Variation In Trophic Position Research Articles

Trophic Structure and a Temporal Shift in Trophic Diversity of Mobile Consumers in a Subarctic Estuary

Quantifying trophic relationships within and between species in terms of trophic position, ontogeny and trophic diversity provides information on community-level structure and function. Little attention has been focused on examining the trophic structure and temporal changes of communities in freshwater-marine coupled systems in the subarctic and associated impacts of anthropogenic activity on trophic interactions. Therefore, the objective of this study was to quantify the trophic position of mobile consumers (15 species in total) within the lower Churchill River area (Churchill, Manitoba, Canada), examine trophic position variation in relation to ontogeny, and measure the trophic diversity of a fish assemblage before (1993–1995) and after (2019–2020) the installation of the Churchill River weir in the late 1990s. We used stable isotopes (δ13C and δ15N) to quantify individual and group-level variation in trophic position of thirteen fish species and two seal species and also assessed six community-level metrics of a three fish species assemblage between time periods. Overall, species that mainly foraged on freshwater resources occupied lower trophic positions than species that mainly consumed marine resources. Trophic position increased with fish age only in cisco, fourhorn sculpin, Greenland cod and northern pike. A temporal shift from a trophically diverse to a more trophically redundant fish assemblage occurred between 1993–1995 and 2019–2020. As a result, these predator species now play similar trophic roles. Information on the long-term change in trophic structure of this sub-Arctic estuarine system may help with understanding how anthropogenic activity and climate change have influenced the trophic diversity of a fish assemblage inhabiting this system.

On the evolution of trophic position.

The trophic structure of food webs is primarily determined by the variation in trophic position among species and individuals. Temporal dynamics of food web structure are central to our understanding of energy and nutrient fluxes in changing environments, but little is known about how evolutionary processes shape trophic position variation in natural populations. We propose that trophic position, whose expression depends on both environmental and genetic determinants of the diet variation in individual consumers, is a quantitative trait that can evolve via natural selection. Such evolution can occur either when trophic position is correlated with other heritable morphological and behavioural traits under selection, or when trophic position is a target of selection, which is possible if the fitness effects of prey items are heterogeneously distributed along food chains. Recognising trophic position as an evolving trait, whose expression depends on the food web context, provides an important conceptual link between behavioural foraging theory and food web dynamics, and a useful starting point for the integration of ecological and evolutionary studies of trophic position.

Intra‐ and interspecific variation in trophic ecology of ‘predatory’ ants in the subfamily Ponerinae

1. The characterisation of energy flow through communities is a primary goal of ecology. Furthermore, predator–prey interactions can influence both species abundance and community composition. The ant subfamily Ponerinae includes many predatory species that range from generalist insectivores to highly specialised hunters that target a single prey type. Given their high diversity and ubiquity in tropical ecosystems, measuring intra‐ and interspecific variation in their trophic ecology is essential for understanding the role of ants as predators of insect communities.2. The stable isotopic composition of nitrogen of 22 species from the ant subfamily Ponerinae was measured, relative to plants and other predatory and herbivorous insects at two Atlantic Forest sites in Argentina. The study tested the general assumption that ponerine ants are all predatory, and examined intra‐ and interspecific variation in trophic ecology relative to habitat, body size and cytochrome c oxidase subunit 1 sequences (DNA barcoding).3. Stable isotope analysis revealed that most ponerines occupy high trophic levels (primary and secondary predators), but some species overlapped with known insect herbivores. Species residing at low trophic levels were primarily arboreal and may rely heavily on nectar or other plant‐based resources in their diet. In addition, larger species tend to occupy lower trophic positions than smaller species.4. Although some of the species were divided into two or more genetic clusters by DNA barcoding analysis, these clusters did not correspond to intraspecific variation in trophic position; therefore, colony dietary flexibility most probably explains species that inhabit more than one trophic level.

Trophic Trait Evolution Explains Variation in Nutrient Excretion Stoichiometry among Panamanian Armored Catfishes (Loricariidae)

Variation in nutrient excretion rates and stoichiometric ratios (e.g., nitrogen to phosphorus) by consumers can have substantial effects on aquatic ecosystem function. While phylogenetic signals within an assemblage often explain variation in nutrient recycling rates and stoichiometry, the phylogenetically conserved traits that underlie this phenomenon remain unclear. In particular, variation in nutrient excretion stoichiometry across a phylogeny might be driven by phylogenetic patterns in either diet or body stoichiometry. We examined the relative importance of these traits in explaining variation in nutrient recycling rates and stoichiometry in a diverse family of Neotropical-armored catfishes, Loricariidae, in Panamanian streams. We found significant variation in nutrient mineralization traits among species and subfamilies, but variation in nutrient excretion stoichiometry among species was best explained by trophic position rather than body stoichiometry. The variation in trophic position among Panamanian species was consistent with variation in the trophic niche of their genera across South America, suggesting that phylogenetic patterns underpin the evolution of trophic and nutrient excretion traits among these species. Such geographical variation in nutrient mineralization patterns among closely related species may be common, given that trophic variation in fish lineages occurs widely. These results suggest that information on trophic trait evolution within lineages will advance our understanding of the functional contribution of animals to biogeochemical cycling.

Hatchery‐reared endangered Colorado pikeminnow (Ptychocheilus lucius) undergo a gradual transition to piscivory after introduction to the wild

Abstract Disruption of ecosystems by human activities has caused worldwide extinction threats, which has prompted conservationists to implement captive breeding programmes that aid the recovery of imperilled species. Understanding factors that limit the survival of hatchery‐spawned fishes after stocking is critical to future conservation efforts using captive populations. As the size at which juvenile piscivorous fishes shift to consuming other fish can influence their survival, the transition to piscivory by hatchery‐spawned Ptychocheilus lucius Girard, 1856 (Colorado pikeminnow) was investigated after stocking the San Juan River in New Mexico, Colorado, and Utah, USA. The stable isotope 15N was used to track ontogenetic changes in the trophic position of individuals stocked at age 0, with the expectation that they would become fully piscivorous by age 2. By sampling the isotopic signatures of Colorado pikeminnow across multiple years, the ability of river discharge and densities of fish prey to explain interannual variation in trophic position was also explored. Annual variation in river flow and the densities of fish prey had little predictive power in explaining variation in δ15N of age‐1 or age‐2 Colorado pikeminnow. After assessing the isotopic signatures of potential prey, a Bayesian isotopic mixing model suggested that invertebrates comprised nearly 25% of the diet of of both age‐1 and age‐2 individuals. The relationship between Colorado pikeminnow size and δ15N within stocked cohorts indicated that juveniles slowly transitioned to consuming fish prey as they grew, rather than abruptly switching, as indicated from a limited number of dietary studies using wild fish. Together, these results suggest that Colorado pikeminnow stocked into the San Juan River slowly transition to consuming fish prey. If this pattern leads to poor survival and recruitment, the effectiveness of this management action for conservation could be compromised.

A global meta‐analysis of marine predator nitrogen stable isotopes: Relationships between trophic structure and environmental conditions

AbstractAimWe examined potential environmental drivers of broad‐scale spatial patterns in the trophic structure of marine ecosystems as represented by nitrogen stable isotopes in globally distributed marine predators. Additionally, we assessed the effects of spatial scale on the predictive capabilities of environmental variables.LocationGlobal oceans.Time period2000 to 2015.Major taxa studiedTunas: Thunnus albacares, Thunnus obesus, Thunnus alalunga.MethodsWe undertook a global compilation and meta‐analysis of the bulk nitrogen stable isotope ratios (δ15N values) of three tuna species (n = 4,281). After adjusting for regional variations in baseline δ15N values using a global ocean biogeochemistry model, generalized additive mixed models were employed to infer global‐scale oceanographic controls of trophic structure, using cosmopolitan tuna species as a model.ResultsFor the three tuna species, variation in trophic position estimated using bulk δ15N values was largely explained by geographical location and the corresponding oxygen minimum layer depth. Tuna trophic positions declined in areas with reduced oxygen at depth. Food‐chain length, as captured by maximum trophic position, was longer in areas of the western Pacific Ocean and shorter in the northern Atlantic and eastern Pacific Oceans. Trophic adaptability of the tuna predators, as indicated by intraspecific variability, was highest in the western and central Pacific Ocean and lowest in the northern Atlantic Ocean. Our analysis demonstrated that while tunas share similar functional trophic roles, deeper‐foraging tuna species had higher trophic positions globally. The predictive capacity of environmental variables decreased at finer (regional) spatial scales.Main conclusionsOur work suggests that habitat compression resulting from the predicted global expansion of oxygen minimum zones with ocean warming will impact the trophic structure of marine food webs and the corresponding foraging habits of marine predators. Spatial scale analyses highlighted the importance of representing differences in regional ecological dynamics in global‐scale trophic and ecosystem models.

Resource composition mediates the effects of intraspecific variability in nutrient recycling on ecosystem processes

Despite the growing evidence for individual variation in trophic niche within populations, its potential indirect effects on ecosystem processes remains poorly understood. In particular, few studies have investigated how intraspecific trophic variability can modulate the effects of consumers on ecosystems through potential changes in nutrient excretion rates. Here, we first quantified the level of intraspecific trophic variability in 11 wild populations of the omnivorous fish Lepomis gibbosus. Outputs from stomach content and stable isotope analyses revealed that the degree of trophic specialization and trophic positions were highly variable between and within these wild populations. There was intrapopulation variation in trophic position of more than one trophic level, suggesting that individuals consumed a range of plant and animal resources. We then experimentally manipulated intraspecific trophic variability to assess how it can modulate consumer‐mediated nutrient effects on relevant processes of ecosystem functioning. Specifically, three food sources varying in nutrient quality (e.g. plant material, macro‐invertebrate and fish meat) were used individually or in combination to simulate seven diet treatments. Results indicated that intraspecific variability in growth and nitrogen excretion rates were more related to the composition of the diet rather than the degree of specialization, and increased with the trophic position of the diet consumed. We subsequently used microcosms and showed that critical ecosystem functions, such as primary production and community respiration, were affected by the variability in excretory products, and this effect was biomass‐dependent. These results highlight the importance of considering variation within species to better assess the effects of individuals on ecosystems and, more specifically, the effects of consumer‐mediated nutrient recycling because the body size and the trophic ecology of individuals are affected by a large spectrum of natural and human‐induced environmental changes.

Geographic and temporal variation in the trophic ecology of a small-bodied shark: evidence of resilience to environmental change

Shark dietary patterns can determine how they will respond to changes in prey availability and biodiversity. Geographic variation in diet can also indicate if species have unique structuring roles or feeding strategies in different environments. Unfortunately, little is known about the diet of most shark species and how diet varies over time and space. This study used stable isotope analysis to assess the diet of the Australian sharpnose shark (Rhizoprionodon taylori). Plasma and muscle δ13C and δ15N of R. taylori were compared with δ13C and δ15N baselines from multiple embayments to determine the isotopic niche, trophic position, and benthic and pelagic contributions to diet over time and space. Overall, R. taylori had a wide trophic position range and consumed prey from benthic and pelagic sources. However, there was geographic and temporal variation in trophic position and benthic and pelagic contributions. These findings indicate R. taylori is a dietary generalist, but different populations may have unique effects on distinct ecosystems. Geographic variation in diet also suggests R. taylori may be adaptive to changes in prey availability.

Trophic dynamics of several fish species in lakes of a climatically sensitive region, the Hudson Bay Lowlands

Freshwater lakes in the Hudson Bay Lowlands (HBL) area of Ontario are expected to undergo considerable physical, chemical and biological changes related to climatic change; however, the nature of those changes is still very uncertain. As a first step to improve our understanding of fish communities within these subarctic lakes, we aimed to: (a) characterize trophic dynamics of several large-bodied species within three HBL lakes and (b) determine whether trophic dynamics of selected species in the HBL lakes differed from the same species in Southern Ontario lakes. We found that species-specific trophic position and littoral resource reliance varied significantly within and among the HBL lakes of differing biological communities, chemistry and morphometry. Although several significant differences were evident among lakes in the northern and southern regions, we did not find striking consistent differences in trophic dynamics. Based on observations of high variation in trophic position and/or littoral reliance, we can hypothesize that changes in food resources resulting from climatic change would have little impact on most of the large-bodied species.

Variation in the trophic position of common stream fishes and its relationship to the presence of a rare fish, northern leatherside chub (Lepidomeda copei)

AbstractVariation in trophic position can be caused by structural changes in food webs that may affect the presence of, or be affected by the presence of, individual species. We examined variation in the trophic position of fishes across 14 stream sites in the Bear River drainage, WY, USA. This drainage is the focus of ongoing conservation of northern leatherside chub (Lepidomeda copei). Our goals were (i) to describe variation in trophic position of individual species and (ii) to determine whether these measures differed between sites with and without northern leatherside chub. Mean trophic position of individual fish species varied between 0 and 3 trophic positions across sites. For two of these species, trophic position declined at sites without northern leatherside chub. Importantly, habitat surveys from a previous study at 10 of these sites revealed no differences in habitat suitability for northern leatherside chub. This suggests that trophic position revealed systematic differences among sites that were not apparent based on traditional species‐habitat modelling. We outline possible mechanisms behind these patterns and argue that monitoring variation in trophic position can complement traditional, habitat‐based methods for understanding species distributions.

Spatiotemporal Variation of Juvenile Common Carp Foraging Patterns as Inferred from Stable Isotope Analysis

AbstractOntogenetic diet shifts from zooplankton to macroinvertebrates represent an important transition for many fishes during early life stages. However, some fishes have plastic feeding habits that allow them to take advantage of abundant prey resources or display diet specialization instead of undergoing distinct diet shifts. In this study, we evaluated ontogenetic changes in energy sources (based on the carbon isotope signature, δ13C) and trophic position (based on the nitrogen isotope signature, δ15N) for age‐0 Common Carp Cyprinus carpio from eight natural lakes in eastern South Dakota. In general, Common Carp trophic position decreased and energy sources shifted from pelagic to littoral prey resources from July to August. As anticipated, substantial inter‐ and intrapopulation variation existed in Common Carp isotope signatures. Common Carp trophic position increased with increasing fish TL in three lakes but decreased with fish TL in two systems. Intrapopulation variation in trophic position was related to Common Carp relative abundance during July, indicating that individuals may have specialized on different energy resources as a result of intraspecific interactions. The δ13C values decreased with increasing fish size in two populations during July, suggesting a shift toward pelagic resources; δ13C increased with fish size in six populations during August, suggesting a shift toward littoral resources. Our results indicate a high degree of trophic variation in juvenile Common Carp populations, indicative of a foraging generalist. Year‐class strength may increase trophic specialization of individuals but without negative effects on fish growth, suggesting that diet plasticity is an important factor contributing to the success of this invasive species.

Contrasting C and N isotope ratios from sperm whale skin and squid between the Gulf of Mexico and Gulf of California: effect of habitat

To investigate feeding variation between populations of an apex oceanic predator, stable isotope ratios of carbon (δ13C) and nitrogen (δ15N) have been compared in skin of female and immature male sperm whales (Physeter macrocephalus) from the northern Gulf of Mexico (GoM) and the Gulf of California (GC). Whale sexes were determined genetically. The δ13C and δ15N values from squid muscle were used from the GC, and from inshore and offshore sites in the GoM. We documented contrastingly lower δ13C and δ15N from whales and squid of the GoM compared with those from the GC. While this difference may be associated with variation in trophic position, geographic variation in biochemical cycling influenced significantly the contrasting isotope values between gulfs. Within the northern GoM, the highly distinct δ15N values of neritic squid versus mesopelagic squid provide further evidence of habitat specificity in δ15N.

Trophic plasticity in the Atlantic sharpnose shark (Rhizoprionodon terraenovae) from the north central Gulf of Mexico

Quantifying the trophic role of sharks in coastal ecosystems is crucial for the construction of accurate ecosystem models. This is particularly important for wide-ranging species like the Atlantic sharpnose shark (Rhizoprionodon terraenovae), ubiquitous across the northern Gulf of Mexico. We used gut content and stable isotope analyses to determine if differences in abundance of Atlantic sharpnose sharks in the waters around Mobile Bay, Alabama translated into differences in dietary sources or trophic position among sharks sampled east and west relative to the mouth of the bay. Gut content analysis suggested that Atlantic sharpnose sharks eat primarily teleost fishes (%IRI > 90% across size classes), and both stomach content and stable isotope analyses highlighted an ontogenetic shift in diet. Nitrogen stable isotope data from liver and muscle tissues indicated regional shifts in trophic position for Atlantic sharpnose sharks. The mixing model SIAR (stable isotope analysis in R) v.4.0.2 was used to suggest possible contributions from likely prey items for Atlantic sharpnose sharks sampled east and west of Mobile Bay. Portunid crabs and shrimp made higher contributions to the diet of Atlantic sharpnose sharks in the western region, compared to higher and more variable contributions from fish like croaker (Micropogonias undulatus) and hardhead catfish (Arius felis) in the eastern region. Our results suggest trophic plasticity in Atlantic sharpnose sharks, findings that emphasize the importance of examining regional variation in trophic position when constructing coastal foodweb models.

Anthropogenic alterations of lotic food web structure: evidence from the use of nitrogen isotopes

Understanding the processes that regulate food chain length in nature is a classic theme in ecology. Two factors expected to explain variation in food chain length are resource availability (productivity) and environmental stress. We examined the impact of anthropogenic activities, both a source of stress and productivity, on lotic food webs using two simple food web descriptors based on stable nitrogen isotopes (δ15N). We used 1) trophic position of small fish and 2) slopes of δ15N‐size class relationships in the invertebrate community as variables related to food chain length. Trophic positions and δ15N‐size slopes changed significantly across the 23 study sites selected to vary in productivity and impact from industrial activities (trophic position=0.4 to 1.9; slopes=−1 to +2.8). Fish trophic position was not affected by productivity (chlorophyll a) or environmental stress (number of factories on the watershed) and was not modulated by the proportion of predatory invertebrates in the food web leading to fish. This lack of relationship suggests that variation in trophic position is likely influenced by fish feeding behaviour and is therefore not a good integrator of changes in food chain length induced by anthropogenic activities. δ15N‐size slopes significantly increased with productivity and decreased with stress, suggesting that trophic organization by size within invertebrate communities is affected by these factors. These slopes also tended to be greater when the proportion of predators in the community increased in larger size classes. These results suggest that when productivity is high and stress is low, proportions of higher δ15N‐value predators in lotic food webs tend to increase, contributing to the lengthening of food chains. They also suggest that δ15N‐size relationships of invertebrate communities may be used as indicators of food web structure alteration by human activities.

Influence of lateral gradients of hydrologic connectivity on trophic positions of fishes in the Upper Mississippi River

Summary1. Riverscapes consist of the main channel and lateral slackwater habitats along a gradient of hydrological connectivity from maximum connection in main channel habitats to minimum connection in backwaters. Spatiotemporal differences in water currents along this gradient produce dynamic habitat conditions that influence species diversity, population densities and trophic interactions of fishes.2. We examined the importance of lateral connectivity gradients for food web dynamics in the Upper Mississippi River during spring (high flow, moderately low temperatures) and summer (low flow, higher temperatures). We used literature information and gut contents analyses to determine feeding guilds and stable isotope analysis to estimate mean trophic position of local fish assemblages. During June and August 2006, we collected over 1000 tissue samples from four habitats (main channel, secondary channels, tertiary channels and backwaters) distributed within four hydrologic connectivity gradients.3. Mean trophic position differed among feeding guilds and seasons, with highest values in spring. Mean trophic position of fish assemblages, variability in trophic position and food chain length (maximum trophic position) of the two dominant piscivore species (Micropterus salmoides and M. dolomieu) in both seasons were significantly associated with habitat along the lateral connectivity gradient. Food chain length peaked in tertiary channels in both seasons, probably due to higher species diversity of prey at these habitats. We infer that food chain length and trophic position of fish assemblages were lower in backwater habitats in the summer mainly because of the use of alternative food sources in these habitats.4. A greater number of conspecifics exhibited significant among‐habitat variation in trophic position during the summer, indicating that low river stages can constrain fish movements in the Upper Mississippi River.5. Results of this study should provide a better understanding of the fundamental structure of large river ecosystems and an improved basis for river rehabilitation and management through knowledge of the importance of lateral complexity in rivers.

Within- and among-population variation in the trophic position of a pelagic predator, lake trout (Salvelinus namaycush)

Many aquatic consumers have flexible feeding habits, and the diet and trophic position of a species can be expected to vary both within and among populations. In this study, we quantify the importance of both within- and among-population trophic variation for lake trout (Salvelinus namaycush) using stable isotope designations of trophic position from 13 Ontario and Quebec lakes. Lake-to-lake differences explained 78% of the total variation in lake trout trophic position. Analysis using both stable isotopes and published dietary data demonstrated that the trophic position of lake trout failed to increase appreciably as a function of animal body size. This finding was attributed to weak predator size - prey size relationships as well as to there being no relationship between prey fish trophic position and body size. The variance in trophic position of a population reflects the extent to which individuals forage as trophic specialists; however, we did not identify any one factor that was correlated with within-population trophic variation. Our finding that much of the total variation in trophic position represents among-population differences indicates that considering the average trophic position of a population does not mask substantial within-population trophic variation.

Within- and among-population variation in the trophic position of a pelagic predator, lake trout (<i>Salvelinus namaycush</i>)

Many aquatic consumers have flexible feeding habits, and the diet and trophic position of a species can be expected to vary both within and among populations. In this study, we quantify the importance of both within- and among-population trophic variation for lake trout (Salvelinus namaycush) using stable isotope designations of trophic position from 13 Ontario and Quebec lakes. Lake-to-lake differences explained 78% of the total variation in lake trout trophic position. Analysis using both stable isotopes and published dietary data demonstrated that the trophic position of lake trout failed to increase appreciably as a function of animal body size. This finding was attributed to weak pred - ator size - prey size relationships as well as to there being no relationship between prey fish trophic position and body size. The variance in trophic position of a population reflects the extent to which individuals forage as trophic special - ists; however, we did not identify any one factor that was correlated with within-population trophic variation. Our find - ing that much of the total variation in trophic position represents among-population differences indicates that