While conservation and fisheries management are often concerned with changes in population abundance and distribution, shifts in population age–size structure are commonly observed in response to human and environmental stressors. Chinook salmon (Oncorhynchus tshawytscha) have experienced widespread declines in mean age and size throughout their North American range. We investigated the consequences of declines in body size for spawner reproductive potential in terms of total egg mass per female. Our case study is the Yukon River where Chinook salmon have supported subsistence, commercial, and recreational fisheries. Using historical observations on individual body size from throughout the Yukon River and the relationship between female size and total egg mass from the Canadian portion, we estimate a decline in average female reproductive potential of 24%–35% since the 1970s. Because spawner abundances and the population sex ratio have not shown clear trends over time, our results suggest a reduced total population reproductive potential. Changes in spawner quality should be considered when developing management reference points, and conservation of population demographic structure may be necessary to sustain productive Chinook salmon systems.

RationaleA liquid chromatography/tandem mass spectrometry (LC/MS/MS) method was validated and utilized to measure and analyze four steroid hormones related to stress and reproduction in individual samples from a novel tissue, Pacific walrus (Odobenus rosmarus divergens, herein walrus) bone. This method determines steroid hormone concentrations in the remote walrus population over millennia from archaeological (>200 bp), historical (200–20 bp), and modern (2014–2016) time periods.MethodsLipids were extracted from walrus bone collected from these periods using methanol before LC/MS/MS analysis. Isotopically labeled internal standards for each target hormone were added to every sample. Analytical and physiological validations were performed. Additionally, a tissue comparison was done among paired walrus bone, serum, and blubber samples. A rapid resolution liquid chromatography system coupled to a QqQ mass spectrometer was used to analyze all samples after derivatization for progesterone, testosterone, cortisol, and estradiol concentrations. Multiple reaction monitoring was used for MS analysis and data were acquired in positive electrospray ionization mode.ResultsProgesterone, testosterone, cortisol, and estradiol were linear along their respective standard calibration curves based on their R2 values (all > 0.99). Accuracy ranged from 93–111% for all hormones. The recovery of extraction, recovery of hormones without matrix effect, was 92–101%. The overall process efficiency of our method for measuring hormones in walrus bone was 93–112%. Progesterone and testosterone concentrations were not affected by reproductive status among adult females and males, respectively. However, estradiol was different among pregnant and non‐pregnant adult females. Overall, steroid hormones reflect a long‐term reservoir in cortical bone. This method was also successfully applied to walrus bone as old as 3585 bp.ConclusionsLC/MS/MS analysis of bone tissue (0.2–0.3 g) provides stress and reproductive data from elusive walruses that were alive thousands of years ago. Based on physiological validations, tissue comparison, and published literature, steroid hormone concentrations measured in walrus cortical bone could represent an accumulated average around a 10–20‐year time span. By investigating how stress and reproductive physiology may have changed over the past ~3000 years based on bone steroid hormone concentrations, this method will help answer how physiologically resilient walruses are to climate change in the Arctic.

Declining sea ice is expected to change the Arctic's physical and biological systems in ways that are difficult to predict. This study used stable isotope compositions (δ13C and δ15N) of archaeological, historic, and modern Pacific walrus (Odobenus rosmarus divergens) bone collagen to investigate the impacts of changing sea ice conditions on walrus diet during the last ~4000 yr. An index of past sea ice conditions was generated using dinocyst-based reconstructions from three locations in the northeastern Chukchi Sea. Archaeological walrus samples were assigned to intervals of high and low sea ice, and δ13C and δ15N were compared across ice states. Mean δ13C and δ15N values were similar for archaeological walruses from intervals of high and low sea ice; however, variability among walruses was greater during low-ice intervals, possibly indicating decreased availability of preferred prey. Overall, sea ice conditions were not a primary driver of changes in walrus diet. The diet of modern walruses was not consistent with archaeological low sea ice intervals. Rather, the low average trophic position of modern walruses (primarily driven by males), with little variability among individuals, suggests that trophic changes to this Arctic ecosystem are still underway or are unprecedented in the last ~4000 yr.

Analysis of stable carbon and nitrogen isotope values (δ 13 C and δ 15 N) of animal tissues can provide important information about diet, physiology, and movements. Interpretation of δ 13 C and δ 15 N values, however, is influenced by factors such as sample lipid content, tissue-specific isotope discrimination, and tissue turnover rates, which are typically species- and tissue-specific. In this study, we generated lipid normalization models for δ 13 C and investigated the effects of chemical lipid extractions on δ 13 C and δ 15 N in Pacific walrus ( Odobenus rosmarus divergens ) muscle, liver, and skin. We also evaluated tissue-specific isotope discrimination in walrus muscle, liver, skin, and bone collagen. Mean δ 13 C lipid-free of skin and bone collagen were similar, as were mean δ 15 N of muscle and liver. All other tissues differed significantly for both isotopes. Differences in δ 13 C lipid-free and δ 15 N among tissues agreed with published estimates of marine mammal tissue-specific isotope discrimination factors, with the exception of skin. The results of this work will allow researchers to gain a clearer understanding of walrus diet and the structure of Arctic food webs, while also making it possible to directly compare the results of contemporary walrus isotope research with those of historic and paleoecological studies.

Age at maturity is an important parameter in many demographic models and, for some species, can be difficult to obtain using traditional methods. Incremental growth structures act as biological archives, recording information throughout an organism’s life and possibly allowing for the reconstruction of life history events. Concentrations of zinc (Zn) in animal tissues are known to be linked to life history, physiology and reproduction and may be retained in incremental growth structures. This study reconstructed lifetime Zn concentrations in teeth (n = 93) of female Pacific walruses (Odobenus rosmarus divergens) collected from 1932–2016. Zn displayed a characteristic pattern of accumulation, with a change point marking the beginning of a lifelong, linear increase in Zn concentrations. We hypothesized that this change point marks the onset of reproductive maturity. The age at which the change point occurred (agecp) was estimated by counting tooth cementum growth layers. These estimates closely matched literature values of timing of first ovulation in female walruses. Total number of ovulations (estimated from ovary corpora counts from paired tooth/ovary specimens) was closely related to reproductive lifespan (total lifespan – agecp; R2 = 0.70). Further, agecp tracked changes in Pacific walrus population size as a proportion of carrying capacity, decreasing when the population was depleted by commercial hunting and peaking when carrying capacity was exceeded. This novel approach will aid walrus management, and is likely applicable to other species, offering a potentially powerful tool for research, management and conservation of wildlife populations.