Stable isotopes in physiological ecology and food web research

Mention the word 'isotopes' and most people will think of short-lived radioactive isotopes. Yet far more abundant than the radioactive forms and perhaps much more useful for ecological studies are the stable isotopes. Early interest in stable isotope analyses developed in the geological sciences, and their application in environmental biology has developed slowly until recently. Over the past few years, innovative applications of stable isotope analyses to studies of biological processes have been expanding rapidly, and there is every indication that stable isotope approaches will lead to major advances over the next decade in our understanding of physiological processes and fluxes through ecological systems. Studies of plant and animal physiological ecology and food webs will benefit greatly.     

Testing the long-term stability of marine isoscapes in shelf seas using jellyfish tissues

Maps of the spatial distribution of stable isotope ratios across wide geographic areas (isoscapes) are increasingly used to study mechanisms of nutrient flux, movements of animals, and to improve trophic information derived from stable isotope analyses. Isoscapes are usually constructed from reference samples collected from known geographic positions, a time consuming and costly process. In this study, we test the temporal stability of isoscapes of carbon and nitrogen isotopes across the North Sea over a ten-year period. Using jellyfish tissues as reference organisms, we show that hydrodynamic and biogeochemical processes controlling the distribution of carbon and nitrogen isotope values, and thus that the underlying isoscapes, are temporally stable. Remarkably, broad geographic variations in stable carbon and nitrogen isotope distributions across the North Sea are consistent with previously published variations seen in archaeological cod bones from the ninth to seventeenth centuries, despite dramatic changes in land use in the past 1,000 years. Stable isotope-based studies of trophic interactions or movements in animals with lifespans of a year or more in shelf ecosystems can consequently be referenced to previously published isoscape models, and do not require construction of temporally explicit isotope baseline corrections. Scyphomedusan jellyfish are excellent reference organisms for ecological stable isotope analyses in pelagic ecosystems, due to their widespread distributions, well-defined life histories, and fast tissue growth.     

Beyond trophic morphology: stable isotopes reveal ubiquitous versatility in marine turtle trophic ecology

The idea that interspecific variation in trophic morphology among closely related species effectively permits resource partitioning has driven research on ecological radiation since Darwin first described variation in beak morphology among Geospiza. Marine turtles comprise an ecological radiation in which interspecific differences in trophic morphology have similarly been implicated as a pathway to ecopartition the marine realm, in both extant and extinct species. Because marine turtles are charismatic flagship species of conservation concern, their trophic ecology has been studied intensively using stable isotope analyses to gain insights into habitat use and diet, principally to inform conservation management. This legion of studies provides an unparalleled opportunity to examine ecological partitioning across numerous hierarchical levels that heretofore has not been applied to any other ecological radiation. Our contribution aims to provide a quantitative analysis of interspecific variation and a comprehensive review of intraspecific variation in trophic ecology across different hierarchical levels marshalling insights about realised trophic ecology derived from stable isotopes. We reviewed 113 stable isotope studies, mostly involving single species, and conducted a meta‐analysis of data from adults to elucidate differences in trophic ecology among species. Our study reveals a more intricate hierarchy of ecopartitioning by marine turtles than previously recognised based on trophic morphology and dietary analyses. We found strong statistical support for interspecific partitioning, as well as a continuum of intraspecific trophic sub‐specialisation in most species across several hierarchical levels. This ubiquity of trophic specialisation across many hierarchical levels exposes a far more complex view of trophic ecology and resource‐axis exploitation than suggested by species diversity alone. Not only do species segregate along many widely understood axes such as body size, macrohabitat, and trophic morphology but the general pattern revealed by isotopic studies is one of microhabitat segregation and variation in foraging behaviour within species, within populations, and among individuals. These findings are highly relevant to conservation management because they imply ecological non‐exchangeability, which introduces a new dimension beyond that of genetic stocks which drives current conservation planning. Perhaps the most remarkable finding from our data synthesis is that four of six marine turtle species forage across several trophic levels. This pattern is unlike that seen in other large marine predators, which forage at a single trophic level according to stable isotopes. This finding affirms suggestions that marine turtles are robust sentinels of ocean health and likely stabilise marine food webs. This insight has broader significance for studies of marine food webs and trophic ecology of large marine predators. Beyond insights concerning marine turtle ecology and conservation, our findings also have broader implications for the study of ecological radiations. Particularly, the unrecognised complexity of ecopartitioning beyond that predicted by trophic morphology suggests that this dominant approach in adaptive radiation research likely underestimates the degree of resource overlap and that interspecific disparities in trophic morphology may often over‐predict the degree of realised ecopartitioning. Hence, our findings suggest that stable isotopes can profitably be applied to study other ecological radiations and may reveal trophic variation beyond that reflected by trophic morphology.     

Assessing temporal and spatial trends in estuarine nutrient dynamics using a multi-species stable isotope approach

Coastal urbanisation can alter estuarine nutrient dynamics through the input of point-source and diffuse pollutants, and nutrient concentrations can be highly influenced by seasonal and episodic rainfall and river flow. Understanding of both the spatial and temporal variability of nutrient dynamics is therefore critical to managing these estuaries. This can be achieved by periodically analysing the stable isotopes a range of aquatic taxa with variable nutrient turnover rates, mobility and distribution within the estuary. In two subtropical urban estuaries with different land use patterns, we analysed the carbon and nitrogen stable isotopes of phytoplankton, shrimp, prawns and fish at various proximities to pollution sources in dry and wet seasons. The fast nutrient turnover rates and ubiquity of phytoplankton in the estuary resulted in stable isotopes varying over fine-scale spatial scales, particularly in relation to proximity to point-source pollution. The slower nutrient turnover rates and localised habitat use of prawns, resulted in stable isotopes varying over larger spatial (between pollution sources) and temporal (seasonal) scales. The much slower nutrient turnover rates and high mobility throughout the estuary of fish resulted in stable isotopes varying over very large-scale spatial scales (between estuaries). These results illustrate a wide range of spatial and temporal changes to estuarine nutrient dynamics in subtropical urban estuaries in relation to rainfall conditions and nutrient inputs. This research also highlights the application of stable isotopes in assessing estuarine trophodynamics, and provides direction on the types of organisms that should be used to assess different spatial and temporal trends.     

Stable isotopes of carbon and nitrogen in soil ecological studies

The development of stable isotope techniques is one of the main methodological advances in ecology of the last decades of the 20th century. Many biogeochemical processes are accompanied by changes in the ratio between stable isotopes of carbon and nitrogen (¹²C/¹³C and ¹⁴N/¹⁵N), which allows different ecosystem components and different ecosystems to be distinguished by their isotopic composition. Analysis of isotopic composition makes it possible to trace matter and energy flows through biological systems and to evaluate the rate of many ecological processes. The main concepts and methods of stable isotope ecology and patterns of stable isotope fractionation during organic matter decomposition are considered with special emphasis on the fractionation of isotopes in food chains and the use of stable isotope studies of trophic relationships between soil animals in the field.     

Stock structure of Grey Mackerel, Scomberomorus semifasciatus (Pisces: Scombridae) across northern Australia, based on otolith stable isotope chemistry

The stable isotopes of δ¹⁸O and δ¹³C in sagittal otolith carbonates were used to determine the stock structure of Grey Mackerel, Scomberomorus semifasciatus. Otoliths were collected from Grey Mackerel at ten locations representing much of their distributional and fisheries range across northern Australia from 2005 to 2007. Across this broad range (～ 6500 km), fish from four broad locations—Western Australia (S1), Northern Territory and Gulf of Carpentaria (S2, S3, S4, S5, S6, S7), Queensland east coast mid and north sites (S8, S9) and Queensland east coast south site (S10)—had stable isotope values that were significantly different indicating stock separation. Otolith stable isotopes differed more between locations than among years within a location, indicating temporal stability across years. The spatial separation of these populations indicates a complex stock structure across northern Australia. Stocks of S. semifasciatus appear to be associated with large coastal embayments. These results indicate that optimal fisheries management may require a review of the current spatial arrangements, particularly in relation to the evidence of shared stocks in the Gulf of Carpentaria. Furthermore, as the population of S. semifasciatus in Western Australia exhibited high spatial separation from those at all the other locations examined, further research activities should focus on investigating additional locations within Western Australia for an enhanced determination of stock delineation.     

Stable isotope views on ecosystem function: challenging or challenged?

Stable isotopes and their potential for detecting various and complex ecosystem processes are attracting an increasing number of scientists. Progress is challenging, particularly under global change scenarios, but some established views have been challenged. The IX meeting of the Spanish Association of Terrestrial Ecology (AAET, Úbeda, 18–22 October 2009) hosted a symposium on the ecology of stable isotopes where the linear mixing model approach of partitioning sinks and sources of carbon and water fluxes within an ecosystem was challenged, and new applications of stable isotopes for the study of plant interactions were evaluated. Discussion was also centred on the need for networks that monitor ecological processes using stable isotopes and key ideas for fostering future research with isotopes.     

Variation in stable isotope (δO and δC) signatures in the sagittal otolith carbonate of king threadfin, Polydactylus macrochir across northern Australia reveals multifaceted stock structure

Otoliths of king threadfin, Polydactylus macrochir were collected from 2007 to 2009 at nine locations across northern Australia representing most of their distributional range and areas where fisheries are active. Measurement of the stable isotope ratios of δ¹⁸O and δ¹³C in the sagittal otolith carbonate from assemblages of P. macrochir revealed location-specific signatures and indicated that adult fish sampled from representative sites across their range were significantly different. The significant differences in the isotopic signatures of P. macrochir demonstrated that population subdivision is evident and there is unlikely to be substantial movement of fish among these distinct adult assemblages. The stable isotopic signatures for the fish from the different locations were persistent through time, and therefore it could be concluded that they comprise separate stocks for many of the purposes of fisheries management. The spatial separation of these populations indicates a complex stock structure across northern Australia with stocks of P. macrochir associated with large coastal beaches and embayments on a fine spatial scale. These results indicate that in order to achieve optimal fisheries management, the current spatial management arrangements need to be reviewed, particularly the potential for localised depletion of stocks on small spatial scales. This study has provided further evidence that measurement of the stable isotopes ratios in teleost sagittal otolith carbonate can be a valuable tool in the delineation of fishable stocks or fishery management units of adult fish and that widely distributed fish can nonetheless show strong localised population structure.     

Stable isotope ecology in insects: a review

1. The use of stable isotope analysis (SIA) in ecological research has dramatically increased in recent years largely because it allows researchers to investigate ecological questions that have been previously difficult to address. 2. Ecological applications of SIA include estimating fundamental niche space and overlap, evaluating trophic or species level interactions, and investigating food web structure. Increasingly, researchers have been incorporating SIA in studies of animal migration, disease transmission, diet composition, nutrient assimilation, and body condition among others. 3. Studies using SIA to evaluate the ecology of terrestrial insects have lagged behind other taxonomic groups. This poor representation of stable isotope studies in publications likely stems from a lack of familiarity of entomologists with this technique. 4. An improved understanding of SIA, as well as the advantages and disadvantages specifically related to insect research, will benefit the field of entomology. In addition, insect-model systems provide unique opportunities for entomologists to incorporate SIA in their research to advance our knowledge of insect biology and the stable isotope ecology of insects. 5. We provide background information on stable isotopes, explain sources of isotopic variation, describe the processes of how isotopes are differentially routed and incorporated into an individual's tissues, explain the principles that influence isotopic fractionation and discrimination, highlight different methods and advancements in SIA, review innovative stable isotope studies, and provide an overview of common mistakes, considerations, and future directions entomologists can explore.     

Vegetation controls vary across space and spatial scale in a historic grassland‐forest biome boundary

Ecological boundaries are critical landscape regions of transition between adjacent ecological systems. While environmental controls of boundaries may operate in a scale‐dependent manner, multiple‐scale comparisons of vegetation–environment relationships have been characterized for few boundary systems. We used approximately 250 000 point records on the occurrence of woody versus grassland vegetation in conjunction with climatic, topographical, and soils data to evaluate scale effects and spatial heterogeneity in a 650‐km section of the historic prairie–forest biome boundary of Minnesota, USA. We chose this as a model system because of the availability of historical vegetation data, a considerable spatial extent, a sharp ecological transition, and the ability to avoid confounding from more recent anthropogenic land use change. We developed modeling techniques using hierarchical variance partitioning in a spatially‐structured format that allowed us to simultaneously evaluate vegetation–environment relationships across two‐dimensional space (i.e. the prairie‐forest boundary) and across spatial scales (i.e. varying extents). Soils variables displayed the least spatial autocorrelation at shortest lag distances and tended to be the least important predictors of woody vegetation at all spatial extents. Topographical variables displayed greater spatial heterogeneity in regions dominated by forest compared with prairie and were more important at fine‐intermediate spatial scales, highlighting their likely control on fire regimes. An integrated climatic variable (precipitation minus potential evapotranspiration) displayed a trend of increasing spatial variance across the study region and was unambiguously the strongest biome boundary control, although its joint influence with fire was difficult to characterize. Spatially heterogeneous vegetation–environment relationships were observed at all scales, especially at finer scales. Our results suggest that the importance of environmental controls changes smoothly rather than discretely across scales and demonstrate the need to account for spatial non‐stationarity and scale to predict and understand vegetation distribution across ecological boundaries.     

Revisiting “what do tadpoles really eat?” A 10‐year perspective

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