause erroneous predictions in any downstream analyses such as conservation planning or paleohabitat reconstructions. We propose a cross‐scale framework for conceptualizing diet suitable for both modern ecologists and paleontologists and provide recommendations for any studies involving dietary data. Treating diet in this temporally explicit framework and matching the scale of our questions with the scale of our data will lead to a much richer and clearer understanding of ecological and evolutionary processes.     

Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient

1. Variation among individuals within size or age classes can have profound effects on community dynamics and food‐web structure. We investigated the potential influence of habitat disturbance on intrapopulation niche variation. 2. Amphibians occupy a range of lentic habitats from short‐hydroperiod intermittent ponds to long‐hydroperiod permanent ponds. We quantified ontogenetic diet variation and individual specialisation in wood frog tadpoles (Lithobates sylvaticus) and blue‐spotted salamander larvae (Ambystoma laterale) to investigate the influence of hydroperiod on population niche width across a natural hydroperiod gradient using stable isotope and gut content analyses. In one of the few tests using larval forms, we tested the niche variation hypothesis, which predicts that populations with larger niche widths also have increased individual variation. 3. Our results support the niche variation hypothesis, indicating that more generalised populations exhibit higher within‐individual diet variation. We report gradual changes in the relative importance of diet items, decreased dietary overlap and increased trophic position in L. sylvaticus throughout development. A. laterale became more enriched in δ¹³C and increased in δ¹⁵N throughout its larval period. We did not find a relationship between hydroperiod and niche parameters, indicating that niches are conserved across heterogeneous habitats. In contrast to most documented cases, we estimated low levels of individual specialisation in amphibian larvae. 4. Amphibians are an important link between aquatic and terrestrial ecosystems, whereby diet shifts can influence food‐web structure by altering energy flow pathways and the trophic position of higher consumers, ultimately changing food‐chain length.