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Stable isotope analysis has the potential to expand our understanding of elasmobranch ecology. However, elasmobranchs share unique traits (i.e., retention of urea, lack of adipose tissue, cartilaginous skeletons) that require modified preparation techniques. Alternative tissue collection and preservation methods would allow sampling from ichthyology collections and at remote locations. We compared different collection, preservation, and preparation techniques to identify treatments that yielded robust isotopic data. Blood components collected in tubes coated with lithium heparin (an anti-coagulant) were not isotopically distinct from blood collected in no-additive tubes. Compared to frozen muscle, ethanol-treated muscle had altered δ 13 C values, but similar δ 15 N values. Finally, we removed lipids and urea with petroleum ether and deionized water, respectively. Although untreated and treated muscle had similar amino acid compositions, treated muscle preferentially lost 14 N and had greater C:N ratios. These results indicate that urea affects isotope ratios and that water treatment removes urea without altering muscle protein composition. Although not exhaustive, our study begins to address the need for elasmobranch-specific methods.

There is growing evidence for individuality in dietary preferences and foraging behaviors within populations of various species. This is especially important for apex predators, since they can potentially have wide dietary niches and a large impact on trophic dynamics within ecosystems. We evaluate the diet of an apex predator, the white shark (Carcharodon carcharias), by measuring the stable carbon and nitrogen isotope composition of vertebral growth bands to create lifetime records for 15 individuals from California. Isotopic variations in white shark diets can reflect within-region differences among prey (most importantly related to trophic level), as well as differences in baseline values among the regions in which sharks forage, and both prey and habitat preferences may shift with age. The magnitude of isotopic variation among sharks in our study (>5‰ for both elements) is too great to be explained solely by geographic differences, and so must reflect differences in prey choice that may vary with sex, size, age and location. Ontogenetic patterns in δ(15)N values vary considerably among individuals, and one third of the population fit each of these descriptions: 1) δ(15)N values increased throughout life, 2) δ(15)N values increased to a plateau at ∼5 years of age, and 3) δ(15)N values remained roughly constant values throughout life. Isotopic data for the population span more than one trophic level, and we offer a qualitative evaluation of diet using shark-specific collagen discrimination factors estimated from a 3+ year captive feeding experiment (Δ(13)C(shark-diet) and Δ(15)N(shark-diet) equal 4.2‰ and 2.5‰, respectively). We assess the degree of individuality with a proportional similarity index that distinguishes specialists and generalists. The isotopic variance is partitioned among differences between-individual (48%), within-individuals (40%), and by calendar year of sub-adulthood (12%). Our data reveal substantial ontogenetic and individual dietary variation within a white shark population.

Diet is a crucial trait of an animal’s lifestyle and ecology. The trophic level of an organism indicates its functional position within an ecosystem and holds significance for its ecology and evolution. Here, we demonstrate the use of zinc isotopes (δ66Zn) to geochemically assess the trophic level in diverse extant and extinct sharks, including the Neogene megatooth shark (Otodus megalodon) and the great white shark (Carcharodon carcharias). We reveal that dietary δ66Zn signatures are preserved in fossil shark tooth enameloid over deep geologic time and are robust recorders of each species’ trophic level. We observe significant δ66Zn differences among the Otodus and Carcharodon populations implying dietary shifts throughout the Neogene in both genera. Notably, Early Pliocene sympatric C. carcharias and O. megalodon appear to have occupied a similar mean trophic level, a finding that may hold clues to the extinction of the gigantic Neogene megatooth shark.

The white shark (Carcharodon carcharias) is a wide-ranging apex predator in the northeastern Pacific (NEP). Electronic tagging has demonstrated that white sharks exhibit a regular migratory pattern, occurring at coastal sites during the late summer, autumn and early winter and moving offshore to oceanic habitats during the remainder of the year, although the purpose of these migrations remains unclear. The purpose of this study was to use stable isotope analysis (SIA) to provide insight into the trophic ecology and migratory behaviors of white sharks in the NEP. Between 2006 and 2009, 53 white sharks were biopsied in central California to obtain dermal and muscle tissues, which were analyzed for stable isotope values of carbon (δ(13)C) and nitrogen (δ(15)N). We developed a mixing model that directly incorporates movement data and tissue incorporation (turnover) rates to better estimate the relative importance of different focal areas to white shark diet and elucidate their migratory behavior. Mixing model results for muscle showed a relatively equal dietary contribution from coastal and offshore regions, indicating that white sharks forage in both areas. However, model results indicated that sharks foraged at a higher relative rate in coastal habitats. There was a negative relationship between shark length and muscle δ(13)C and δ(15)N values, which may indicate ontogenetic changes in habitat use related to onset of maturity. The isotopic composition of dermal tissue was consistent with a more rapid incorporation rate than muscle and may represent more recent foraging. Low offshore consumption rates suggest that it is unlikely that foraging is the primary purpose of the offshore migrations. These results demonstrate how SIA can provide insight into the trophic ecology and migratory behavior of marine predators, especially when coupled with electronic tagging data.

The Cookiecutter shark ( Isistius brasiliensis ) is an ectoparasitic, mesopelagic shark that is known for removing plugs of tissue from larger prey, including teleosts, chondrichthyans, cephalopods, and marine mammals. Although this species is widely distributed throughout the world’s tropical and subtropical oceanic waters, like many deep-water species, it remains very poorly understood due to its mesopelagic distribution. We used a suite of biochemical tracers, including stable isotope analysis (SIA), fatty acid analysis (FAA), and environmental DNA (eDNA), to investigate the trophic ecology of this species in the Central Pacific around Hawaii. We found that large epipelagic prey constituted a relatively minor part of the overall diet. Surprisingly, small micronektonic and forage species (meso- and epipelagic) are the most important prey group for Cookiecutter sharks across the studied size range (17–43 cm total length), with larger mesopelagic species or species that exhibit diel vertical migration also being important prey. These results were consistent across all the tracer techniques employed. Our results indicate that Cookiecutter sharks play a unique role in pelagic food webs, feeding on prey ranging from the largest apex predators to small, low trophic level species, in particular those that overlap with the depth distribution of the sharks throughout the diel cycle. We also found evidence of a potential shift in diet and/or habitat with size and season. Environmental DNA metabarcoding revealed new prey items for Cookiecutter sharks while also demonstrating that eDNA can be used to identify recent prey in stomachs frozen for extended periods. Integrating across chemical tracers is a powerful tool for investigating the ecology of elusive and difficult to study species, such as meso- and bathypelagic chondrichthyans, and can increase the amount of information gained from small sample sizes. Better resolving the foraging ecology of these mesopelagic predators is critical for effective conservation and management of these taxa and ecosystems, which are intrinsically vulnerable to overfishing and exploitation.

Trophic ecology and resource use are challenging to discern in migratory marine species, including sharks. However, effective management and conservation strategies depend on understanding these life history details. Here we investigate whether dental enameloid zinc isotope (δ 66 Zn en ) values can be used to infer intrapopulation differences in foraging ecology by comparing δ 66 Zn en with same-tooth collagen carbon and nitrogen (δ 13 C coll , δ 15 N coll ) values from critically endangered sand tiger sharks ( Carcharias taurus ) from Delaware Bay (USA). We document ontogeny and sex-related isotopic differences indicating distinct diet and habitat use at the time of tooth formation. Adult females have the most distinct isotopic niche, likely feeding on higher trophic level prey in a distinct habitat. This multi-proxy approach characterises an animal’s isotopic niche in greater detail than traditional isotope analysis alone and shows that δ 66 Zn en analysis can highlight intrapopulation dietary variability thereby informing conservation management and, due to good δ 66 Zn en fossil tooth preservation, palaeoecological reconstructions. Sex- and ontogeny-related differences in diet and habitat use of endangered sand tiger sharks from Delaware Bay revealed by analysis of shark teeth zinc isotope values.

Stable oxygen isotope (δ18O) compositions from vertebrate tooth enamel are widely used as biogeochemical proxies for paleoclimate. However, the utility of enamel oxygen isotope values for environmental reconstruction varies among species. Herein, we evaluate the use of stable oxygen isotope compositions from pronghorn (Antilocapra americana Gray, 1866) enamel for reconstructing paleoclimate seasonality, an elusive but important parameter for understanding past ecosystems. We serially sampled the lower third molars of recent adult pronghorn from Wyoming for δ18O in phosphate (δ18OPO4) and compared patterns to interpolated and measured yearly variation in environmental waters as well as from sagebrush leaves, lakes, and rivers (δ18Ow). As expected, the oxygen isotope compositions of phosphate from pronghorn enamel are enriched in 18O relative to environmental waters. For a more direct comparison, we converted δ18Ow values into expected δ18OPO4* values (δ18OW‐PO4*). Pronghorn δ18OPO4 values from tooth enamel record nearly the full amplitude of seasonal variation from Wyoming δ18OW‐PO4* values. Furthermore, pronghorn enamel δ18OPO4 values are more similar to modeled δ18OW‐PO4* values from plant leaf waters than meteoric waters, suggesting that they obtain much of their water from evaporated plant waters. Collectively, our findings establish that seasonality in source water is reliably reflected in pronghorn enamel, providing the basis for exploring changes in the amplitude of seasonality of ancient climates. As a preliminary test, we sampled historical pronghorn specimens (1720 ± 100 AD), which show a mean decrease (a shift to lower values) of 1–2‰ in δ18OPO4 compared to the modern specimens. They also exhibit an increase in the δ18O amplitude, representing an increase in seasonality. We suggest that the cooler mean annual and summer temperatures typical of the 18th century, as well as enhanced periods of drought, drove differences among the modern and historical pronghorn, further establishing pronghorn enamel as excellent sources of paleoclimate proxy data. We evaluate the use of stable oxygen isotope compositions (δ18O in phosphate; δ18OPO4) from pronghorn (Antilocapra americana Gray, 1866) enamel for reconstructing paleoclimate seasonality by serially sampling lower third molars of recent adults (died 1970, 1972, 2010) from Wyoming. We found that pronghorn δ18OPO4 values from tooth enamel record nearly the full amplitude of seasonal variation from Wyoming. Collectively, our findings establish that seasonality in source water is reliably reflected in pronghorn enamel, providing the basis for exploring changes in the amplitude of seasonality of ancient climates.

The capacity of terrestrial ecosystems to sequester carbon dioxide (CO2) from the atmosphere is expected to be altered by climate change and CO2 fertilization, but this projection is limited by our understanding of how the soil system interacts with plants. Understanding the soil–vegetation interactions is essential to assess the magnitude and response of terrestrial ecosystems to the changing climate. Here, we used soil profile and satellite data to explore the role that soil properties play in regulating water and carbon use by plants. Data obtained for 19 terrestrial ecosystem sites in a warm temperate and humid climate were used to investigate the relationship between remotely sensed data and soil physical and chemical properties. Classification and regression tree results showed that in situ soil carbon isotope (δ13C), and soil order were significant predictors (r2 = 0.39, mean absolute error (MAE) = 0 of 0.175 gC/KgH2O) of remotely sensed water use efficiency (WUE) based on the Moderate Resolution Imaging Spectroradiometer (MODIS). Soil extractable calcium (Ca), and land cover type were significant predictors of remotely sensed carbon use efficiency (CUE) based on MODIS and Landsat data-(r2 = 0.64–0.78, MAE = 0.04–0.06). We used gross primary productivity (GPP) derived from solar-induced fluorescence (SIF) data, based on the Orbiting Carbon Observatory-2 (OCO-2), to calculate WUE and CUE (referred to as WUESIF and CUESIF, respectively) for our study sites. The regression tree analysis revealed that soil organic matter and soil extractable magnesium (Mg), δ13C, and soil silt content were the important predictors of both WUESIF (r2 = 0.19, MAE = 0.64 gC/KgH2O) and CUESIF (r2 = 0.45, MAE = 0.1), respectively. Our results revealed the importance of soil extractable Ca, soil carbon (S13C is a facet of soil carbon content), and soil organic matter predicting CUE and WUE. Insights gained from this study highlighted the importance of biotic and abiotic factors regulating plant and soil interactions. These types of data are timely and critical for accurate predictions of how terrestrial ecosystems respond to climate change.

Precipitation is a major driver of ecosystem change and the physiochemical characteristics of soil. Under different climate change scenarios, increased drought frequency and changing precipitation are predicted to impact Mediterranean ecosystems, including those in Northern California. Subsoils are large carbon (C) reservoirs; however, most studies that have investigated precipitation effects on soil organic matter (SOM) have primarily focused on near-surface soils. Recent studies have indicated different responses to environmental perturbation in surface (<30 cm) versus deep (>30 cm) soils due to important differences in physiochemical characteristics. Here, we present soil data at depth (∼300 cm) from a 20-year precipitation manipulation experiment. We determined changes in total elemental concentration and stable isotope composition of soil C, N, δ13C, and δ15N for an ambient control versus additional precipitation in the winter and spring months. The addition of winter precipitation resulted in the largest cumulative C stock (0–300 cm); however, there were no statistically significant changes in C stock throughout the depth profile. Nevertheless, there was evidence of the vertical translocation of C to deep soil layers, specifically of plant-derived C, with both winter and spring precipitation additions. The precipitation addition in winter also resulted in the highest subsoil C stock compared to the control (ambient) and spring treatments. Overall, added winter precipitation led to the best conditions for C accumulation, as the added precipitation coincided with lower temperatures and improved growing conditions at our field site. This study highlights the importance of the timing of precipitation events, especially with regard to deep C stocks (>1 m).

Stable isotopes are important ecological tools, because the carbon and nitrogen isotopic composition of consumer tissue reflects the diet. Measurements of isotopes of individual amino acids can disentangle the effects of consumer physiology from spatiotemporal variation in dietary isotopic values. However, this approach requires knowledge of assimilation patterns of dietary amino acids. We reared leopard sharks (Triakis semifasciata) on diets of squid (Loligo opalescens; 1250 days; control sharks) or squid then tilapia (Oreochromis sp.; switched at 565 days; experimental sharks) to evaluate consumer-diet discrimination factors for amino acids in muscle tissue. We found that control sharks exhibited lower nitrogen isotope discrimination factors (Δ¹⁵N) than most previous consumer studies, potentially because of urea recycling. Control sharks also had large carbon isotope discrimination factors (Δ¹³C) for three essential amino acids, suggesting microbial contributions or fractionation upon assimilation. Compared to controls, experimental sharks exhibited higher Δ¹³C values for four amino acids and Δ¹⁵N values for seven amino acids, corresponding with differences between diets in δ¹³C and δ¹⁵N values. This suggests that not all amino acids in experimental sharks had reached steady state, contrary to the conclusion of a bulk isotope study of these sharks. Our results imply that (1) the magnitude of a shift in dietary δ¹³C and δ¹⁵N values temporarily influences the appearance of discrimination factors; (2) slow turnover of amino acid isotopes in elasmobranch muscle precludes inferences about seasonal dietary changes; (3) elasmobranch discrimination factors for amino acids may be affected by urea recycling and microbial contributions of amino acids.